! DENATURING NATURE ! Philosophical and Historical Reflections on the Artificial-Natural Distinction in the Life Sciences ! ! by ! S. Andrew Inkpen ! B.Sc. Saint Mary’s University 2008 ! ! ! ! ! A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF ! Doctor of Philosophy ! in ! The Faculty of Graduate and Postdoctoral Studies ! (Philosophy) ! ! ! ! ! THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver) ! August 2014 ! © S. Andrew Inkpen, 2014 ! ! ABSTRACT ! ! The philosopher Georges Canguilhem observed that the “physician’s thought and activity are incomprehensible without the concepts of the normal and the pathological.” I argue similarly regarding the biologist, only it is “the artificial” and “the natural” that are indispensable. Whether it is their objects of study, the methods used to investigate those objects, or even fellow researchers, biologists have habitually classified aspects of their discipline in a way that reflects the artificial-natural distinction. Why this way of classifying? What purpose does it serve? What principles guide its application? With what repercussions? Tracing the transformation of these concepts through a series of historical episodes, I explore the reasons why biologists use this distinction and how it has influenced the practices and directions of certain biological fields—specifically evolutionary biology and ecology. The argument of this dissertation is that in biology decisions concerning the choice and evaluation of experimental and evidential practices, objects of study, and even assessments of scientific personas betray the artificial-natural distinction. Invocations of this distinction, like the normal-pathological, code normative contentions about proper biological practice. “The natural,” I argue, often functions as an epistemic authority. The methodology I employ in this dissertation is conceptual and historical. The arguments marshalled are supported by conceptual-philosophical analysis, close readings of primary texts, and archival work. In the end I aim to problematize a set of widely invoked, but heterogeneously used, biological concepts. My arguments undermine a commonplace view according to which the collapse of the artificial-natural distinction is a prerequisite for contemporary science. This distinction is not, I argue, an outdated, pernicious relic; it has continued to exert a significant influence on scientific practice, and should not be ignored.

!ii ! PREFACE ! ! This dissertation is original, unpublished, independent work by the author, S. Andrew Inkpen. A shorter version of Chapter 4 has been accepted for publication in the December 2014 issue of Endeavour as “‘The Art Itself is Nature’: Darwin, Domestic Varieties, and the Scientific Revolution.” ! ! ! !

!iii ! TABLE OF CONTENTS

ABSTRACT ...... ii

PREFACE ...... iii

TABLE OF CONTENTS ...... iv

LIST OF TABLES ...... viVI

LIST OF FIGURES ...... viiVII

ACKNOWLEDGEMENTS ...... viiiVIII

DEDICATION ...... xX

CHAPTER 1 ...... 1 The Artificial-Natural Distinction An Introduction

CHAPTER 2 ...... 32 Art, Nature, and the Scientific Revolution Making Modern Experimental Science

CHAPTER 3 ...... 57 Denaturing Nature Disturbing Conditions and Classifications

CHAPTER 4 ...... 84 The Art Itself is Nature Darwin, Domestic Varieties and the Mechanical Philosophy

CHAPTER 5 ...... 112 Selection in the State of Nature Darwin and Wallace on Domestication

CHAPTER 6 ...... 159 Searching for What Nature has Wrought Dobzhansky and the “Natural” Experimental Fruit Fly

!iv CHAPTER 7 ...... 219 Communities, Natural Experiments, and “Soft” Science Jared Diamond’s Community Ecology

CHAPTER 8 ...... 267 Conclusion

REFERENCES ...... 275 APPENDIX 1 ...... 305 A Note on the Artificial-Natural Classifications of Persons

APPENDIX 2 ...... 311 What is a Natural Experiment?

APPENDIX 3 ...... 316 Diamond on Experimental Tradeoffs ! ! ! ! ! !

!v ! LIST OF TABLES ! ! Table 7.1 Abridged table from Diamond 1986a comparing the advantages and disadvantages of different types of experiments in ecology...... 251

Table A3.1 Full table from Diamond 1986a comparing the advantages and disadvantages of different types of experiments in ecology...... 318 !

!vi ! LIST OF FIGURES ! ! Figure 2.1 The Mirror of Nature and the Image of Art...... 43

Figure 2.2 An Air Pump Used by Robert Boyle...... 45

Figure 4.1 An Elaborate Astronomical Clock in Strasbourg (c. 1875)...... 89

Figure 4.2 Clockmaker About to Fix a Clock...... 92

Figure 4.3 A Mechanical Toy...... 94

Figure 4.4 Victorian’s in Domesticated Nature, “Botanising”...... 95

Figure 4.5 The Polish Pigeon or Polish Cock...... 102

Figure 5.1 Wallace’s Variation in the State of Nature...... 128

Figure 5.2 Darwin’s Variation in the State of Domesticity...... 129

Figure 5.3 An Argus Pheasant Sitting on a Branch of a Tree...... 144

Figure 6.1 The Mutant Type: “eyeless”...... 171

Figure 6.2 The First Linear Linkage Map...... 172

Figure 6.3 A Postcard from Dobzhansky to L. C. Dunn...... 204

Figure 7.1 Diamond’s Geometric Principles for Nature Reserve Design...... 227

Figure 7.2 Distributions of Two Macropygia Cuckoo-Dove Species...... 229

Figure 7.3 Experimental Types...... 313

!vii ACKNOWLEDGEMENTS ! ! I owe a tremendous debt to two people in particular. First, my supervisor, John Beatty. Some time ago John took a risk on an eager biology undergraduate with little training in philosophy or history and who was, probably unbeknownst to John, predisposed to intellectual wandering. John has seen me through every step of graduate education and has been a good friend as well. The best parts of this project are a result of his insight. The worst parts are of my own doing. I owe him a debt I could not possibly repay. Without him, this project would never have begun. Second, my partner in crime, Dani Hallet. She has unscrambled more drafts and heeded more complaining than anyone should be asked to bear, and through this has remained her marvelous and clever self. Her mark is indelibly left on the pages that follow. Without her, this project would never have concluded. I have accrued less tremendous, but still significant, debts to a number of other people. My transition to philosophy can be blamed on Lisa Gannett and John MacKinnon at Saint Mary’s University. The former has become a close friend and has provided needed feedback on this project. The other members of my dissertation committee, Alan Richardson and Chris Stephens, were helpful throughout course work and with aspects of this project as well. Other members of the UBC philosophy faculty have been influential on the direction I have taken, in particular, Margaret Schabas and Sylvia Berryman. Finally, I would like to thank those members of my dissertation examination committee not mentioned above, Bob Brain (university examiner) and Garland Allen (external examiner). Thanks to all my friends at UBC, particularly Chris French, Taylor Davis, Tyler DesRoches, Alirio Rosales, and Jiwon Byun. You have made graduate school both stimulating and enjoyable. Also to all members of the UBC philosophy of biology group. Nissa Bell and Rhonda Janzen have an organizational wherewithal I have no hope of comprehending or approaching, but of which I have made great use and am sincerely thankful. At Harvard University, I’d like to thank, in particular, Janet Browne, Sam Schweber Everett Mendelsohn, and the Modern Sciences Reading Group, the Losos Biological Laboratory, and the Incubator Series for reflecting on the material that formed the foundation of these chapters. Janet and Sam provided feedback on aspects of this dissertation dealing with the 19th century and Charles Darwin, and Janet provided me with a much-needed job.

!viii Thanks to support from the University of British Columbia, the Social Sciences Research Council of Canada, the Darwin Correspondence Project, and the American Philosophical Society.

!ix DEDICATION ! ! ! ! ! ! ! ! For Dani and Rook, Steve and Kathy

!x ! ! CHAPTER 1 ! The Artificial-Natural Distinction An Introduction ! ! ! ! ! ! ! it is the duty of the naturalist [...] to separate artifice from Nature; and never to confound the animal with the slave, the beast of burden with the creature of God Georges-Louis Leclerc, Comte de Buffon, 1749-88, Histoire Naturelle! Domestic animals are abnormal, irregular, artificial; they are subject to varieties which never occur and never can occur in a state of nature: their very existence depends altogether on human care Alfred Russel Wallace, 1858, On the Tendency of Varieties! we do not have any basis for seeing in the process of origin of mutations the result of the artificial influence of man [...] this process goes on just as regularly under natural conditions Sergei Chetverikov, 1926, On Certain Aspects of the Evolutionary Process! studies dealing with natural populations did far more to convince the “Darwinians” of the Mendelian nature of selectively important “natural” variation than either the “artificial” (as they called them) Drosophila mutations or mathematical calculations Ernst Mayr, 1973, The Recent Historiography of Genetics1 ! The philosopher of medicine Georges Canguilhem observed that a “physician’s thought and activity are incomprehensible without the concepts of the normal and the pathological.”2 I argue similarly regarding the biologist, only it is “the artificial” and “the

1 All emphases mine. 2 Canguilhem 2008, 121.

!1 natural” that are indispensable. Whether it is their objects of study, the methods they use to investigate those objects, or even fellow researchers, biologists have habitually classified aspects of their discipline in a way that reflects the artificial-natural distinction.3 Why this way of classifying? What purpose does it serve? What principles guide its application? With what repercussions?

Tracing the transformation of these concepts through a series of historical episodes, I will explore the reasons why biologists use the artificial-natural distinction to think through and classify aspects of their science, as well as how these classifications have influenced the practices and directions of certain biological fields—in particular evolutionary biology and ecology. The argument of this dissertation is that in biology decisions concerning the choice and evaluation of objects, evidential and experimental practices, and even assessments of scientific personas betray the artificial-natural distinction. I will argue that invocations of the artificial-natural distinction, like the normal-pathological, code normative contentions about proper biological practice. “The natural,” I will argue, often functions as an epistemic authority. To the extent that we understand science to be knowledge-making through the investigation of phenomena according to specific methods and practices done by individuals, these scientific decisions, and thus the artificial-natural distinction framing them, shape

3 The third category I highlight here, “researchers themselves,” may cause some head-scratching. I have two things in mind, as explained below. First is that biologists often categorize other biologists according to a dichotomy of naturalist versus experimentalist. This was especially common in the early decades of the 20th century and still happens today. Second, and relatedly, whether biologists draw the artificial-natural distinction or deny it, they will be viewed by the other camp as being naive, uninformed, cognitively or methodologically limited etc. Naturalists are more likely to draw this distinction than experimentalists. Chapter 3 and 6 consider these issues in more detail. As an illustration of the extent to which this language commonly but is diversely used, a 2010 JSTOR search of the journals Ecology and Evolution showed that the artificial-natural distinction was used to categorize the following things (this from only the first 50 results): selection, environment, organisms, life, populations, ponds, calls, hybridization, survival, manipulation, species, habitats, settings, ecosystems, variation, community, strains, conditions, soil, forest, enemies, patterns, plants, nests, fertilization.

!2 biological science.4 These arguments are made ostensively, by providing examples which show these points to be true of biologists and their science.

The argument of this dissertation undermines a commonplace view according to which the collapse of the artificial-natural distinction was essential for the rise of modern science, and furthermore, according to which the distinction currently lacks significance and relevance. I argue, instead, that this distinction is not merely an outdated, pernicious relic left-over from antiquated anxieties, but has continued to exert a significant influence on scientific practice, thus meriting philosophical and historical treatment.

The origins of the artificial-natural distinction are ancient, pre-dating the origins of biology. Its antiquity and embeddedness makes the invocation of these categories obvious in many cases, yet nonetheless consequential. Motivating this dissertation is the belief that sometimes the most fascinating aspects of scientific discourse and practice are concealed by their everyday familiarity. As 17th century scholar Edward Tayler observed, “ages, like men, rarely demand that the language they allow to think for them be brought to the surface of consciousness.”5 My aim is to bring these concepts to the surface so as to better understand their significance.

The methodology I employ in this dissertation is an amalgamation of philosophical- conceptual analysis and intellectual history; I analyze manifestations of these concepts as words in historical sites.6 The concepts with which I deal are “artificial” and “natural”; the

4 A note about my use of “shape” here. These are the sorts of decisions that we look to in order to distinguish the science of biology from other disciplines or other systematic knowledge-making activities. In this sense they shape what we mean by “biology.” 5 Tayler 1966, 25. 6 See Hacking 2001, Chapter 3, for a discussion of the methodology draw on.

!3 sites are case studies in the history of life sciences up to the present day. The evidence marshaled is supported by conceptual-philosophical analysis, close readings of primary texts, and archival work.

What follows are reflections on a set of commonly but heterogeneously used concepts. It is perhaps helpful to begin at the end, by summarizing seven general conclusions that will emerge throughout this dissertation: ! 1. Invoking the artificial-natural distinction is common in biology and has been central to recalcitrant debates. Biologists, of the stripe considered here, have a habit of approaching their discipline and its puzzles with these two alternatives in mind. To steal a helpful metaphor from historian William Cronon, it serves as their “conceptual map” for understanding the world.7 2. There is a standard historical narrative, which serves as a touchstone for discussions of the artificial-natural distinction in science, that relates the dissolution of the distinction to the rise of modern science. This narrative is highly problematic and cannot accommodate the life sciences. In the life sciences this distinction has remained central to its modern history. 3. The distinction is commonly invoked in biology because it piggybacks on a more fundamental concern about how human actions relate to nature’s “normal” development. A common sentiment encountered in biology is that investigations involving significant manipulations of nature are unwarranted—human manipulation “denatures nature” and misleads investigators with regard to their object of study. I argue that in many cases when this distinction is drawn, humans are treated as, what philosophers call, “disturbing conditions.” 4. The concepts “Nature” and “the natural” do not refer to a physical place, as one might prima facie expect, but to an ideal state free from human disturbance. 5. Artificial things are often treated as lacking the reality of natural things, and this is because the former are, in some way, human disturbed. “The natural” is often, for this

7 Cronon 1995c.

!4 reason, given a privileged epistemic position in the biological sciences: it is the “ultimate” object of knowledge. 6. As Canguilhem showed for “normal” and “pathological,” underlying the artificial- natural distinction are normative concerns about, for example, what objects and methods should be used to gain valid biological knowledge. The dichotomy influences scientific practice because “artificial” and “natural” are analogous to what moral philosophers call “thick concepts.” They are, as Bernard Williams said, both world-guided and action-guiding.8 It follows that classifications made according to the artificial-natural dichotomy are not innocuous: how we classify things changes how we act towards them. 7. These concepts, as well as their relation to one another, have been in a process of continual reconceptualization. Phenomena, practices, etc., once deemed artificial may no longer be so called, which again changes how biologists act towards them. ! This chapter has two further divisions. The first introduces recurrent themes, and summarizes my methodology and the theoretical resources I draw on and develop. The second provides a layout of the dissertation. !

A GENERAL PRIMER INTRODUCING RECURRENT THEMES AND THEORETICAL RESOURCES

The artificial-natural distinction is discussed in a variety of ways and in diverse literatures. As these have collectively influenced my thoughts in this dissertation, I will highlight those most pertinent and explain how the present study builds on and departs from them. This will help to situate what follows as well as highlight its novelty. !

DEFINITIONAL ATTEMPTS AS IDEAL TYPES

8 See Williams 1985, Chapter 8.

!5 There have been many attempts to provide necessary and sufficient conditions for distinguishing between artificial and natural things. All of these draw attention to human influence. The Oxford English Dictionary shows that going back to the 15th century,

“artificial” has commonly referred to that which is made or constructed by human skill or involving human intervention. The natural, when opposed to the artificial, is that which is independent of human influence or contrivance.9 Immediately problems arise. In what way does human influence stamp an object as artificial? What degree of human influence is required for something to count as artificial?

Aristotle answered these questions in terms of “natures”: natural objects have natures, that is, internal impulses to change in a goal-directed manner; artificial objects, say, beds or clothing, on the other hand, may be said to have goals only insofar as humans have thwarted nature’s goals to meet their own.10 Artifacts do not have natures; there is nothing that has as its nature to develop into a coat. Whether Aristotle consistently adheres to this ontological categorization is a problem for his historical commentators. What is certain is that this straightforward classification of natural and artificial things did not endure. By 1740 the philosopher David Hume warned of the ambiguity of “nature,” it is the “repository of anything and everything.”11 Likewise, the 19th century philosopher John Stuart Mill’s reflections make it clear that he not only concurred with Hume, but considered the artificial- natural relationship to be particularly problematic.12

9 See Sagoff 2001 plus the literature below for further discussions. 10 By “change” Aristotle has in mind change by place, growth, or other alteration; not just, that is, locational change. 11 Schabas 2008, 72. 12 Mill 1874. In 1965 Arthur Lovejoy documented 66 different meanings of nature! See Lovejoy 1965.

!6 Although Aristotle’s demarcation criteria did not endure, the idea that artificial things are made by humans and so lack ontological identity has been a common and important one, particularly in biology. To many 18th and 19th century naturalists, for example, domestic animals were ignored or even treated with scorn because they were created by man and thus not a part of the natural, that is, God’s order. Buffon expresses this sentiment in the epigraph.

In terms of more recent definitional attempts, the polymath Herbert Simon is at least prima facie more successful than Aristotle.13 In The Sciences of the Artificial (1981) he discusses the relation between “natural worlds” and “artificial worlds,” demarcating between the objects of the natural and the design sciences (engineering, architecture, business, etc.).

While many today would question his distinction between epistemic cultures, his definitional criteria provide ideal types (or an ideal space) useful for thinking about the artificial-natural distinction.14 ! 1. Artificial things are synthesized (though not always or usually with full forethought) by man. 2. Artificial things may imitate appearances in natural things while lacking, in one or many respects, the reality of the latter. 3. Artificial things can be characterized in terms of functions, goals, adaptation.15

13 For another entertaining attempt, see Jacques Monod Chance and Necessity (1971), Chapter 1. The Nobel prize-winning biologist Monod introduces his famous discussion of chance and necessity in biology with a failed “imaginary experiment” to distinguish artificial from natural objects. Because these terms cannot be distinguished “objectively” he is compelled to consider the unique characteristics of living systems, in particular, their (apparent) purposiveness (Monod 1971, 13). 14 See Pauly 1987 and Chapter 7 of Daston and Galison 2007. But see Dear 2003 on differences between these epistemic cultures tracing back to differences between “natural philosophy” and “instrumentality” in ancient and early modern thought. 15 For difficulties maintaining the distinction between natural objects and artifacts in terms of their functions, see Sperber 2007.

!7 4. Artificial things are often discussed, particularly while they are being designed, in terms of imperatives as well as descriptives (how they should be as well as how they are).16 ! We will see these demarcation criteria invoked by biologists throughout the discussions that follow. The connection between the first and second should again bring to mind Aristotle and Buffon—things made lack the reality of things found. However, my aim in this dissertation is not to offer, much less evaluate, definitions of these concepts. Nor is it to survey uses, and extract minimal definitional criteria that are collectively and timelessly held amongst biologists. I am concerned with a different enterprise, to understand how and why biologists invoke this distinction as well as how it influences scientific practice.

Instances of dispute or inconsistency often illuminate best what is at stake.17 Aristotle’s and

Simon’s criteria are, in other words, helpful only insofar as they help illuminate our understanding of biologists’ own reasoning. !

THE EPISTEMIC AUTHORITY OF NATURE?

The artificial-natural distinction is often discussed in a different way in environmental history and policy.18 For example, when Jared Diamond, an influential ecologist and conservation biologist, was asked to reflect on the ends of restoration ecology, he drew attention to a widely held goal “to restore communities to their natural state.”19 “The

16 Simon 1981, 8. 17 Controversies “often involve disagreements over the reality of entities or propriety of practices whose existence or value are subsequently taken as unproblematic or settled” (Shapin and Shaffer 1985, 7). See also, Hacking 2001, 189, and Chapter 12; Cronon 1991, xix. 18 A very good collection is Cronon 1995a; see especially Cronon 1995b. 19 Diamond 1987b, 334. For examples of this sort of thinking throughout ecology see the provocative essay “Natural Landscapes, Natural Communities, and Natural Ecosystems” by Shrader-Frechette and McCoy (1995).

!8 problem,” he said, “is with the very idea expressed in the word ‘natural’. What do we mean by this word?”20 The goal is arbitrary and yet potentially dangerous in its consequences. He continued, ! There are simply too many problems and ambiguities about such a goal. Over most of the surface of the earth there are no really ‘natural’, undisturbed communities left. If we arbitrarily define natural to mean ‘as first seen by Europeans’, we face major ambiguities. Even if we did define some nineteenth century condition as the ‘natural’ condition, there are many communities whose species composition we couldn’t restore even with an infinite amount of time and money.21 ! The artificial-natural distinction is of consequence in this case because it dictates what we see as worthy of protection or as the goal of restoration, and can distract us from what is really feasible.22 In other words, wider considerations about what counts as natural or what counts as artificial can influence practical decisions made by restoration ecologists. Is the rainforest more natural than the surrounding farmlands? Are indigenous song birds more natural than introduced starlings?23 Human disturbed environments or organisms may be neither the goal at which to aim nor worthy of protection in their own right, but such decisions ultimately depend on, or are entangled with, how the artificial-natural distinction is drawn.

20 Ibid., 331 21 Ibid., 334 22 It is also of course possible that the distinction is dictated by what we see as worthy of protection—we call something artificial, that is, because we don’t value it. Rather than consider this Euthyphro-ian dilemma a problem, let me just allow for an obvious feedback loop: we protect things because they are natural, and likewise they are natural because we want to protect them. This sort of problem is considered by Daston and Galison (2007) with regards to the virtue of objectivity. 23 See a recent issue of The Chronicle Review: The Chronicle of Higher Education for an interesting article about the “Battle for the Soul of Conservation” related to the first question (Nov. 15th, 2013). For the second, see Davis et al. 2011.

!9 Seen as both an important environmental category, and yet one that is conceptually problematic and liable to mislead, or worse, put to unwanted political use, many environmentalists and environmental historians try to salvage the category of “the natural” by endorsing a distinction made famous by Karl Marx between first nature (“original, prehuman nature”) and second nature (“the artificial nature that people erect atop first nature”).24 These two concepts are subsumed under the larger blanket concept “Nature,” meaning, “all the powers [or laws of nature] existing in either the outer or the inner world and everything which takes place by means of those powers [or laws of nature].”25 I flag this distinction for a particular purpose: when biologists draw the artificial-natural distinction, they are not denying that humans are a part of Nature in the broadest sense—that human actions are supernatural. They are drawing a distinction between first nature and second nature. The

French political philosopher Georges Sorel meant to draw this same distinction but did so in an atemporal way.26 He called this distinction “artificial nature” versus “natural nature.”

When I speak of the distinction, I have his categories in mind.

Considerations like Diamond’s bring to light the normative aspects inherent in the concept of nature. Historians and philosophers in recent decades have drawn attention to the moral authority of nature: the ways in which “nature” and “the natural” have been, and still

24 See Cronon 1991, xix. See also Marx 2005, 61 for the context. 25 Mill 1874 [1969]. Or Darwin’s, “I mean by Nature, only the aggregate action and product of many natural laws, and by laws the sequence of events as ascertained by us” (Darwin 1872 edition of the Origin). 26 This way is preferable because first nature and second nature make it sound as though the artificial-natural distinction has a temporal dimension, which I think it easier done without. After all, first nature is meant to evoke the feeling of the “untouched” New England wilderness, or at least for William Cronon’s influential usage.

!10 are, used as a cultural value or social norm.27 Such is the basis for the famous naturalistic fallacy—the is-ought gap—in moral theory, which holds that it is a fallacy to assume that what is natural is for that reason good and dictates what we should do, how we should act. A cursory look at history or at contemporary political debates suggests that we have a particular knack for committing this fallacy.28

While it is commonly acknowledged by historians and philosophers that “natural” carries normative weight, these accounts rarely observe, for example, that the phrase “in nature” is a common expression in both ecology and evolutionary biology which has descriptive and normative dimensions interlinked specifically in the context of knowledge- making. As do its antonyms (e.g., “artificial,” “laboratory”). I will argue in this dissertation that “the natural” is also an epistemic authority: just as decisions about which types of organisms or communities we should protect have been made in the broader context of the artificial-natural dichotomy in environmental thinking, so to have decisions about what is to count as a valid object or method in biology.29 This argument is founded on the combination of two literatures: one devoted to understanding the moral authority of nature and the

27 See Daston and Vidal 2004 and Bensaude-Vincent and Newman 2007 for excellent historical-philosophical discussions and examples. For similar issues in environmental science from a variety of perspectives, see Shrader-Frechette and McCoy 1995, Cronon 1996a, Ereshefsky 2007, Davis et al. 2011. See also Sagoff 2001 for these issues in the context of genetic engineering. See Haraway 1991 for (challenging) discussions about narratives of “nature” and their political-scientific intentions and consequences. 28 For examples, see Sagoff 2001. 29 I have found three studies particularly helpful in this regard. Two are about the 18th century (“Human experimentation in the eighteenth century: natural boundaries and valid testing,” by Londa Schiebinger, and “Attention and the Values of Nature in the Enlightenment” by Lorraine Daston). While these studies touch on the evaluation of objects and methods of study, they are primarily concerned with uncovering the sources of the moral authority of nature, and so are more useful as formal guides rather than helpful with regard to content. The final paper, by historian Graeme Gooday, entitled “‘Nature’ in the Laboratory,” considers mid-19th century rhetorical and pedagogical uses of “nature” in arguments over the legitimacy of the microscope. Thomas Henry Huxley and others argued that the microscope had privileged access to nature by drawing on discourse traditionally found in natural history: one only truly experiences nature by looking through the microscope. With its focus on valid methods and instruments of study this study comes the closest to the topic of this dissertation.

!11 complex mechanisms which imbue nature with value; the other directed at more straightforwardly philosophy of science problems about the evaluation of scientific objects, methods, and evidential, experimental, and explanatory practices.

Let us consider a few examples. When geneticist Theodosius Dobzhansky referred to another biologist’s work derisively as being based on merely “laboratory flies!,” he was both describing the fact that these flies were bred in captivity rather than taken directly from the field (a description that depends as we will see on how the artificial-natural distinction is drawn), and exploiting the negative connotations that accompanied the term “laboratory,” a place of artificiality and artifice, in the eyes of some naturalists.30 To the consumate naturalist, what happens in the lab should stay in the lab—laboratories are not for studying natural things. As we will see in Chapter 6, particular views about what was “natural” and what was “artificial” played a role in structuring Dobzhansky’s research practice and his evaluations of others and their work.

The artificial-natural distinction also influences the choice and classification of research site and organism. When ecologist and co-founder of the Natural Capital Project,

Gretchen Daily, began ornithological research as a graduate student she “only had eyes for the rain forest.” It alone seemed to offer a pristine and untouched natural world, perfect for a budding conservationist. But following a failed attempt at studying bird species in the Costa

Rican rain forest, and unwilling to waste a summer of research, she turned to the farmland and surrounding countryside. And what she found, to her surprise, was that the “natural

30 Dobzhansky to Ernst Mayr, March 29, 1975. William Provine has argued that Dobzhansky was wrong about the flies because Alfred Sturtevant, the biologist he is referring to, used flies Dobzhansky had sent him directly from the mountains east of Pasadena (Provine 1981). I believe that Dobzhansky is more likely referring to the fact that Sturtevant used free-living populations of D. melanogaster, which he thought involved problems of human interaction, and were unsuitable for this reason. See Chapter 6 and Dobzhansky 1939, 358.

!12 world had never left this man-made system.” So began her life-long project to change the way we understand conservation by restoring an anthropocentric perspective (to the great frustration of her 1980s forebears).31 While she is now happy with the outcome, her initial disappointment in not being able to study undisturbed nature, was, and still is, a common reaction amongst conservation biologists—something she is attempting to change.32 This is to say, her initial choice of research site was influenced by what she at the time believed to be artificial and natural and her present work aims to undermine these widespread beliefs.

The biologist, and now philosopher, Massimo Pigliucci recalls a similar story from his graduate student days. When he decided to switch research organism from the

Connecticut native perennial, Lobelia, to the domesticated “weed,” and model organism,

Arabidopsis thaliana, his dissertation committee was appalled. The latter, they insisted,

“wasn’t a real plant.” Here, again, we see the interplay of Simon’s first and second criteria: this plant’s unnaturalness makes it less real. The resistance to Pigliucci’s decision is put forward against a backdrop of the artificial-natural dichotomy, in which “the natural” is understood to be preferable.33 In both of these cases—that of Daily and of Pigliucci—it is not just material or scientific features of the organisms or sites in question that determine biologists’ evaluation of them, but also wider considerations about what counts as artificial or natural and what this entails. Some objects and places are thought to have scientific or epistemic value precisely because they are thought not to be human disturbed.

31 We should not divorce the choices of research objects and sites made by conservation biologists and ecologists in the 1980s from the conviction, held among many college students in the 1960s and 70s, that everything artificial is bad, and everything natural is good (see Yankelovich 1972, 167-85. See also Rome 2013, 46.). 32 The story and quotes come from Voosen 2013. 33 The story and quotes from Pennisi 2000. See also Leonelli 2007, 212.

!13 Classifications according to the artificial-natural distinction extend beyond the objects, sites, methods, and other aspects of the biological “toolbox,” to styles of researchers themselves (or their typical personas). This case is much less straightforward (see Appendix

1). In the late-19th and early-20th centuries, for example, the terms “naturalist” and

“experimentalist” were used to classify two sets of biologists.34 The former studied nature

“undisturbed,” the latter made use of experimental methods. The former tended to draw a distinction between the artificial and the natural and approached experimental methods with skepticism; many of the latter did not draw the distinction, and few saw it as impinging on the validity of experimental methods. Such classifications similarly conceal normative dimensions behind seemingly descriptive claims.35 Each camp viewed the other as being naive, uninformed, or cognitively or methodologically limited; to classify one’s opponent as a member of the other category did not simply indicate different methodological commitments, but called into question her ability to produce valid research.36

The part played by concepts of the artificial and natural in biology is analogous to that played by what Bernard Williams called “thick concepts.” Williams coined this term in

34 See Allen 1979. 35 The case becomes even more complicated when these are used as historical categories as opposed to actors categories. For example, evolutionary ornithologist Ernst Mayr’s uses of the terms “naturalist” and “experimentalist” to classify sets of biologists’ contributions to the evolutionary synthesis movement of the 1930-40s, conceals a similarly normative dimension, as one acute commentator—the historian William Coleman—pointed out: according to Mayr, the naturalist had the right scientific character to properly recognize what the experimentalist could not. 36 In all of these ways, concepts of the artificial and the natural are used to express epistemic value. Such uses are not a new development in science. We might return to Aristotle. On a common reading, medieval Aristotelianism respected an ontological distinction between artificial and natural objects, as I said above. Natural objects possessed natures that marked them as one type of thing rather than another. Artificial objects, lacking such natures, also lacked ontological identity. Since natural philosophy—our “science”—aimed to describe “natures,” it focused on natural objects and the sorts of things they ordinarily do. Artificial objects, lacking natures, were not the object of study, nor were they helpful for learning about the objects of study. In this case, the artificial-natural distinction corresponds to one between invalid and valid objects of study. Given that our intellectual landscape has changed considerably since this time, to what extent does a similar sort of thing happen in current science and why? See Dear 1995, 2005.

!14 the context of moral theorizing to describe concepts such as treacherous or courageous.37

These are both “world-guided” in the sense that they refer to features of the world to guide their application, but they are also “action-guiding” insofar as they give us reason to, or not to, act. This is what I mean when I say that the artificial-natural distinction conceals underlying descriptive and normative dimensions. Classifying something as “natural” is to evaluate its scientific validity, not just to describe its properties. When embedded in larger norms of scientific behavior, as we saw in the case of Daily and Pigliucci’s choice of research organism and site, the artificial-natural distinction can have a direct impact on scientific practices. In other words, as a thick concept that goes beyond superficial description, natural, in opposition to artificial, acts as an epistemic authority that makes itself known through real decisions and practices. !

REVERSED ROLES FOR NATURAL AND ARTIFICIAL

When “natural” is invoked in the context of environmental thinking it typically corresponds to “good.” This is not the case for either term in the artificial-natural distinction in science. The early-20th century experimentalist Jacques Loeb, for example, lauded the artificial and promoted a “trivialization of nature” as part of his engineering ideal for biology.38 To Loeb, “the natural”—the nonhuman object of biological study—was just one amongst a set of possible states for investigation, and not one he considered particularly useful or interesting. What humans did in the laboratory was in no way “natural”—Loeb was

37 Thick concepts are contrasted with “thin concepts,” such as “good” or “right.” These concepts are action- guiding but not world-guided. See Williams 1985, Chapter 8. See also Jenkins 2006, Chapter 6, for a general discussion. 38 As it is put by Philip Pauly (1987).

!15 creating animals with bums on each end of their bodies—but this only added to the potential value of such work, as he saw it. For Loeb, “natural” was value-neutral at best.

In this dissertation, however, the cases I consider are biased towards those in which, when opposed to artificial, “natural” corresponds to good, valid, or advantageous, and

“artificial,” bad, invalid, or disadvantageous. There are three deliberate reasons for this bias.

First, I am primarily interested in the field practices of ecology and evolutionary biology, and in these disciplines, “natural” typically carries positive connotations. The easiest way to define the subfield of biology of which I’m interested is to follow an arbitrary division common to major biology departments, between (i) organismal and evolutionary biology and

(ii) molecular and cellular biology. While this leaves some fields unspecified it is fair to say that I focus predominantly on biologists in the former category. Second, I want to complicate a trend in historical-philosophical research that focuses on the rise of laboratory science in the 20th century and emphasizes the accompanying unnaturalness of much of modern science.39 It’s not that this history is misrepresentative, but that the problems it seeks to grapple with are not always exportable to the sciences of which I’m interested. To a laboratory experimentalist, many of the issues I consider may appear unintelligible, but that is the point. Third, and finally, I simply have insufficient space, time, and mental capacity to do justice to all subfields of biology. !

39 Perhaps the earliest example is Garland Allen’s Life Sciences in the Twentieth Century. We will consider this further in Chapter 6. The historical philosopher Hans-Jorg Rheinberger’s work is a good example of this focus on laboratory science. See Rheinberger 2010.

!16 THE COMMONPLACE VIEW: THE DISSOLUTION OF THE ARTIFICIAL-NATURAL DISTINCTION

Within historical literature about the origins of modern science, the artificial-natural distinction is often discussed in a different way still. A common narrative portrays the collapse of this distinction during the early modern period as central to the rise of modern experimental science. On this view, Aristotle’s distinction was nothing but a hindrance. It took the rise of the “mechanical” philosophy in the 17th century to put Aristotle’s distinction finally to rest and to found experimental science on its collapse. The recurrence of the distinction throughout the modern history of the life sciences should then be somewhat unexpected. Are the life sciences a “modern” science?

Furthermore, it is often suggested by early modern scholars that these concepts structured thought and practice in the ancient and early modern worlds more fundamentally and less vaguely than they do today.40 In an intellectual world that was dominated by order, hierarchies, correspondences and analogies, understanding the relation between “the vast and all-comprehending Dominions of Art and Nature” was part and parcel of understanding other similar categories: particular-universal, conventional-natural, passive-active, primitive- progressive, pastoral-courtly, etc.41 Discussions of the artificial-natural distinction were commonplace and radiated into many areas of social-political life. When Shakespeare toys with these concepts in The Winter’s Tale to produce both wit and social commentary he is assuming an audience which has been well-versed in how these concepts purportedly provide the world with a particular order. Historian Lorraine Daston writes that, compared to the

40 See Tayler 1961. See Daston and Park 1998. 41 Tayler 1961, 22ff. See also Foucault 2008, Chapter 5. See many examples throughout Daston and Vidal 2004 of the importance of these categories during the enlightenment.

!17 early modern period, “in the current metaphysical vernacular, the artificial has been swallowed up by the natural.”42 In other words, in the modern world, these categories have become less distinct and are thus, it often follows, less significant.43

These historical perspectives combine with one that focuses on the rise of laboratory experimentalism to support a view of contemporary science as fundamentally artificial, in which the artificial-natural distinction lacks relevance.44 This view grew out of a healthy reaction to earlier historical and philosophical literature that had a predilection for discovery in science. Although this is still a controversial topic in some quarters, many would now agree that much of current science is about creating, manufacturing or constructing phenomena. It is hard to deny this change in perspective when one considers the amount of human work necessary for designing and creating scientific apparatuses, producing and interpreting results, stabilizing results, as well as the rarity or even absence of the phenomena produced in the universe outside the laboratory.45 Science no longer works on Aristotle’s undisturbed nature, it is said, but on “artificial phenomena.”46 The epistemological advantages of the laboratory arise precisely from their not having to study natural objects as

42 Daston 1998, 153. 43 One way to push against this narrative would be to point to the work of Leo Marx (see Marx 2005). Marx shows that the idea of “nature,” along with its opposite “artificial,” were a powerful categories structuring early american social and political life (from the 18th century through to 1920 at least). The early puritans, for example, saw America as a “state of nature,” and took it as their religious duty to domesticate and civilize the savage land. 44 One edited volume which attempts to move past this view is Bensaude-Vincent and Newman’s The Artificial and the Natural: An Evolving Polarity (2007). Most of the essays, however, are about ancient and early modern periods, and there is nothing about the life sciences. None of the essays consider the concern that humans denature nature or, as I explain below, act as disturbing conditions, and thus it is only so helpful with regard to the following study. 45 See Hacking 1983 for early summary. See Golinski 2005, and Sismondo 2010, Chapter 14, “The Unnaturalness of Science and Technology,” for recent summaries. 46 See Hacking 1991, Kohler 1992, Knorr Cetina 1999; see Sismondo 2010, Chapter 14 for summary of literature.

!18 they are, where they are, or when they are.47 As philosopher of physics Nancy Cartwright writes, “nature in the mundane world seems obstinately unruly. Outside the supervision of the laboratory [...] what happens in one instance is rarely a guide to what will happen in others.”48 When it comes to the rise of model organisms as standing for wild ones, philosopher Sabina Leonelli argues that it is precisely their ability to transcend the artificial- natural boundary that is central to their scientific success.49 According to this recently popular view of science, the artificial-natural distinction is eroding away as modern science moves further from the unruly nature it once studied.50

Such perspectives might be contrasted with one arising from a smaller literature that often focuses on the field sciences and treats the artificial-natural distinction as both prevalent and important. Historian Robert Kohler concludes a study on the laboratory-field border—one of the modern manifestations of the artificial-natural distinction—with the following remark, “The line between nature and artifice can be blurred but not erased.

Natural places cannot be made so lablike that they become unnatural; laboratories cannot be made so natural that they lose the artifice that gives them their power.”51 Kohler’s remark should remind us that just because scholars of science understand the sharpness of the

47 See Knorr Cetina 1992, 117, as just one example. See references in Sismondo 2010, Chapter 14. 48 Cartwright 1999, 86. 49 Leonelli 2007. Donna Haraway’s work on cyborgs also draws attention to the multiple things which call this distinction into question. See Haraway 1991. 50 The question raised by this literature is whether science still studies “natural” objects and whether we are still primarily concerned with learning about the “natural” world rather than the engineered world. The microbiologist Carl Woese describes many of his molecular biologist colleagues as having lost the vision of understanding nature, instead caring only about technological advance (Woese 2004). Many science studies scholars agree with Woese’s assessment and see the phenomena studied as essentially artifactual, and thus also see current science as primarily focused on the artificial, not the natural—or, maybe more accurately, they understand the secret of modern science to have been the dissolution of these categories altogether. 51 Kohler 2002, 308.

!19 distinction to be somewhat dulled, and have rightly called attention to its permeability, we need not conclude that the distinction is unimportant, especially to field biologists themselves. Instead we should ask why the distinction remains important to these biologists, given that we see many problems with its conceptual coherence. For many working biologists, the artificial-natural distinction is real because it determines the conditions in which careers and decisions are made. What is real, as the Thomas Theorem conjectures, is real in its consequences. Moreover, the artificial-natural distinction might matter more to some sciences (and scientists) than to others, having to do with a complicated mix of history, ontology, and objectives. To many of the biologists we will consider, some parts of nature count as more natural and thus also as epistemically more valuable. !

METHODOLOGICAL ORIENTATION: HISTORICAL CONCEPTUAL ANALYSIS

As a philosopher, my analysis is fundamentally conceptual. This is not to suggest that

I am interested in seeking timeless definitions for “natural” and “artificial.” Rather, I am interested in how changes in their relationship have influenced the practice of biology from the 19th century to the present. As I have said above, I am most interested in those places in which “natural” is endowed with epistemic authority, and in which “the artificial” is approached uneasily, as if it possessed an air of unreality, or even deceitfulness. The present study aims to uncover some of the sources of this characterization.

I am indebted to three methodological approaches that combine the insights of history and philosophy. The first is the most straightforward. Philosophers often take it as their primary objective to question the hidden assumptions that inform arguments. I see myself as

!20 applying this methodological orientation to historical examples. In the following chapters I seek the hidden assumptions informing uses of the artificial-natural distinction.52

The second approach is best summarized by philosopher Alan Richardson:

only by attending to very specific themes and argumentative moves can one tease out quite different connotations in philosophical terms that one might be tempted to think were used identically across great stretches of time. [...] One virtue of the historiography of philosophy that I offer is that it makes the past history of philosophy more interesting, more nuanced, and more subtle; it is not the hack work of hundreds of years of people trying to work out one or two mistaken ideas. A more important virtue is that it can make the future history of philosophy more interesting. Just now it is hard to say that epistemology and philosophy of science have quite enunciated the problem(s) of knowledge appropriate to our time and place. If the historical record can reveal five or fifteen rather than one or two accounts of the problem of knowledge, we have that many more epistemological resources with which to think. I can think of nothing for which we contemporary philosophers could be more grateful.53 ! Richardson, and others like him, hold that detailed studies of history provide us with resources for thinking about our present state of affairs. The ways in which the artificial- natural distinction has been conceptualized in the past, for instance, might provide us with resources for thinking about the distinction at present. Of course, Richardson is discussing a more traditional philosophical problem—the problem of knowledge and the place of experience—but I see no reason why the artificial-natural distinction is not equally amenable

52 The difference between what I’m doing and history of ideas is a matter of degree. I am both more concerned with context than traditional history of ideas (in this case, my methodology is more similar to intellectual history), and more concerned with hidden assumptions informing the use of concepts than their broad transformations over time. Harvard history professor Peter Gordon has a nice essay on his faculty page, which I have found useful, entitled, “what is intellectual history.” This is not published and I’m not allowed to cite it without permission. 53 Richardson 2003, 66.

!21 to this manner of philosophical interrogation. We might ask, how does our current distinction differ from how Aristotle drew that distinction? This may give rise to a new way of thinking about our current distinction, or simply prompt us consider the assumptions behind our distinction, regardless of its similarity to Aristotle’s. Surely even this second scenario is enough to warrant the attempt.

The final methodology informing this work is that used by philosophers such as

Georges Canguilhem, Michel Foucault, and Ian Hacking, which often goes under the label

“history of the present.” This method pushes one to seek the historical conditions that made possible our present organization of concepts.54 One might think that in applying this method to problematic concepts like “child abuse” or “madness,” by telling their history and unearthing their preconditions, we would make them less problematic. My presentism, like

Hacking and Foucault, has a different aim. This is to problematize a distinction that many would see as mundane, inconsequential, or possibly even dissolved. For example, consider the phrase “in nature.” It is a common clause appended to the end of sentences within, and titles of, biological papers. I argue that the phrase “in nature” does not refer to a physical place, but to an ideal state involving no human disturbance. This should cause a moment’s pause and a question, why is this phrase so common? With what sort of epistemic weight is evidence valid “in nature” endowed? Why is it important to flag something that seems self- evident? Why not also add “is true”? We can only make sense of uses of the artificial-natural distinction by paying attention to how these concepts are and have been used and by thinking about how the history of their usage obliges us, as its heir, to think in a certain way.

54 Canguilhem 2008; Foucault 2007, Chapter 5; Hacking 2001, Chapter 3; See Rheinberger 2010, on Canguilhem.

!22 !

THEORETICAL RESOURCE 1: HUMANS AS “DISTURBING CONDITIONS”

I introduce and develop two theoretical resources that help facilitate the discussions that follow. The first treats human as “disturbing conditions.” In general terms I see contestations over the artificial-natural distinction in modern biology as negotiations about how and why human actions count as interferences or disturbances in the systems of biological study under discussion. The Oxford English Dictionary defines “disturbance” as

“interference with the regular or due course or continuance of any action or process.”

Philosophers of science also have a convenient and general way of characterizing such interfering factors. They call them “disturbing conditions.” Throughout the recent history of biology I argue (subject to necessary refinement in Chapter 3) that humans themselves have been thought of as disturbing conditions, in a more narrow philosophical sense, and that this is reflected in the artificial-natural language commonly used. It must be kept in mind, however, that what counts as “human disturbance,” “artificial,” or “natural” is constantly being redefined. Throughout this dissertation we will look at some of the ways in which humans have been thought of as disturbing conditions. !

THEORETICAL RESOURCE 2: META-LEVEL CLASSIFICATION

The second resource I develop treats the artificial-natural distinction as a classification system. In the past, philosophers have often been concerned with

“classification” in a narrow sense. For example, classification within the sciences, as when individuals are classified as tokens of a species type. Classification of this type is often

!23 discussed in relation to natural kinds: whether there are more objective or natural ways of carving up the world and whether we can know what these ways are.55 I am concerned with a much broader notion of “classification,” as discussed in science and technology studies literature, which is concerned about the kind of work that classification practices can do.56

This way of looking at classification focuses on how classifying things changes the way we act towards them and how particular ways of classifying become standardized, rather than whether there are “objective” classifications. We will look at how objects, broadly construed, in the life sciences are classified according to the artificial-natural distinction. Classifications, it will be argued, are consequential. They have the power to define disciplines. !

RELATION TO RECENT PHILOSOPHY OF BIOLOGY

Biology differs from other sciences in a number of ways. In comparison to physicists, biologists tend to be wary of grand theories and explanations, are not driven by the search for universal laws, use a set of heterogeneous methods and evaluation criteria, and are more concerned about the actual than the possible.57 A friend once wrote to me, “who wants to be a philosopher of biology if biology is just physics applied to organisms?” Indeed. I think that the artificial-natural distinction has an epistemic relevance in biology that it does not have in some other sciences, particularly the so-called “harder” sciences. And because philosophers of biology have been particularly invested in arguments about what does and does not make biology unique, this dissertation is properly in the scope of philosophy of biology.

55 See David Hull’s paper in Douglas and Hull 1992. 56 See Hacking 2001, Chapter 6; Bowker and Star 1999; Epstein 2007. 57 See for example Beatty 1995, Keller 2002, Waters 2007, Woodward 2010.

!24 Yet, my goal, and this I think is different from a currently popular approach—one

Michael Ruse characterized as “handmaiden to the science”—is not to aid the sciences or scientists so much as understand them and their historical development.58 The concepts I focus on are not concepts used by biologists, such as “natural selection” or “fitness.”

“Natural” and “artificial” are meta-level concepts that I argue are central to biology whether biologists recognize this or not (although some certainly do).

This study also impinges on a number of general themes in the philosophy of science.

My concern to understand how the artificial-natural distinction influences practical decisions about choice of object, site, or method, is driven by a concern to understand how biologists evaluate evidence. Controversies within biology are often about “relative significance,” for example the relative significance of natural selection versus random genetic drift. What is at issue in such cases is “the extent of applicability of a theory or mechanism within a domain

[...] not whether the theory or mechanism is the correct account of the domain.”59 Rather than performing crucial experiments, biologists often therefore end up amalgamating evidence from a number of various sources, like mathematics, computer simulations, laboratory experiments, field experiments, natural experiments, etc. to support a theory of their choice— biologist Jonathan Losos compares this approach to detective work.60 The question of how they evaluate different types and sources of evidence is a pressing one, and one that I think the artificial-natural distinction has some bearing on. !

58 Ruse 2008. 59 See Beatty 1995, 229. 60 See Losos 2007.

!25 LAYOUT OF THE DISSERTATION

Rather than following a single culminating narrative, the chapters of this dissertation offer reflections on the artificial-natural distinction from different philosophical and historical perspectives. The second chapter, Art, Nature and the Scientific Revolution, begins with the touchstone for discussions of the artificial-natural distinction in science: the

“scientific revolution.”61 I ask, what is it about this time period that compels historians and philosophers to consider how the artificial-natural distinction was drawn? The answer will be that the ideological origins of modern science—in particular the origin of modern experimentalism—is regarded as intimately tied to the collapse of an ancient ontological distinction between nature and artifice. I will develop this point through a brief intellectual history of this time period. This history will (i) introduce different ways of thinking about the artificial-natural relationship—many of which are still with us today—and help conceptually clarify these relationships for future discussions, and (ii) provide a reason why the artificial- natural distinction is often ignored or regarded as a relic of ancient debates no longer relevant.

At the end of Chapter 2 I draw out three philosophical tropes that will reemerge throughout the thesis. These are phrased as questions and are as follows: (i) how does the breakdown of the distinction relate to the rise of manipulative approaches to the study of nature? (ii) how do different ways of thinking about art and nature relate to one another and continue to exist in tandem? (iii) how did the ontological collapse of this distinction make room for the distinction to be drawn on epistemic grounds?

61 One exception is Bensaude-Vincent and Newman (2007).

!26 I begin Chapter 3, Denaturing Nature, with another intellectual history. I argue that, contrary to the narrative developed in Chapter 1, there has been a continuous invocation of the artificial-natural distinction in the biological sciences since the time of the scientific revolution when it was supposedly dissolved in all of science. This also shows that the artificial-natural distinction we draw on today is old, and that the problems surrounding it are long-standing. I provide a number of examples throughout the history of the life sciences where the artificial-natural distinction arises, saving the details of these cases for later chapters.

In the second half of Chapter 3 I will ask whether there is a common, general thread guiding the historical cases considered in the first half. I will suggest two. First, I explain what the artificial-natural distinction consists in, and argue that the distinction is a manifestation of the extent to which humans count as, what philosophers call, “disturbing conditions.” To draw the artificial-natural distinction is to acknowledge that humans are—for reasons which are context-dependent—disturbing conditions. Second, I explain that the artificial-natural distinction is used as a classification system; biologists use it widely to classify various aspects of their discipline. Drawing on literature in philosophy and science and technology studies, I explain that such classifications are not innocuous—they have practical consequences for the way in which we act towards the objects we classify. The chapters that follow aim to show some of these consequences, as they play out in the constituting of biological thought and practice.

Common to both the scientific revolution and the Darwinian revolution of the 19th century were discussions impinging on the relationship between art and nature. Was art a part

!27 of nature? Could art be used as a model for nature? In Chapter 4, The Art Itself is Nature, I consider this intellectual congruence and suggest that it is more than just nominal. Charles

Darwin and Asa Gray, for example, were well-aware of the 17th century debates which preceded them about the relation between art and nature through the works of such revered

English writers as William Shakespeare and Thomas Browne. Furthermore, they used their understandings of these debates to inform and express their own thinking about the relation between artificial and natural selection. Darwin’s reliance on the art of breeding as a central element in his theory of natural selection cohered well with 17th century texts that argued for a breakdown of the art-nature distinction by claiming that Nature was simply the art of God.

This chapter demonstrates one of the ways in which the artificial-natural distinction made its way from the literature we considered in Chapter 2 to the 19th century: the concepts are transferred, even though the exact concerns they are used to express change.

Building on the previous chapter, Chapter 5, Selection in the State of Nature, focuses more closely on how the artificial-natural distinction was understood and used by naturalists in the 19th century. I investigate a disagreement between Darwin and Alfred Russell Wallace with regard to Darwin’s famous analogy between artificial and natural selection. Though a proponent of Darwin’s views throughout his life, Wallace remained uneasy about Darwin’s principal analogy. I first show that the reason commonly provided by scholars for the disagreement is historically unfounded. I then approach their disagreement in a different light, by looking at how each of them reflected more broadly on the artificiality of domestication and on how these reflections interacted with the conclusions they drew about whether the process of domestication was representative of natural selection in the “state of

!28 nature.” Wallace—drawing on a long naturalist tradition—upholds the artificial-natural distinction and thus opposes Darwin’s analogy, whereas Darwin’s discussions act to collapse the distinction itself. In effect, Darwin’s discussions render unanswerable questions such as,

“is this a product of art or nature?” From these considerations, as well as those discussed in the last chapter, I conclude that different ways of conceiving of the artificial-natural distinction were used to think through and express deep seated differences of opinion about whether domesticated organisms could be used to shed light on natural species change.

Jumping ahead to the 1930s, Chapter 6, Searching for what Nature has Wrought, considers a period known as the “evolutionary synthesis.” Following a 1974 conference on the synthesis, the historian William Provine wrote to conference attendee Theodosius

Dobzhansky asking him how it felt to be singled out as “the perpetrator of the great synthesis of the 1930’s and 40’s.” Dobzhansky’s work was seen to have fruitfully blurred the line between the outdoor, “natural” world of naturalists and the indoor, “artificial” world of experimentalists—a line that had impeded the synthesis of evolution and genetics. The naturalist Ernst Mayr wrote to him, “I still remember how delighted I was when I read your first papers [...] I exclaimed at the time ‘Finally a geneticist who talks sense!’” In what way did Dobzhansky talk sense, while his experimental colleagues did not? To Mayr, his work on natural populations proved that he had an acquaintance with “nature” which was generally lacking among geneticists. Although Dobzhansky was perhaps concerned with “the natural” in a way that his immediate predecessors were not, his experimental practice was fundamentally similar to theirs and manipulative laboratory work continued to play a dominant and necessary role throughout his career.

!29 Questions abound: Why was “the natural” meaningful and useful for Dobzhansky?

Why did Dobzhansky believe that a study of evolution based on natural populations would provide different and better results than one on artificial populations? Given that his practice remained fundamentally laboratory-based, in what way did he think his experimental practice met this “natural” goal? I address these questions by considering a crucial experimental move

Dobzhansky made in the 1930s: a switch of research organism, from the—in his words

—“domestic,” “artificial,” or “cosmopolitan” fruit fly, D. melanogaster, to its “wild,”

“natural,” or “noble” cousin D. pseudoobscura. I argue for two theses: (i) that assumptions about what was artificial and what natural played a role in structuring Dobzhansky’s research practice and his evaluations of others and their work; (ii) Dobzhansky was skeptical of studies that were too artificial, and this skepticism was reinforced throughout his career, because he believed they promoted both misrepresentative and undesirable views of the nature and evolutionary consequences of genetic variation.

In the final episode, Chapter 7, Communities, Natural Experiments, and “Soft”

Science, I consider 1980s community ecology and the use of natural experiments therein. In all its oxymoronic guise, the idea of the “natural” experiment has been a valuable one throughout the history of biology. It simultaneously possesses both the authority to speak about nature as well as the rigor associated with the rise of laboratory experimentalism. Thus, bringing the artificial-natural distinction back into the heart of the territory in which it was apparently collapsed. But, why exactly is it advantageous that an experiment be “performed” by nature rather than by human experimenters? And why has the artificial-natural distinction been a useful and commonplace way for biologists to classify the types of experimental work

!30 done in biology? Community ecologists confronted these questions in the 1980s during a significant internal dispute about what it meant to do “good” science. This chapter will address these questions by focusing on ecologist Jared Diamond’s use of natural experiments and his widely cited and disputed reflections on proper experimental methodology. I will argue that, contra any narratives about the collapse of the distinction due to the rise of experimentalism, concepts of “the natural” and “the artificial” have been central to experimentation in ecology, particularly as a way to clarify and maintain methodological distance between ecology and the so-called “harder” sciences. This will also give us reason to think that the artificial-natural distinction in biology is here to stay.

The final chapter, Chapter 8, Conclusion, is a summary which showcases limitations and possible future research directions.

!31 ! ! CHAPTER 2 ! Art, Nature, and the Scientific Revolution Making Modern Experimental Science ! ! ! ! ! ! ! ! For even as in the business of life, a man’s disposition and secret workings of his mind and affections are better discovered when he is in trouble than at other times, so likewise the secrets of nature reveal themselves more readily under the vexations of art than when they go their own way. Francis Bacon, 1620, Novum Organum, Book One !

The touchstone for discussions of the artificial-natural distinction in science continues to be the scientific revolution.62 In addressing the literature on this topic I seek to understand what it is about this time period that compels us to consider the artificial-natural distinction?

I will (i) introduce different ways of thinking about the artificial-natural relationship and help clarify these relationships for future discussions, and (ii) provide a reason why in current discussions of science the artificial-natural distinction is ignored or seen as a relic of

62 One discussion which goes beyond the scientific revolution is Bensaude-Vincent and Newman’s 2007 edited volume The Artificial and the Natural: An Evolving Polarity. This volume is the first historical treatment of the distinction and while it does consider more than the scientific revolution, such discussions are rare. Most of the volume is about ancient science and the scientific revolution period.

!32 antiquated, irrelevant debates. The business of the next chapter will be to complicate and undermine this idea.63 !

THE MYTHIC ORIGINS OF MODERN SCIENCE

What was the scientific revolution? Historian Steven Shapin opens his recent introduction to the topic with the following provocative assertion: “There was no such thing as the Scientific Revolution, and this is a book about it.”64 This is as good a characterization of the current state of affairs as any. It reflects the paradoxical place of revolutions in the history of science.

On the one hand, there is general agreement that between roughly the 16th and 18th centuries large-scale transformations occurred in discourses, practices, institutions, and attitudes that aimed at explaining, understanding and controlling nature.65 It is believed that through many of these transformations aspects of our current scientific methodology become visible, if only in the broadest outlines. Three famous examples will make this point: (i) the term “law” was applied to regularities found in the natural world and nature itself thus came to be seen as lawful or law-governed; (ii) mathematics was applied liberally to describe phenomena throughout the natural world. The Italian astronomer Galileo Galilei, for

63 One may well ask, why bother to tell the historical story at all? One reason is that the stories we tell about the past significantly influence how we view the present. This is especially so for deeply engrained stories, such as those about the “scientific revolution” and the origins of “modern science.” By reevaluating a theme central to these stories, I think we will come to see the present in a new light and in a way that raises a host of philosophical issues. This is not to say that the following is a Whig history—one that assesses the accomplishments of the past in terms of current standards. I will suggest rather that we alter our current standards through a study of their prehistory. 64 Shapin 1996, 1. 65 See Cohen 1994 and the references therein. For classic, but not so old, discussions see, Westfall 1971; Kuhn 1976; Wilson 1995 as well as footnotes below.

!33 example, is famously paraphrased as saying that the book of nature was written in the language of mathematics; (iii) the merging of the mechanical arts with natural philosophy led to an increased use of experiments and instruments.66

Yet, the once popular grand narratives about the scientific revolution, citing such canonical names as Nicolaus Copernicus, Johannes Kepler, Galileo, René Descartes, Francis

Bacon, Robert Boyle, and Isaac Newton, no longer convince.67 Early on, many studies of the scientific revolution sought to explain how today’s science came to be. The emphasis has shifted towards understanding what these transformations meant for those living in the 17th century, rather than how they do or do not contribute to the progress towards modern science.

Hardly anyone today would disagree that the activities of early moderns were too various to be penned into a single, coherent story about the genesis and rise of modern science.

Furthermore, today’s science does not map onto early modern science in any simple intelligible fashion. Our “science” was spread out over natural philosophy, natural history, alchemy, mixed mathematics, and the mechanical arts, each with its own styles of reasoning,

66 For these reasons, historians spent a good chunk of the 20th century sorting out both what was so “new”—as the early modern thinkers themselves put it—about these ways of studying nature, and the ever-increasingly complicated mishmash of social-cultural, scientific, technological and political causes that gave rise to such novelty. For a nice overview see Wilson 1995, Chapter 1. For laws see Ruby 1986; Milton 1998; for mathematics see Westfall 1971, Dear 1995; for experimentation see Shapin & Schaffer 1985. See Kuhn for 1976 for a classic discussion of the differences between the experimental and mathematical traditions and the changes occurring in each. Although it existed before, the phrase “Scientific Revolution” was given wider readership by Alexandre Koyré in 1939. See Cohen 1994 and Shapin 1996 for excellent discussions of these causes and their historiographies. See Lindberg and Westman 1990 and Osler 2000 for reappraisals of the themes that dominated early historical research surrounding the Scientific Revolution. See Park and Daston 2008 for the early modern use of “new.” 67 See Dijksterhuis 1961; Koyré 1957; Burtt 1925; Butterfield 1950; Cohen 1994.

!34 methods, and goals.68 Our idea of the “renaissance man” is likely more a manifestation of our ignorance of how early modern areas of knowledge were sub-divided, than it is of the abilities of early moderns to transcend disciplines.

As with many other grand narratives, such as those of the Enlightenment or

Modernism, the scientific revolution has taken on a mythic quality.69 In spite of, or perhaps because of, its paradoxical nature, the scientific revolution still enters into academic and popular discussions of the rise of modern science and in doing so reflects what we take to be important about science and influences how we think about current science.70 This “myth,” as historians Katherine Park and Lorraine Daston put it, “expresses in condensed and sometimes emblematic form themes too deep to be unsettled by mere facts, however plentiful and persuasive,” and “holds both proponents and opponents in its thrall.”71 As an origin myth, it acts to reinforce our current conceptions of the essence of science and, if only implicitly, what marks science from other intellectual endeavors and cultural practices.72 The philosopher Evelyn Fox Keller wrote that “Unexamined myths, wherever they survive, have a subterranean potency; they affect our thinking in ways we are not aware of, and to the

68 See Park and Daston 2008, 3-5, for a discussion of the different knowledge-producing bodies/practices/ disciplines in the early modern period. See Willson 1995, 11ff for a discussion of how these different disciplines entered into university curriculum. See Hacking 2001 for a discussion of different “styles of reasoning.” Each discipline typically dealt with the following: natural philosophy with organic and physical change, natural history described nature’s particulars often for medical purposes, alchemy with distillation and the transmutation of metals, mixed mathematics with geometry, astronomy, optics, and harmonics, and the mechanical arts with architecture, engineering, clock-making, agriculture, breeding, horticulture. 69 See Park and Daston 2008. See also Wilson 1995, who also refers to the scientific revolution as mythic. 70 Peter Dear’s recent Revolutionizing the Sciences (2009) is another example. See for a reflection on this issue Cunningham and Williams 1993. 71 Park and Daston 2008, 15. See also a great discussion by Cunningham and Williams (1993) about the ‘big picture’ version of the scientific revolution, and why this is inadequate in light of our current historiographical aims and more recent history of science. 72 For some thoughts on the function of myths see Kirk 1970.

!35 extent that we lack awareness, our capacity to resist their influence is undermined.”73 While the scientific revolution is hardly unexamined, there are aspects of it that continue to influence our thinking in mythic ways.

Here I characterize one particular aspect of the mythology. By rehearsing the myth my point is not to reinforce a just-so story about the early modern period, but to tell an intentionally selective story that illuminates what many have found to be a particularly compelling feature.74 Doing so will put us in a better position to reflect philosophically on the extent to which this aspect of the making of modern science is indeed true of science and to assess the influence it has had on our thinking about science.

The aspect of the scientific revolution I want to analyze is the “breakdown” of the artificial-natural distinction. This breakdown enters into discussions of early modern science in several ways. Lorraine Daston writes, “The insight that art was merely a part of nature, governed by the same rules was of cardinal importance to the early stages of the Scientific

Revolution [...] The central precept of the mechanical philosophy, that nature was actually composed of microscopic machines, presupposed the destruction of the boundary between works of nature and handiwork.” Steven Shapin writes, “Unless it was accepted that there was a basic similarity between the products of nature and those of human artifice, experimental manipulations with machines could not stand for how things were in nature [...] there could be no secure inference from what experimental apparatus made manifest to the natural order of things.” Ian Hacking says that Robert Boyle’s air-pump “inaugurated

73 Keller 1985, 76. 74 For a much better introduction to the origins of modern science, that is pluralistic both in terms of episodes, time periods, sciences, and larger historiographic themes see Bowler and Morus 2005.

!36 laboratory science. Before the air pump, one aimed at solving the phenomena given in nature

[...] Afterwards a new kind of science answered to [...] phenomena that fleetingly exist by artifice.” Today, Hacking continues, “the category of matters of fact, answering to artificially produced phenomena, is sacrosanct.” Richard Westfall writes, “It is difficult to speak of natural phenomena in relation to [Newton's experiments ...] the design of the experiment determined that nature had no choice but to answer 'yes' or 'no.' [...] By the end of the 17th century, the scientific revolution had forged an instrument of investigation that it has wielded ever since.” And finally, H. Floris Cohen writes that “Every interested thinker in the 17th century had to grapple with the distinction between ‘natural’ and ‘artificially produced’ nature and the root issue of how one was related to the other.”75

We can see that the collapse of the artificial-natural distinction is an oft-cited idea in the secondary literature. Let me move to the history of the breakdown of this distinction and its relevance for science in the early modern period. !

THE ONTOLOGICAL BREAKDOWN OF THE ARTIFICIAL-NATURAL DISTINCTION

The early moderns responsible for the scientific revolution inherited the distinction between the artificial and the natural from a scholastic understanding of natural philosophy.

The text widely cited, in this regard, throughout the 17th century was Aristotle’s treatise on

Physics.76 Consider the following passage, !

75 Daston 1988, 464; Shapin 1996, 97-8; Hacking 1991, 235 and 240; Westfall 1971, 116; Cohen 1994, 188 (respectively) 76 Daston and Park 1998, 263-4. See Close 1969.

!37 Of the things that exist, some exist by nature, some from other causes. By nature the animals and their parts exist, and the plants and simple bodies (earth, fire, air, water) - for we say that these and the like exist by nature. All the things mentioned plainly differ from things which are not constituted by nature. For each of them has within itself a principle of motion and of stationariness (in respect of place, or of growth and decrease, or by way of alteration). On the other hand, a bed and a coat and anything else of that sort, qua receiving those designations - i.e. in so far as they are products of art - have no innate impulse to change.77 ! For Aristotle, distinguishing between art and nature meant distinguishing between external sources of change and internal sources of change.78 The natural object contained within itself a principle of motion to change or develop in accordance with its nature. The nature of an acorn is to develop into an oak tree. The acorn, in other words, does this regardless of any external influence. The “natures” of natural objects were to be determined, according to this tradition, by their ends.79

The artificial object, in contrast, does not have an internal nature or essence. Its source of change is imposed externally by us—what Aristotle calls accidental, rather than substantial. There is nothing that has as its nature to develop into a coat or a bed. We make these things by using, combining and in some cases thwarting, the natures of natural objects.

Artificial objects are thus “necessarily posterior to and parasitic upon natural objects.”80

Artificial objects, since they lacked natures, also lacked a clear ontological identity. What kind of thing an artifact was could not be specified by its nature, since it did not have one.

77 Aristotle Physics in Works, vol. 1, 329; quoted in Daston and Park 1998, 264. 78 See discussions of this passage as compared to Aristotle’s work in other areas in Schiefsky 2007. 79 See Dear 1995, 153-5. 80 Daston and Park 1998, 264.

!38 The artificial-natural distinction was an ontological distinction; artificial and natural objects differed in kind, not in degree.81

The understanding of “natural philosophy” that dominated in medieval universities prior to the scientific revolution, was Aristotelian in the above sense. It was, according to historian Peter Dear, a contemplative endeavor, aimed at understanding the natural world in terms of how things happened most of the time or as per their usual course.82 This contemplative endeavor can be juxtaposed with how “Art” was understood in classical antiquity and preserved through the renaissance: art was “any rationally organized activity which has a practical rather than a speculative end (e.g. rhetoric, carpentry, politics, painting, drama), and as the system of theoretical knowledge or the intellectual expertise or the technical proficiency which such activities presuppose.”83 Art was practical and instrumental.

In contrast to the instrumentality of art, natural philosophy promoted an understanding that was intricately tied to the Aristotelian idea of “natures.” It aimed at providing an account of the natural world in terms of the natures of objects. This difference lay at the heart of distinctions in natural philosophy between, for instance, violent motion and natural motion. Smoke rises, it was supposed, because it is composed primarily of the element air whose natural place is above the earth, which in turn falls to its natural place at the centre of the universe. Since the Aristotelians were concerned with understanding the natural motion of physical bodies, the study of human artifacts—which operated according to

81 Because of the entrenchment of this view, historians call the opposition of art and nature during the renaissance a “habit of the understanding” or “conceptual reflex.” See Park and Daston 2006, and also Tayler 1964 and Close 1969. 82 See Dear 1990, 680ff; Dear 2005, 393-4; Hansen 1986, 129-30; Daston 1991, 340-1; Shapin 1996, 30-46; Newman 2004; Reill 2008, 26; Merchant 1980. 83 Close 1969, 467.

!39 violent motion—was of less value.84 The launched projectile, which disobeyed its natural inclination to fall, told of what humans could do rather than what happened by nature. One

17th century thinker, reflecting on this tradition and its particular distinction between types of motion, wrote, this “seems much to favour the Opinion of the Naturists, since ‘tis grounded upon a Supposition, that what is violent, is, as such, contrary to Nature.”85 Similarly machines and other artificial constructions functioned because humans had constrained their natural motion.86 In order for these types of motion to fall under the purview of natural philosophy, its epistemological and ontological foundations had to be reconceptualized.87

The ontological opposition between art and nature began to break down as the goal of natural philosophical knowledge changed from one of contemplation to one of instrumentalism.88 This gradual placing of natural knowledge in the service of the improvement of arts and trades was brought on, as historians have pointed out, by a plethora of factors present by the early modern period, including the increasing role that machines— such as large clocks erected in town centers—played in day-to-day human affairs, the wide- spread beliefs that we can securely know only what we construct by hand and that the authority of ancient texts was secondary to the “authority of nature” known by personal experience, influential observations of various heavenly phenomena through the telescope and other discoveries using mechanical instruments, and political instability that called into

84 Violent motion is also called “constrained” or “artificial” motion. 85 Boyle [1686] 1999, 507. 86 DesChene 2007. 87 Also required was a new understanding of “experience” itself. See Dear 2005. Add a note here about this. 88 Daston and Park (1998) examine how this change was preceded by challenges to the art-nature distinction through the Wunderkammern. See Daston 1995, 40ff and especially Dear 2005, 2006. As shown by Sylvia Berryman (2009; see especially the Appendix), the mechanical philosophers were not reacting to or rejecting ancient philosophy directly—in fact, many of the ways in which “mechanical” was used in the early modern period have their roots in antiquity.

!40 question the established uses of natural philosophical knowledge, and created a climate in which political authority could be symbolized by the sovereignty’s dominion over nature.89

An early example of this instrumentalism combined with a reexamination of the art-nature distinction can be found in the work of philosopher Francis Bacon.

Bacon opposed the glorification of Aristotelian natural philosophy at the expense of the mechanical arts. He accused his philosophical predecessors of illegitimately employing the art-nature divide as an excuse to focus on nature in its unrestrained state (natura in cursu), at the expense of nature “constrained, moulded, translated, and made as it were new by art and the hand of man” (natura vexata).90 In doing so, Bacon believed they had failed to grasp the benefits of the crafts for improving the human condition, and even worse, had founded philosophy on an “impoverished and inadequately evaluated, stock of experience,” comparable to founding a kingdom upon the mere “gossip of the streets.”91 “The secrets of nature,” he wrote in opposition to the Aristotelian view, “reveal themselves more readily under the vexation of art than when they go their own way.”92

This was more than just an opposition, it was a complete reversal of the aim of traditional natural philosophy. Bacon was clear that the ontological distinction between art and nature, that which had given rise to a reverence for “nature” over art, was no longer tenable.93 He wrote,

89 See broad social histories of the period: Shapin 1996, 123ff and his bibliographic essay on this topic. See also Shapin and Shaffer 1985. 90 Bacon 1861-1879, 506. See also Newman 2004, 258 91 Bacon 1620 [1999], Book 1, §63 92 Ibid. 93 The reverence for nature Bacon attributed to natural philosophers is more a reaction to that dominant in his own time than to what the ancients actually did or said. See Schiefsky 2007 for a discussion of this point.

!41 ! it is the fashion to talk as if art were something different from nature, so that things artificial should be separated from things natural, as differing totally in kind; whence it comes that most writers of natural history think it enough to make a history of animals or plants or minerals, without mentioning the experiments of mechanical arts (which are far the most important for philosophy); and not only that, but another and more subtle error finds its way into men’s minds; that of looking upon art merely as a kind of supplement to nature; which has power enough to finish what nature has begun or correct her when going aside, but no power to make radical changes, and shake her in the foundations; an opinion which has brought a great deal of despair to human concerns. [...] things artificial differ from things natural only in the efficient [cause]94 ! This discussion is unmistakably directed at Aristotelian understandings of natural philosophy based on a distinction between art and nature. Bacon, redraws the distinction between art and nature not in terms of natures or essences, but in terms of their efficient causes, that is, the immediate processes which bring them about.95 Nature, personified, is now an artisan; just as an artisan makes an artificial object, so nature makes a natural object (Figure 2.1).96 It is in this sense which the natural world is, for Bacon, highly wrought—the world is contrived in the same way that humans contrive mechanical engines. This lead Bacon to claim that his history of the arts, an important part of his redefined philosophy, was a “mechanical” history.97 The word “mechanical” came to greater prominence in the later 17th century, but in

94 Bacon 1861-1879, 506. 95 In this way he is taping into a distinct ancient tradition of thinking about the creative forces in the cosmos and their relation to one another. Commonly these were: art, nature, and chance. See Close 1969. 96 See Daston 1995. 97 Ibid., 506.

!42 a way subtly yet importantly different from what Bacon intended, and corresponding to a different understanding of the relation between art and nature. !

! Figure 2.1: The Mirror of Nature and the Image of Art. During the renaissance it was common to portray Nature, or Natura, as an artisan giving the task of looking over the earthly realm. This image comes from Robert Fludd’s 1617 Utriusque cosmi maioris scilicet et minoris metaphysica, physica atque technica historia. The hand of God, emerging from the cloud in the upper middle, holds a chain that symbolizes His mastery of Natura. Natura similarly holds a chain attached to simian, Ars, holding a globe. This symbolizes the attempts of art to ape the creations of Nature and ultimately God. (From: Tayler, Nature and Art in Renaissance Literature.) (© Wellcome Library, London) !

!43 Bacon’s philosophy was promoted most ardently in the later 17th century by members of the Royal Society of London (founded 1660) through the so-called “experimental” or

“mechanical” philosophy. These philosophers advocated similarly instrumental goals. One prominent member, Joseph Glanvill, writes that the objective of natural philosophy is to

“enlarge knowledge by observation and experiment [...] so that nature being known it may be mastered, managed, and used in the services of human life.”98

The mechanical philosophy also took their means—the experiment—from Baconian philosophy. As the quote above suggests, Bacon significantly elevated the role and status of experiments in natural philosophy.99 He saw experiments as not just “thought experiments” and not just for demonstrating a conclusion known in advance or deduced from theory, but as a means to see how nature would behave under previously unobserved circumstances that could not have been brought about without human intervention.100 This method he called,

“twisting the lion’s tail.”101 When later members of the Royal Society experimented by putting rodents into an air pump and evacuated the air inside, they followed in the wake of

Bacon’s experimental methodology (Figure 2.2).

98 Glanvill 1668 [1958]. See Merchant 1980 for a broader discussion. 99 See Kuhn 1976, 41ff for a discussion of these themes. 100 As an example of simply demonstrating prior knowledge, Kuhn writes, “Roger Bacon writes that, though one can in principle deduce the ability of flame to burn flesh, it is more conclusive, given the mind’s propensity for error, to place one’s hand in the fire” (Kuhn 1976, 43). 101 Kuhn 1976, 44.

!44 Figure 2.2: An Air Pump Used by Robert Boyle. If you look closely you’ll notice that there is an unhappy rodent in the glass chamber. From: Boyle’s 1669 A Continuation of New Experiments Physico-Mechanical, Touching the Spring and Weight of Air. (© Wellcome Library, London) ! However, the mechanical philosophy advocated by the Royal Society was more than just an elaboration of Baconian methods. It involved a new understanding of the relation between art and nature that grounded arguments for the primacy of experimentalism as a method of gaining natural knowledge. This new understanding of nature was influenced by

French thinkers like René Descartes, and while it was nominally similar to Bacon’s—it was

!45 also called “mechanical”—it proposed something very different.102 Robert Boyle was the foremost proponent of this mechanical-experimental philosophy and he is for this reason often praised as the father of experimental science. Rather than an artisan, Boyle likened nature to an intricate, but passive, work of art: an engine, such as a clock. He offered the following understanding of nature: ! ‘tis like a rare Clock, such as may be that at Strasbourg, where all things are so skillfully contriv’d, that the Engine being once set a Moving, all things proceed according to the Artificers first design, and the Motions of the little Statues [...] do not require [...] the peculiar interposing of the Artificer, or any intelligent agent imployed by him, but perform their functions on particular occasions, by vertue of the General and Primitive Contrivance of the whole Engine.103 ! For Boyle, nature was artificial in the sense that it was God’s work of art. This collapsed the gap between art and nature more fundamentally than had Bacon: the natural was essentially subsumed under the artificial.104 Like Bacon, nature was highly wrought; but unlike Bacon, this was not because Nature, or Natura, brings about its objects in the same contrived way as humans bring about artificial objects.105 It was because nature was nothing other than a very complicated mechanical engine. Against the distinction between natural and violent motion,

102 Regarding “Mechanical”: it has four separate senses throughout the scientific revolution that can be confusing. (i) Mechanical as in the mechanical arts; both highly wrought and made with hands (e.g., Francis Bacon). (ii) Mechanical as in like a machine; being characterized by uniformity and regularity (e.g., Robert Boyle); (iii) Mechanical as in made of parts or corpuscles (e.g., René Descartes); (iv) Mechanical as in can be described by the language of mathematics (e.g., Isaac Newton). Regarding the influence of René Descartes; he wrote for example that “it is no less natural [...] for a clock constructed with this or that set of wheels to tell the time than it is for a tree which grew from this or that seed to produce the appropriate fruit” (Descartes 1998[1644], 209[326]). 103 Boyle 1686 [1999], 448. 104 This is the starting place for natural theological arguments from design. Nature’s contrivance is indicative of its creator, in the same way that a watches contrivance is indicative of a watchmaker. 105 See Merchant 1980, 6-7, on the related distinction between natura naturata (the natural creation) and natura naturans (nature as a creative force).

!46 Boyle wrote, “It may be justly Question’d [...] Whether there be any Motion, among

Inanimate Bodies, that deserves to be call’d Violent, in Contradistinction to Natural”, since

“among such, all Motions [...] are made according to Catholick, and almost, if not more than almost, Mechanical Laws.”106

This account, as Boyle laid out in his Vulgarly Received Notion of Nature (1686), was prompted by fears that ancient understandings of nature, along with those of Bacon, were anthropomorphic and stole praise away from God by portraying Nature as an artisan or as having some degree of autonomy or vitality. To Boyle, God needed no intermediary; nature was simply a machine and God the engineer that put the clockwork system in motion. (We will return to Boyle’s view in Chapter 4.)

The mechanical philosophers argued that traditional natural philosophical explanations of nature were—to borrow their favorite terms of abuse—unintelligible, animistic and anthropomorphic. The ancient saws “Nature does nothing in vain” and “Nature abhors a vacuum” lacked explanatory power, they argued. To the explanation that bodies descended because they were heavy, philosopher Thomas Hobbes responded, “But if you ask what they mean by heaviness, they will define it to be an endeavour to go to the centre of the earth [...] which is as much to say that bodies descend, or ascend, because they do.”107 The

French natural philosopher Pierre Gassendi wrote of Aristotle’s definition of motion, “the act of being in potentiality insofar as it is in potentiality”: “Great God! Is there any stomach strong enough to digest that?”108 In contrast, many mechanical philosophers offered bottom-

106 Boyle 1686 [1999], 507. 107 Hobbes 1651 [2002], 504 108 Cited in Cartwright 1999, 78.

!47 up explanations involving the interaction of variously shaped, invisible, micro-level corpuscles. The interactions proposed were analogous to the ways in which everyday, macro- level bodies interacted and hence, they argued, intelligible.

The view of nature as a passive work of artifice cohered well with the rise of experimentalism and with the belief that nature, highly regular and uniform, can be described according to laws. When nature became machine-like, it was easy to see why experiments— and in general methods involving significant manipulations of nature’s mechanisms—were deemed useful.109 Recall the oft-cited analogy to a clockmaker. Just as a clock is the product of a designer, so nature was seen as a great engineering feat set in motion by the hand of

God. If nature was like a clock, one should approach it in the same way as a clockmaker approaches a foreign watch: determine the laws which govern the regular and uniform movement of its variously sized and shaped parts by taking it apart and studying its pieces.

Joseph Glanvill writes, for instance, that anatomy was useful because it tended

“mightily to the eviscerating of nature, and disclosure of the springs of its motion,” and the microscope important because “the secrets of nature are not in the greater masses, but in those little threads and springs which are too subtle for the grossness of our unhelped senses.”110 The metaphor runs deep since it could be employed to justify experimental or manipulative practices and the uses of instruments which helped one get beyond the unmediated senses or created unnatural environments. As the distinction between the artificial and the natural broke down, the methods appropriate for the study of machines and

109 There is a difference to be pointed out here. See the beginning of Wilson 1995. Nature as a machine versus nature is to a machine as a clock-face is to clock mechanism. 110 Glanvill 1668 [1958], 1668.

!48 the mechanical arts, and the rigor these philosophers saw accompanying these methods, could be applied to the study of nature. And so, it is said, begins modern experimentalism.

The narrative I have summarized concerns the ideological origins of the experimental methods we see as partly constitutive of modern science. The scientific revolution is characterized as a period when the study of nature changes from one of passive observation to one of experimental manipulation and control; from the study of “natural phenomena” in their due course to the study of “artificial phenomena” hidden from the immediate senses but brought out through human intervention under artificial circumstances.111 This was the scientific revolution because we seen in it the dim origins of our own scientific attitudes, discourses, and practices.112 Accompanying this change is a reconceptualization of the art- nature relationship, from the ontological distinction drawn by Aristotle, through Bacon’s likening of nature to a human artisan, to nature as artificial, in the sense of being highly regular and uniform, describable in terms of laws of nature, and modular or reducible to independent parts.

Before analyzing this narrative, it is useful to take stock of another important and somewhat paradoxical idea which is often said to emerge during the scientific revolution: the idea that humans are separate from, or outside of, nature.113 This theme is evident in the political sphere, for instance, when Thomas Hobbes theorized that the formation of human society was a flight from the “state of nature.” The idea is also instantiated in Descartes' res

111 See amendments to this standard history from especially William Newman: Newman 2004, 2006; Bensaude- Vincent and Newman 2007; see also Moran 2005. 112 See Wilson 1995. 113 Paradoxical because this is seen to arise through the same historical forces that resulted in the breakdown of the art-nature distinction. See Merchant 1980, Williams 1980; Daston 1995; Cronon 1995a, b.

!49 cogitans and mind-body dualism. As a theme of the literature on method in the scientific revolution, however, it is captured by the discourse of “experimentation on nature,”

“manipulation of nature,” and “intervention in nature” which became prevalent during this time. Humans were intelligent and could choose to act in certain ways, but the nature that they they studied and experimented on was “‘brute, passive, stupid matter’ in seventeenth century parlance.”114 This view of nature was in a sense necessary for the experimental methodology of the scientific revolution. The methodology required that humans view nature as controllable, lacking a will of its own, and thus also to a large extent as separate from themselves.115 !

ANALYZING THIS MAKING OF MODERN SCIENCE

In analyzing this narrative, the historian would point out the ways in which it misrepresents the diversity of projects of the early moderns or their predecessors.116 I am not qualified to analyze the narrative in this way, and it is not my intention to do so. I am interested in what it advocates as the central aspects of modern science, here represented in their proto-form. In this regard, the narrative prompts three philosophical questions to which we will return throughout the dissertation. The questions are, (i) how does the collapse of the distinction relate to the justification of manipulative approaches to the study of nature? (Or, its converse: how does drawing the distinction relate to arguments against the use of manipulative approaches?) (ii) how do these different ways of thinking about art and nature

114 Daston 1995, 39. 115 Carolyn Merchant’s work has focused most specifically on the alienation of nature during this time period. See Merchant 1980. 116 The clearest example of this project is Newman 1997.

!50 relate to one another? (iii) how did this ontological breakdown make room for the distinction to be drawn on other grounds?

I mean to pose the first question in a philosophical manner, rather than historical; the question is about justification in general, rather than what Boyle or Glanvill thought counted as justification.117 According to the medieval Aristotelian position considered above, for example, there is an ontological difference between artificial and natural objects and this is paired with a particular epistemic goal to understand what happens ordinarily by nature. This pairing counted against the usefulness—in natural philosophy—of experiments in the

Baconian sense of “twisting the lion’s tail.” Such experiments demonstrated human control over nature, useful for practical purposes only.118 Humans are in this case seen as disturbances or accidents in nature’s ordinary course. Consider 16th century natural philosopher Thomas Erastus’ objections to alchemy, and 17th century alchemist Daniel

Sennert’s answers. Erastus denied that “the ‘chymical resolutions’ [...] can ever reveal the true principles of things, because they are ‘not natural, but artificial.’”119 Sennert responded, in a way similar to Boyle, “But if you consider the proximal agent of mixture, it must be denied that chymical resolutions are not natural, even if the artisan participates in his own fashion. They are brought out by fire and heat, by means of a natural cause.”120 For opponents of the alchemical proto-experimentalism, such as Erastus, Sennert’s response

117 Historically, the connection between experimentalism and the breakdown of the artificial-natural distinction is not necessary (although the connection was exploited by the Royal Society). There were certainly experiments being done long before the mechanical philosophy. No one has argued this point more forcefully than William Newman. See Newman 1997, 2006. 118 As one historian writes of pre-Baconian natural philosophy, “Their attitude might best be described as a- practical, since practical purposes are utterly irrelevant to the science of nature as such. [...] there is a certain disdain, almost contempt, for utility in the more practical sense” (Reif 1962, 56). 119 Newman 1997, 313. My emphasis. 120 Newman 1997, 313.

!51 would be meaningless: nature’s ordinary course is the object under investigation and human contrived situations are thus fundamentally misleading. They are considered “violent,” hence opposed to nature, and do not count, as Erastus argued, as revealing the true principles of things, that is, their natures.121

This was not true for humans alone, but all processes which impeded nature’s ordinary course—all “violent” motions or accidents. How do we learn that the nature of an acorn is to develop into an oak tree? We watch its development under ordinary conditions where it actualizes its potential, in other words, we look for what it does most of the time. Of course, most acorns do not end as oak trees. Whatever impedes their normal development counts as “violent” motion and thus as a disturbance. As one 17th century scholastic textbook writer summarizes, ! In the individual kinds of changes two changes are found: the one natural, the other beyond nature or violent. There is natural generation, as when roses blossom in the spring; violent, when they blossom in winter by means of human endeavor. [...] Natural deterioration [occurs] when a man becomes old in his sixtieth year; violent, when he ages more quickly because of sickness or mental anxiety. Alteration is natural, when water cools; violent, when it becomes hot. Locomotion is natural, as when earth descends; violent, when it ascends.122 !

121 Newman 1997, 314. 122 Translated in Reif 1962, 224.

!52 Experiments that twist the lion’s tail involve violent motion and thus teach us about disturbances, not about natures.123

The later scientific revolution involved a significantly different pairing: the collapse of the artificial-natural distinction, by likening nature to artifice, was paired with an instrumental goal to investigate what humans could do, rather than what happened in nature’s due course. This opened the door for manipulative projects. Experimentalism involves the assumption, as historian Garland Allen wrote, that “intervention into the workings of the organism [or nature], although artificial, still reveals something about natural or normal states of nature.”124 The assumption that intervention or disturbance can reveal what is natural conflicts with the Aristotelian position just discussed.125 It involves a fundamental shift in what counts as a “natural” state of nature. The Aristotelian would deny the assumption. The connection between justifying experimentation and collapsing the artificial-natural distinction will be considered throughout what follows.

The second question is prompted by the progressive nature of this narrative. It gives the impression that each new understanding of the art-nature relationship supplants the one that came before; that Aristotle’s distinction was supplanted by Boyle’s collapse. Through

123 See Reif (1962) for a discussion of the common type of Aristotelian Natural Philosophy taught through scholastic textbooks in the 17th century (1962 238ff). As she writes, “due to their [the textbook writers] conception of natural philosophy as a wholly speculative science whose subject is natural, not artificial, bodies, they relegated [...] practical endeavors to the various arts.” Peter Dear also points out that, more often than not, we do not learn about the acorn by actually watching anything. Nature’s are revealed through universal statements of commonly held experience—statements like, “the sun sets in the evening.” A lot more could be said here. There is a further complication as well which I have left out of this discussion and that is the role of experience itself, and its relation to understanding cause and effect as a necessary connection. See the discussion in Dear 1995, 153-5. 124 Allen 1994, 90. 125 Nancy Cartwright (see Cartwright 1999) argues that our current scientific experimentalism is actually much more similar to Aristotle’s than we give it credit for. The difference is that for Aristotle we learned about natures through unmediated, ordinary circumstances, whereas the natures of modern science reveal themselves only under very contrived circumstances. See below.

!53 this dissertation we will see that this is not the case. Each of these different ways of thinking about the art-nature relationship continue to exist in tandem in biology, sometimes creating tension. The arguments between early-20th century naturalists and experimentalists will call to mind those highlighted above between the Aristotelians and Bacon, or Erastus and

Sennert: naturalists were concerned that experimental methods did not teach one about

“undisturbed nature”; experimentalists chided them for their “reverence” of what they considered “natural or ‘normal’ phenomena.”126 Throughout the dissertation we will consider how these understandings of the relationship between natural and artificial relate to one another.127

Finally, the third question, to the extent that the ontological distinction between art and nature was undermined during this period, it was not for this reason simply ousted from scientific discourse, as the historical narrative suggests. The collapse of the ontological distinction only opened the door for the distinction to be drawn on other grounds. Historians have been particularly interested in its being drawn on aesthetic, moral and theological grounds.128 But it was also drawn on epistemic grounds: how can we learn about, as Allen put

126 MacDougal 1909, 121; Kohler 2002, 88; see Kingsland 1991. Livingston 1917; quoted in Kohler 2002, 93-4. 127 Darwin’s Origin of Species, presents examples of all three types of relationship between the artificial and the natural considered above. At times Darwin speaks of nature in the Baconian sense: nature is like a breeder that creates forms fitted to different external circumstances in the same way that humans create pigeons fitted to their (the breeders’) aesthetic sensibilities. He utilizes this notion of nature to dodge the problem that natural species are different from man’s domestic pigeons. Isn’t it clear, he asks rhetorically, that nature creates works of art much greater than man ever could? Darwin also uses nature in Boyle’s sense. He says we can look at each creature as like “any great mechanical invention”, as “the summing up of many contrivances” (Darwin 1859, 486). Maybe most surprisingly he also speaks of nature in an Aristotelian sense, if only to discredit it. This comes out when he speaks of “reversion.” The position he argues against is one that takes it that “no deductions can be drawn from domestic races to species in a state of nature” because domestic races are simply the result of disturbances of nature, and once the disturbance is alleviated, they revert back to their “natural” condition. In other words, this so-called “principle of reversion” posits an internal source of change; an internal potential to return to a natural state. Human interference is simply accidental and cannot teach one about species in the “state of nature.” 128 Daston and Park 1998, 280, 300. For example, in the case of 18th century naturalists separated “artificialia” from “naturalia”; similarly, “natural” is commonly used as a moral authority, and contrasted with the artificial.

!54 it, natural states of nature through the study of objects under artificial circumstances? The collapse of the artificial-natural distinction does not solve this stumbling block; it simply introduces another one. Artificial circumstances become only one possible state of nature— what makes us think they will stand for the others? What makes us think they are representative? This is a problem because what we study under controlled circumstances are rarely the exact phenomena of which we’re interested. What we hope is that they accurately represent the type of phenomena of which we’re interested.129 We will consider these issues in the following chapters.130 !

CONCLUSION

Myths about the origins of modern science reflect what we take to be the salient features of, and influence how we think about, contemporary science. These myths often involve the collapse of the artificial-natural distinction and the ideas that nature became passive, manipulable, and distinct from humans, while human artifice rose to the level of an

129 This obviously depends on what circumstances we’re interested in understanding—what modern commentators call the target system (to be contrasted with the experimental system). 130 One recent attempt to explicate what science learns about natural circumstances from experimental circumstances involves re-invoking Aristotle. Philosopher Nancy Cartwright argues that the best way to understand what physicists do when they perform experiments is to bring back Aristotle’s idea of natures. Physicists perform experiments that reveal the stable natures of the phenomena of which they’re interested. Unlike Aristotle those natures can no longer be revealed in the everyday, common experience of the natural world however. They are only revealed under very specific, particular and idealized circumstances that physicists create in the laboratory. Like Aristotle, however, the knowledge that is gained is of natures: the typical or ordinary behavior of the phenomena under investigation. What physicists do, according to Cartwright, is less the creation of circumstances that are representative of natural states of affairs in the laboratory, and more the understanding of natures of physical phenomena under very unnatural circumstances. These natures can be used to understand more complicated, non-experimental, “natural” circumstances because their typical behavior or nature will remain stable across large changes in the natural systems in which they occur. (See Cartwright 1999a. See Morgan 2003 for a discussion of how this approach fails when applied to biological phenomena.) Beginning with the next chapter we will start to see many examples of biologists who are worried about human disturbance, and thus continue to invoke the artificial-natural distinction. The way in which this might be considered Aristotelian is very different from how Cartwright understands physics to be Aristotelian—many biologists, as we will see, do not think that the nature of the biological phenomena can only be revealed under very contrived circumstances—quite the contrary.

!55 epistemically privileged method. The following chapters will complicate and undermine the collapse narrative. This chapter has also introduced different ways of thinking about the artificial-natural distinction and provided a reason why their discussion is not widely represented in historical and philosophical scholarship. In the end I proposed three questions which arose from this narrative and which we will consider in further detail in what follows.

!56 ! ! CHAPTER 3 ! Denaturing Nature Disturbing Conditions and Classifications ! ! ! ! ! ! ! ! Despite Darwin, we are not, in our hearts, part of the natural process. Lynn White Jr, 1967, The Historical Roots of Our Ecological Crisis !

In the previous chapter I considered a historical narrative that has done more than any other to direct our intuitions about how the artificial-natural distinction impinges on scientific discourse and practice. Because this narrative emphasizes the connection between the breakdown of the artificial-natural distinction and the origin of modern experimental science, it creates the impression that, firstly, the distinction matters little to contemporary science, and secondly, that any science invoking the distinction must be old-fashioned or premodern.131 My goal throughout the rest of this dissertation is to move beyond this narrative.

In the first half of this chapter I provide a general overview of how the artificial- natural distinction is used in the life sciences through a brief, albeit longue durée, intellectual

131 Or more like a design science. By modern science here I really mean something more like “modern natural science.” That “artificial” enters into design sciences—architecture, engineering, business administration, artificial intelligence, etc.—is well known. Simon (1981) argues at great length that such design sciences are “sciences.” The scientific revolution narrative is about modern natural sciences, and largely ignores design sciences.

!57 history spanning from the late-18th century onwards. My argument contrary to the above narrative, is that the artificial-natural distinction has been repeatedly invoked since the time of the scientific revolution when it was supposedly dissolved in all of science. This section will also act as a prelude by introducing the episodes that will occupy later chapters.132 In the second half I ask whether there are any common, general threads to be found among the historical cases. I think there are two: ! (i) We should treat the history of the artificial-natural distinction as a negotiation about the place of human activity within the world of biological study. (ii) The artificial-natural distinction is used to demarcate the contours of the discipline according to multiple axes, in terms of objects, results, methods, and even personas. ! ARTIFICIAL AND NATURAL THROUGHOUT THE HISTORY OF THE LIFE SCIENCES

The breakdown of the artificial-natural distinction that accompanied the rise of

Boyle’s “mechanical” philosophy did not happen as cleanly as the standard myth suggests.

This is especially true for the life sciences.133 A common historiographical sentiment is that the mechanization was “a 'failed' aspiration.”134 Of course, this is not true for today’s biology tout court. Especially in those areas “closest” to the physical sciences, like molecular biology, the boundary between the artificial and the natural doesn't have the same influence

132 This history will be selectively focused on a set of cases where the artificial-natural distinction arises and is contested—in other words, it should not be taken as representative of the history of the life sciences in general. 133 This was not, however, because of some general lack of initiative to collapse the distinction: many of the early mechanists and physiologists—as well as many later figures in biology and philosophy—thought that all of life could and should be understood as like a machine. Moreover, René Descartes, whom we encountered in the last chapter as having a strong influence of the Royal Society’s thinking, was directly influenced by physiologist William Harvey's experiments on the circulation of blood. 134 Ernst Mayr, the evolutionary biologist and philosopher, wrote, “When evolutionary biology was examined for its “scientificness” according to the criteria of mechanics, it flunked the test” (Mayr 1997, 28). Shapin 1996, 185; see Wilson 1995, chapter 1 regarding the historiography of this sentiment.

!58 on practice nor does it play as large a role in theory. Nonetheless, the relationship between these two antithetical terms looms behind many of the major and most stubborn issues in the history of the life sciences. Here I will introduce examples from the recent history of biology to bring out some of the ways in which the distinction has been and still is drawn and contested.135

Late-18th century debates among pre-Darwinian physiologists over the importance of experimentation are especially germane. The Laplacian-style determinists of the 18th and

19th centuries—most famously, Antoine Lavoisier and Claude Bernard—argued that artificial experimentation should be the major methodological approach to physiology because only through experimental means could life-phenomena be isolated, stabilized and calculated; life, for them, was law-governed and could be represented by predictable regularities determined through extensive experimentation.136 Physiologists in the tradition of the Montpellier school of medicine—notably Xavier Bichat—favoring a less manipulative observational physiology, could not disagree more; to isolate and stabilize biological phenomena—to obtain constant outcomes from experiments—was to fail to comprehend living phenomena altogether.137 They believed that there was something spontaneous and indeterminate about life such that one could obtain predictable outcomes only at the expense of “denaturing” the objects under investigation.138 Thus, the human-manipulated, artificial experiment, they argued, was of limited value for the study of nature.

135 As a clarification, in saying that the distinction is contested in biology, I mean to claim that the extent to which the distinction is collapsed is an issue prevalent in biology. 136 Coleman 1971; Westfall 1971, 82ff; Gigerenzer et al. 1989, 124-7; Canguilhem 2008. See Bichat 1801. 137 Williams 2003, 157-8; Reill 2008. We will see how these concerns arise for Darwin in the next chapter. 138 Albury 1977, 61; Gigerenzer et al. 1989, 124-7.

!59 Concerns about the artificial-natural boundary in the pre-Darwinian era did not arise in physiology alone. In natural history, for instance, such concerns can be found throughout the works of both Georges-Louis Leclerc, Comte de Buffon and Georges Cuvier.139 Buffon's natural history, laid out in his multi-volume Natural History (1749–1788), was a direct response to the methods of the mechanical philosophy and its definition of matter and therefore dealt directly with the artificial-natural distinction.140 “The springs that [nature] uses,” he writes, exploiting a favorite analogy of the mechanists, “are living forces,” wholly different from the dead matter of machines.141

While Buffon maintained that the mechanical philosophy impressively dealt with dead matter, living matter, to be comprehended correctly, required a different methodology entirely. In volume five of Natural History, for instance, he warns naturalists about complications posed by domestic animals, which carry the “stigma of slavery.”142 Domestic animals have been shaped by humans under “unnatural” conditions and are misleading with regard to “savage,” “wild” or “natural” animals.143 Their study is therefore of limited use for natural history. Thus it is the duty of the naturalist to “examine them with care [...] to separate artifice from Nature; and never to confound the animal with the slave, the beast of burden with the creature of God.”144

In the introduction to his famous systematic study of the structure of animals, The

Animal Kingdom (1817-1830), Georges Cuvier contrasts natural history with general physics

139 See Sloan 1995 for natural history more generally. 140 Sloan 1976. 141 Quoted in Reill 2005, 47. 142 Buffon 1749-1788, 50. 143 Ibid., 301 144 Ibid., 301-2; my emphasis.

!60 (i.e., at that time: mechanics, dynamics, chemistry, etc.), and relies on a distinction between the artificial and the natural to do so. Although natural history should attempt to apply the same methods as general physics—calculation and experimentation—Cuvier suggested that this would seldom be possible, since life-phenomena must be taken in their entirety and cannot be reduced to their elements.145 When we attempt to isolate and suppress “the numerous phenomena which compose the life of an animal,” life itself is “wholly annihilated.”146 The artificial methodology of general physics which relies on human manipulation is useless for the study of living nature—a concern similar to that expressed by the Montpellier tradition above. Within natural philosophy, that is, both natural history and general physics, each investigative strategy has its particular place: “Calculation, so to speak, commands Nature; it determines phenomena more exactly than observation can make them known: experiment forces her to unveil; while observation watches her when deviating from her normal course, and seeks to surprise her.”147 Adopting an observational, comparative methodology may be the only way to study the natural history of life without annihilating its very naturalness—that is, without annihilating the very object of study.

Later in the 19th century Charles Darwin would find himself involved in debates surrounding the boundary between the artificial and the natural. Traditionally construed, the argument of the Origin of Species (1859) is built on an analogy between natural and artificial selection: Darwin exploits the fact that breeders have had particularly good success in creating new varieties of domesticated plants and animals, through the process of artificial

145 Cuvier 1817-1830, 14. 146 Ibid. 147 Ibid.

!61 selection, to argue for the efficacy of natural selection.148 While the former depends on an anthropogenic power, the latter depends only on a natural process of competition, that is, “the struggle for existence.” Darwin even chose the phrase “natural selection,” “in order to mark its relation to man's power of selection” over domesticated animals and plants.149 Alfred

Russel Wallace, co-founder of the theory of evolution by natural selection, disagreed strongly with Darwin's use of artificial selection (see Chapter 5). From his earliest papers Wallace makes it clear that these two categories of selection are disanalogous, arguing that natural competition between species in the wild is a completely separate process from domestic breeding.150 For instance, ! A wild animal has to search, and often to labour, for every mouthful of food—to exercise sight, hearing, and smell in seeking it, and in avoiding dangers [...] There is no muscle of its body that is not called into daily and hourly activity [...] The domestic animal, on the other hand, has food provided for it, is sheltered, and often [...] carefully secured from the attacks of its natural enemies, and seldom even rears its young without human assistance.151 ! Domestication, practiced by even the best breeders, is unstable and impermanent— domesticated animals revert to their wild type when human selection is alleviated. For these reasons, Wallace saw the artificial process of domestication as unable to tell us anything

148 See chapters 4 and 5. See also Evans 1984; Provine 1986; Waters 1986; Richards 1998; Alter 2007; Burnett 2009. 149 Darwin 1859, 61. 150 Darwin and Wallace 1858, 54. 151 Ibid., 59-60.

!62 about natural processes like natural selection. The debate between Darwin and Wallace about the importance of artificial selection lives on in contemporary evolutionary theory.152

The artificial-natural distinction remains prominent in post-Darwin biology as well and the concerns expressed are quite similar. In Chapter 1 I alluded to the engineering ideal of early-20th century biologist Jacques Loeb. Loeb thought that biology should be reformulated as an engineering science, rather than a natural science; one that aimed, harkening back to Bacon and Glanvill, to produce practical knowledge first-and-foremost. If the aim was to understand the nature of life, for example, the surest way to meet that aim was to try to synthesize life in the laboratory: “we must either succeed in producing living matter artificially, or we must find the reasons why this is impossible.”153 His stance on the proper aims and practices of biology was meant to be in direct opposition to naturalists of the 19th century, such as Darwin and Wallace.154

A similar perspective can be found among many experimentalists of the early decades of the 20th century. The physiologist and ecologist Burton Livingston, for example, tactfully wrote that by 1917 the “older reverence for natural or ‘normal’ phenomena [had] largely disappeared.” And echoing Loeb’s engineering program, ! We have learned that the range of conditions offered by nature does not generally happen to be great enough to allow adequate experimental interpretation of plant processes, [...] if a student has not a liking and talent for creating physical and

152 See reviews by: Hill and Caballero 1992; Falconer 1992; Rice and Hostert 1993; Harshman and Hoffman 2000; Gregory 2008; Garland and Rose 2009; Kawecki et al. 2012. 153 Loeb 1912, 5-6. See also Keller 2002, 18. 154 This perspective was influential on a number of later biologists including the famous geneticist Hermann J. Muller as well as scientists in related disciplines such as the behaviorist B. F. Skinner.

!63 chemical conditions such as never have occurred in nature, he should not cast his lot with plant physiologists, for the next generation.155 ! Livingston’s argument should be seen more as rhetoric than accurate representation—who is this “we” that has “learned,” the experimentalist? The “older reverence” for natural phenomena still existed in many fields and physiological ecologists continued to argue about such issues for many years.156

These engineering and experimentalist conceptions of biology met with antagonism from biologists of other traditions, especially evolutionary naturalists, who considered them to be misguided. Members of Thomas Hunt Morgan’s laboratory—the famous “fly group”— recalled the “violent opposition” they encountered from those who saw the mutations they discovered in the fruit fly Drosophila melanogaster as mere pathological changes, brought about by the laboratory, and having no relevance to what goes on in nature.157 (The mutations referred to included flies with black bodies, white eyes, or no eyes at all.) Henry Fairfield

Osborn, curator at the American Museum of Natural History in New York, for example, wrote that “Speciation is a normal and continuous process,” he argued, “it governs the greater part of the origin of species [...] Mutation [what the fly group investigated] is an abnormal and irregular mode of origin, which while not infrequently occurring in nature is not essentially an adaptive process; it is, rather, a disturbance of the regular course of

155 Livingston 1917; quoted in Kohler 2002, 93-4. 156 For a history of ecology which brings out many of these issues see Kingsland 2005. 157 Quote from an interview between historian Garland Allen and the geneticist Theodosius Dobzhansky, 1966, at Rockefeller University. Dobzhansky worked in Morgan’s laboratory. 158 Emphasis in first quote mine. Osborn 1927, 40-1. The second quote has emphasis throughout in the original which I have removed.

!64 speciation.”158 The idea that mutations were disturbances, and their importance an artifact of laboratory experimentalism, was a commonly held belief among naturalists.159

The distinction played a role in many debates associated with the “modern evolutionary synthesis” of the 1930s and 40s—an influential movement which aimed at unifying a set of seemingly distant sub-disciplines of biology under the general rubric of evolution (see Chapter 6).160 Theodosius Dobzhansky, one of the primary “architects” of the modern synthesis, relied heavily on the “artificial setting” of the laboratory to derive evolutionary conclusions about populations of fruit flies (Drosophila) in the “wild.”161 He often and openly expressed his reservations however about the artificiality of such settings and continually sought to make the laboratory more natural by bringing in flies, for example, directly from the field rather than using strains which have spent long periods of time under artificial conditions.162 In a 1939 paper Dobzhansky expressed his uneasiness: ! The fact that most animal and plant species which have served as material for genetic investigations might be classed as domestic or semi-domestic has repeatedly been used to cast aspersions on the validity of the resulting data for an understanding of the evolutionary process. To some writers the word “domestication” has become a kind of scarecrow. Since there is no evidence that domestication per se either induces or prevents the appearance of any class of genetic changes, this attitude is untenable. In the last analysis domestication is merely a special case of “natural” conditions, this

159 Evolutionary biologist Ernst Mayr recalled that this comment “well expressed the feelings of the naturalists.” See his keynote address for Evolutionary Synthesis Conference, 13, 33, APS. 160 Provine 1971, 1986; Smocovitis 1996; Cain 2003, 2009; Cain and Ruse 2009. 161 Provine 1981. 162 Kohler 1994, 288. Ironically, this practice is now frowned upon because the evolution of flies not yet adapted to laboratory conditions is seen as too artificial!—most laboratory strains have thus been domesticated for a decade or more. See Harshman and Hoffman 2000, 34-5.

!65 latter term being in reality a name subsuming a great variety of diverse conditions. On the other hand, domestication does modify in some ways the balance of forces acting upon the genetic composition of a population, and hence cannot be entirely disregarded in studies concerning population dynamics. In the ecology of an animal, even so little “domesticated” as D. melanogaster, one can perceive certain special features: temporary relaxations of natural selection caused by overabundance of food and lack of enemies, extreme shrinkages and excessive increases of the population size, introduction or removal from a given locality of masses of individuals by man.163 ! As I mentioned in Chapter 1, Dobzhansky disdainfully referred to a paper by another prominent geneticist and evolutionary biologist, Alfred Sturtevant, as being based upon

“laboratory strains!.”164 Dobzhansky's concerns that model organisms, such D. melanogaster, might be purely human artifacts, non-representative of their natural or “wild” counterparts, continues to be hotly debated today.165

The application of the artificial-natural distinction extends to biologists’ personas and their evaluations of one another. The idea that biologists who spend too much time in the laboratory don’t have enough of an appreciation for “nature” to pronounce on evolution persisted throughout Dobzhansky’s lifetime. Dobzhansky himself, for example, wrote to

Ernst Mayr in 1974, “I also agree with you that [geneticist Hermann J. Muller] was ‘rather naive’ in many of his pronouncements on evolution,” but this was to be expected, “from a man of great laboratory achievements but no familiarity with organisms as they live

163 Dobzhansky 1939, 345-6. 164 Provine 1981, 52. 165 See for instance Wolff 2003, Creager et al. 2007, Ankeny and Leonelli 2011.

!66 outside.”166 In other words, to say anything substantial about the natural process of evolution, one needs some acquaintance with nature.

More recently, in ecology debates surrounding the artificial-natural distinction arise not only with regard to model organisms, but with regard to the experimental set-up itself.

Laboratory experiments in ecology are often done with microorganisms or small arthropods kept in bottles because this set-up is cheap, easy to maintain, and easy to control genetically and environmentally. Critics contend that these conditions go little way towards answering ecological questions.167 Ecologist Steven Carpenter puts the worry in the following way: “a molecular biologist who isolates ribosomes is working on ribosomes; an ecologist who isolates organisms in bottles may not be working on communities and ecosystems in any relevant sense.”168 According to Carpenter, the objects of molecular study can be and should be isolated. It makes little difference to the molecular study of ribosomes—their structure and function, for instance—whether they are considered in vitro or in vivo. On the other hand, to isolate the objects of ecological study from their natural context is, in a sense, to sacrifice those objects altogether; they are altered so fundamentally that little worthy of ecological study remains.

Ecologists like Jared Diamond argue, therefore, that for ecology the most fruitful types of experiments are natural experiments, in which biologists study the effects of large natural disturbances (e.g., storm, drought, fire, etc.), rather than the human-induced

166 Theodosius Dobzhansky to Ernst Mayr, 1974, Mather, Sierra, California; APS. It is also worth flagging that Muller was influenced by Loeb’s conception of biology in which the main aim was to control life processes and to create or engineer new biological possibilities (as we saw from the quote above). This aim was inconsistent with many evolutionary biologists who wanted to understand how nature worked undisturbed by human influence (see Pauly 1987, 177-83). 167 Diamond 1983, 1986; Carpenter 1996, 1999. 168 Carpenter 1996, 678.

!67 disturbances of laboratory and field studies (see Chapter 7).169 Natural experiments should not be confused with the study of natural history; that is, natural experiments do not belong to the observational or descriptive side of science. Rather they are often utilized to test specific hypotheses, involve significant manipulations— albeit “natural,” rather than human- induced, manipulations—and aim at determining the causal factors at play in a particular ecological situation. Most natural experiments involve a comparison of two ecological

“places” that ideally differ with regard to only one important factor, e.g., two lakes that differ with regard to mineral composition or two islands that differ with regard to a particular predatory species. A claim is then made about the causal influence—or lack thereof—of this factor. Diamond and others argue that natural experiments far outweigh other types in terms of realism (i.e., whether results can be extrapolated to natural communities), generality (i.e., the number of natural communities to which the study applies) and scope (i.e., the experimental manipulations possible—natural experiments, for instance, do not face as many ethical or environmental obstacles).

Parallel concerns arise in ecology in a number of different venues. Biologist Earl

McCoy and philosopher Kristin Shrader-Frechette have highlighted and problematized the ways in which the natural-artificial distinction arises throughout the entire discourse of ecology. “Natural places,” “natural divisions,” “natural systems,” “natural experiments,”

“natural laboratories,” “natural communities,” are all commonly employed phrases.

Furthermore, natural history and ecology both, they argue, presuppose concepts of “natural

169 Diamond 1983, 1986, 2001; see also Dunning 2008 and Diamond and Robertson 2010 for the use of natural experiments outside of ecology; and see Kohler 2002 for a history of natural experiments in ecology.

!68 place.”170 Should a distinction that is for a number of reasons conceptually incoherent play such a fundamental role in ecology, they provokingly ask?

In a similar fashion, the artificial-natural distinction underlies recent arguments surrounding the distinction between “native” and “non-native” (or “alien”) species. This distinction has acted as a guiding principle in conservation and restoration management, where alien species are seen as disruptive or unwanted and large-scale environmental policies are administered to control or remove them.171 It is argued, for instance by conservation ecologist Daniel Simberloff, that alien species threaten the natural environment.172 Critics of the native-alien distinction, however, point out that the very distinction between native and alien species is itself inconsistent, and that such claims are often not based on any evidence of the disruptive influence of alien species at all. Instead, claims are usually based on the idea that the artificial act of human-dispersal accomplishes a “denaturing” of the dispersed species

—above we saw this phrase used in a similar fashion by Xavier Bichat and the Montpellier school of medicine.173 As it is put in a recent article, Simberloff's claim “makes sense only if a human act of dispersal renders nature unnatural.”174

The issue of native versus non-native species is part of a broad category of what we might call applied issues in which the artificial-natural divide plays a role.For example, the issues emerge clearly from a set of influential articles by Thomas Henry Huxley, the evolutionist and vocal proponent of Darwin’s theory.175 When Huxley was writing these

170 See Shrader-Frechette and McCoy 1995. 171 Davis et al. 2011. 172 Simberloff 2005. 173 Chew and Hamilton 2011, 36. 174 Ibid., 36. 175 Huxley 1893, 1894.

!69 articles, questions were emerging about whether evolution justified a particular social-ethical order—a question which resonates still. Huxley argued that civilized human society was not in the “state of nature,” where the struggle for existence reigned supreme, but instead in the

“state of art.” He likened human civilization to a domestic garden: when we build a garden, he said, we put a wall around it, literally or figuratively, so that natural forces —what he called the “cosmic process”—are kept out; we might control, for instance, the influence that sunlight, temperature, or pests have on our garden. Civilization is currently in a similar state, he argued, and those concerned with ethical progress within society would do well to keep this in mind. “The thief and murderer follow from nature,” he argued, “just as much as the philanthropist.”176 A functioning society therefore depends on its members acting contrary to nature, so that the struggle for existence is alleviated within the artificial conditions of human evolution. For these reasons he professed, “Let us understand, once for all, that the ethical progress of society depends, not on imitating the cosmic (i.e., natural) process, still less in running away from it, but in combating it.”177 For Huxley, and the many that have followed him, human evolution is a process altogether different from natural evolution.178 !

DISTURBING CONDITIONS AND CLASSIFICATIONS

What should we make of these myriad examples? One thing at least should be clear.

In many areas in the history of the life sciences, significant and recurrent issues arise from the different ways in which the artificial-natural distinction is drawn and contested. This

176 Huxley 1893, 80. 177 Ibid., 83 178 See Williams 1992.

!70 distinction has been used to frame the debates even when the stakes are very different. Bichat and Bernard’s concerns about life’s supposed spontaneity are very different from Darwin and

Wallace’s dispute over the status of domestic organisms. In other words, the collapse of the artificial-natural distinction has not occurred in many areas of biology.

Without ignoring their historical specificity, is there anything more we can say about these cases? What threads run through and link them? I think there are two: ! (i) The history of the artificial-natural distinction is a negotiation about the place of human activity within the world of biological study. (ii) The artificial-natural distinction is used to demarcate the contours of the discipline according to multiple axes, in terms of objects, results, methods, and even personas. ! I will explain these below but illustrate them in greater depth throughout the succeeding chapters. !

(I) HUMANS AS DISTURBING CONDITIONS

The reason why the artificial-natural distinction arises frequently in biology may be, at one skeptical extreme, attributed simply to convention. In the distant past something significant was conveyed through its employment, but unlike then, its current prevalence can be attributed to tradition. This dissertation is meant to prove otherwise. The wide-ranging set of examples considered so far are enough to call this into question: the frequency with which the distinction is actively debated would be somewhat mysterious if it were merely

!71 conventional. I believe there are deeper reasons why this language permeates biologists’ characterizations of manifold aspects of their discipline.

It is helpful to reflect on this by thinking about the rhetorical use of the artificial- natural dichotomy. People, Loeb is one, exploit the artificial-natural language to further their own goals against those of their opponents and to evaluate each other’s work or even scientific identity. “Your fruit flies are artificial constructions!,” is a good or bad thing depending on the context. Such rhetorical uses of this distinction, however, are effective— indeed rhetoric in general is effective—only in so far as they take advantage of a tacit but potent common framework of assumptions for understanding and communicating. If there was not already an underlying framework, such uses would not be rhetorically effective. This should push us to ask, what are the features of this underlying framework? How did this framework come into being?

In Chapter 1 I hypothesized that the artificial-natural distinction is useful because it succinctly conveys a more fundamental concern about how human actions relate to nature’s normal development. This concern is a feature of the underlying framework which makes rhetorical uses of the distinction effective. The concern is that “the artificial”—that which is human-mediated—is misleading with regard to “the natural”—that which exists by (or in) nature. This is expressed many times above—think of the complaint that humans “denature” nature. Even those who do not themselves express this concern employ tactics that feed off its prevalence. Loeb exploits it to argue not that the distinction doesn’t exist, but that the artificial is more useful. The artificial-natural distinction is a useful way of sorting out what

!72 is created by humans from what exists by nature; of sorting that which is misleading from that which is useful and valid for biological study.

Of course, all natural science requires sorting artifacts from natural phenomena, where the latter are most commonly the ultimate object of study.179 In this sense, artifacts are things that are observed in a scientific investigation but that are a result of preparation, rather than naturally occurring. Examples from microscopy are perhaps the easiest to grasp. Many artifacts are introduced as a result of the preparative procedure that goes into creating a microscopic slide. These can be as simple as distortion due to the size or color of the specimen, but also include much more misleading and problematic cases, for instance, when an artifact is mistaken for a natural object. What T. H. Huxley thought were organic structures in the late 1850s, possibly even the most primitive and fundamental form of life, turned out to be nothing but preparative artifacts created by immersing the specimen in a particular solution of alcohol and sea water.180 Although the cases vary in complexity, this is a common practical scientific problem.

A distinction thus arises for all sciences between preparative artifacts and natural phenomena, so the interesting question is why do the concerns appear to be more exaggerated, fundamental, and derisive in biology. There are at least two reasons. First, in those areas of biology where this distinction arises most frequently—ecology, organismal and evolutionary biology—the systems studied are extremely complex. Since complete control over all experimental variables is in these cases impossible, and often even undesirable, the

179 But see Herbert Simon on the design or engineering sciences versus the natural sciences (Simon 1981). See Weber 2005, Chapter 9, for a general discussion of artifacts and a study case of the mesosome. See Sperber 2007 on artifacts more generally. 180 Rehbock 1975.

!73 introduction of artifacts is more common and the sorting of artifacts from natural objects is arduous, uncertain and can remain incomplete.

Second, and more provocatively, in these areas of biology the question is often not just about which objects are artifacts, so they can be ignored or removed and relegated to the category of error. The question is whether the artifacts are actually natural phenomena. One reason why this question arises is because humans themselves are often active participants in the biological systems studied. They are arguably parts of ecosystems and causes of natural selection. In such cases, debates over whether something counts as an artifact cannot be resolved simply by appealing to technical or practical factors, such as Huxley’s sea water- alcohol solution. Instead, one is forced to appeal to wider concerns about drawing the artificial-natural distinction. The way the distinction arises in biology thus betrays a more fundamental set of questions about the relation between human activities and the objects of biological study: are humans a part of natural ecosystems or are they a part of the natural evolutionary process? If they are, how are their activities similar to or different from the rest of the system studied?

Consider an example introduced above. One of the debates between Darwin and

Wallace—the focus of Chapter 5—was about whether domesticated varieties are artifacts or natural creations. At the time the answer to this question had a direct bearing on whether they were considered valid objects of study in natural history. For example, Asa Gray, a Harvard botanist and friend of Darwin’s, regretted the fact that domesticated varieties had been neglected by naturalists merely “because these races are not in a state of nature.”181 Were

181 Gray [1860] 1963, 21.

!74 domesticated varieties to be ignored in the study of species evolution or were they examples of species evolution? This disagreement between Darwin and Wallace differs from the microscopy case in that it is not just a sorting of artifacts from natural phenomena, but a negotiation about what is to count as an artifact in a more fundamental sense: how does human breeding relate to natural evolution? Are humans creating artifacts or natural phenomena? In what ways do these things differ? And what does this entail for the ways we might know about them? These are more than just technical or practical questions, but are substantive questions in biological theory.

In very general terms what is negotiated in such cases is whether and in what ways human actions should be seen as interferences in systems of biological study. Philosophers of science have a convenient and general way of characterizing such interfering factors. They call them “disturbing factors” or “disturbing conditions.”182 I think that the ways in which human-mediation is treated in the above cases suggests that it is analogous to the factors philosophers call disturbing conditions. Consider one definition from philosopher of ecology

Chris Eliot. Disturbing conditions, Eliot writes, are ! factors which interfere with the applicability of a model (or model system) to the members of the designated set of instances to which a model can be, is, or is intended to be, applied.183

182 The language of disturbing conditions comes from economics, where it still occurs regularly. John Stuart Mill argued, for instance, that economic laws were true only in the abstract: they describe what would happen in the absence of disturbing causes (Persky 1990, 187-90; Hausman 1992, 123-51). He reasoned that since economic laws depend on a definition of “mankind as solely occupied in acquiring and consuming wealth,” anything that causes mankind to deviate from this definition would count as a disturbing cause (Reutlinger et al. 2011). Ecologists often use similar language, as when they designate a particular ecosystem “disturbed.” The Oxford English Dictionary defines “disturbance” in a similar way as “Interference with the regular or due course or continuance of any action or process.” 183 Eliot 2004, 3.

!75 ! To provide an illustration, during the 1920s the mathematicians Alfred Lotka and Vito

Volterra independently proposed a model that accounts for how populations of predators and prey—mice and owls, say—interact. By building into their model sizes and growth rates of each population as well as the efficiency of both predators and prey, they could provide predictions of oscillations in the size of each population relative to the other.

Although this model does outline a general trend, it will never accurately predict what happens in natural populations of owls and mice. This is because, for example, the size of mice populations depends on more than just owls. It depends on the abundance of their own food sources, the availability of suitable habitat, the presence of other predators, disease, and so on. Relative to this model, all of these factors are disturbing conditions. The application of the model is valid—accurately describes the dynamics of the natural system—only when these factors are excluded (that is, when all disturbing conditions are excluded).

While Eliot’s definition is appropriate for this particular case, to capture the general philosophical use of this phrase we need a definition that is applicable beyond the context of scientific models.184 In addition to characterizing those “outside” factors in mathematical model-building, “disturbing conditions” are used in two other contexts. In the context of laws of nature or scientific generalizations they describe those factors that must be excluded in order for a law or generalization to apply. In the context of experimentation they describe those factors that must be excluded or controlled for in order for a scientific test to be

184 Unless of course “model” is understood very broadly to capture any representation—see below.

!76 reliable.185 What these different uses of disturbing conditions have in common is related to scientific representation. In these instances, a model, generalization, or experimental system is used to represent a part of the natural world. Disturbing conditions more generally are thus factors not included in a representation that, when present, impede its accuracy or suitability.

They impede its ability represent.

The above history suggests that human-mediation has been considered a disturbing condition in certain areas and at certain times in biology. Although I mean the phrase

“disturbing condition” in a general way, I think there is something analogous between how, say, humans are considered in the above examples and how disease is considered in the predator-prey example.186 Human disturbance is the “outside” factor which hinders the validity of an object, method, etc., to represent a portion of nature. The history of the artificial-natural distinction is thus a negotiation over whether and in what ways humans count as disturbing conditions; in what ways human activity is outside of the system of study

(as disease was outside of the Lotka-Volterra model) and in what ways this matters.

It should go without saying that it is not my contention that what is significant about biology is that biologists as a collective consider humans as disturbing conditions. My contention is that throughout the history of biology, concerns about whether humans are

185 In terms of debates over laws of nature more generally, such as whether laws require implicit ceteris paribus clauses that exclude disturbing conditions and whether such laws express empirically testable content, see Earman, Roberts and Smith 2002. In terms of experiments where disturbing conditions are factors that must be excluded or controlled for in order for a scientific experiment to be considered reliable, see Reutlinger et al. 2001. 186 I realize that this characterization is more general than some philosophers would allow, but I believe it is consistent with the diverse way in which this term is used in ecology, economics, and philosophy. See Reutlinger et al. (2011) for many examples that corroborate my claim about this diversity. Most commonly “disturbing conditions” are discussed in terms of laws of nature. The language of “disturbing conditions” (“factors” or “causes”), and the related language of “ceteris paribus clauses,” came into prominence in the nineteenth century, predominantly in economics. See Persky 1990; Hausman 1992, 124; Kaufer 1997.

!77 disturbing conditions have arisen frequently and have been at the heart of a number of notable debates. The discourse of biology is thus permeated by the artificial-natural dichotomy because this language is a useful way to express such concerns.

By way of summary, what I have been pursuing here is the common framework that gives rise to the usefulness of the artificial-natural distinction. We might say that throughout the history of biology humans themselves have been thought of as disturbing conditions and this is reflected in the commonly used language of artificial and natural. We must keep in mind, however, that what counts as “human disturbance,” “artificial,” or “natural” is constantly being redefined. As a result of this discussion we should amend our first thread to say: ! (i) The history of the artificial-natural distinction is a negotiation about whether and in what ways humans count as disturbing conditions. ! (II) THE ARTIFICIAL-NATURAL DISTINCTION AS A META-CLASSIFICATION SYSTEM

In what ways is the artificial-natural distinction deployed? To what objects is it applied? The answer to this question is quite simple, but has interesting consequences. When the distinction is drawn, biologists use the artificial-natural distinction as a classification system; in other words, they classify aspects of their work and discipline according to the artificial-natural distinction. Consider these common classifications made by biologists: ! ! !

!78 Natural Artificial

natural ecosystem artificial ecosystem

natural community artificial or synthetic community

wild organism domestic organism

natural variation artificial variation

natural mutations artificial mutations

natural perturbation artificial perturbation

natural selection artificial selection

natural experiment laboratory experiment

naturalist experimentalist ! Notice first that this classification is wide in scope. It is not on the same level as classification within science, such as classifying a specimen as a particular species. This is a meta-level classification system, applied to a variety of objects of study, processes, experimental perturbations, experiments, and even persons or types of researcher (although the latter becomes slightly more complicated). The concept of human disturbance enters in at all of these levels (for instance, the naturalist, according to common opinion, is one whose object of study is not disturbed nature).

Classification practices have been a central object of study in recent science and technology studies.187 What I borrow from this research is its general definition of classification and two general points about classification systems.188

187 This research builds principally on the work of Michel Foucault, Mary Douglas, and Nelson Goodman. See Foucault 2008 [1966]; Douglas 2002 [1966]; Goodman 1978. 188 This literature focuses on many other issues which are irrelevant for our current discussion. Many of these have to do with classification systems reflecting hierarchies of power, shaping moral and social order, and natural kinds. See Douglas and Hull 1992, Hacking 2001, chapter 6, and Epstein 2007.

!79 In one important analysis Geof Bowker and Susan Leigh Star suggest that a classification is a “set of boxes (metaphorical or literal) into which things can be put to then do some kind of work—bureaucratic or knowledge production.”189 They argue that a classification system exhibits the following abstracted or ideal properties: ! 1. There are consistent, unique classificatory principles in operation. 2. The categories are mutually exclusive. 3. The system is complete. (Provides total coverage of the world it describes.)190 ! Of course very few actual classification systems meet these goals—the classificatory principles are often not consistent or unique, and classification systems gradually change over time—but this account provides a regulative ideal nonetheless. The artificial-natural distinction meets each of these properties as well as one can expect. The classificatory principle in action relates to treating humans as disturbing conditions—in the sense that human-disturbed things are artificial. The categories are mutually exclusive and complete in any given case, albeit quite variable between cases. In other words, the artificial-natural dichotomy is being used as a classification system.

The much more interesting points have to do with common properties of all classification systems. First, classification systems are not innocuous. They have consequences. This is so for everyday things as well as scientific things, because how we classify things changes how we act towards them—this is the point of many classification systems. If something is classified as A we treat it as a member of A, if it is classified as B we

189 Bowker and Star 1999, 10. 190 Ibid.

!80 treat it as a member of B. In Chapter 1, I provided an example of this in terms of the goal of restoration ecology, commonly, to restore communities to their “natural” state. Obviously in this case deciding what is to count as natural has practical consequences.

While this case is perhaps more extreme—and, in this regard, useful for making the present point—I do not think we should overlook that a similar thing happens with regard to the artificial-natural distinction as it is used in biology more widely. For the rest of this dissertation it will be important to keep in mind that classifications have consequences. How biologists use this classification system changes how they act towards those aspects of the discipline classified.

The second point is that, “As classification systems get ever more deeply embedded,” write Bowker and Star, “they risk getting black boxed and thence made both potent and invisible.”191 With embedded, Bowker and Star mean to draw attention to the degree of their commonality in working infrastructures. By infrastructures they have in mind more than just physical structures, and mean to highlight any organizing structure, including beliefs and practices, that support a system (whether bureaucratic, scientific, or belief). The artificial- natural distinction is deeply embedded in the working infrastructures of modern biologists because they use these categories as part of their daily practice and these categories provide important organizational support. They become invisible as their use becomes unconscious: they are invoked without recourse to the principles of their application. In other words, they become invisible as their use becomes obvious. A goal of what follows is to bring the

191 Bowker and Star 1999, 325.

!81 classifications biologists make out in the open, as a way of understanding what consequences such classifications might have.

By invoking the artificial-natural distinction, biologists do more than categorize aspects of their discipline, they are also defining their very discipline. By determining what objects to study, how to do so, and who shall do so, biologists shape the content and scope of their discipline. As a classification system applied to the whole of biology, the artificial- natural distinction is constitutive of biological practice. Especially in biology, where “the natural” can act as an epistemic authority, classification according to the artificial-natural distinction has a strong influence over what is and what is not considered a valid object, site, method, etc., for study. Recall again Daily’s and Pigliucci’s choices of site and research organism, or Dobzhansky’s assessment that biologists who spend too much time in the lab cannot speak authoritatively about evolution. By classifying, biologists are demarcating the contours of their discipline: they are defining what does and does not count as properly scientific. !

CONCLUSION

This chapter was split into two parts. The first was historical and aimed to move beyond the implications of the myth narrated in the previous chapter: the artificial-natural distinction is of relevance to modern science—it has been central to the modern history of biology. In the second half of this chapter I asked whether there were any general threads to be extricated from the historical cases and have argued for two threads which will be central to succeeding chapters, again they were:

!82 ! (i) The history of the artificial-natural distinction is a negotiation about whether and in what ways humans count as disturbing conditions. (ii) The artificial-natural distinction is used to demarcate the contours of the discipline according to multiple axes, in terms of objects, results, methods, and even personas. ! While being central to negotiations about valid objects or methods of study, as the above examples attest, the artificial-natural distinction both reflects and informs our understanding of the relation between human activity and that part of the natural world biologists attempt to understand. As the distinction is defined and redefined so too is what counts as valid for biological study, as are the ways in which humans count as disturbing conditions.

!83 ! ! CHAPTER 4 ! The Art Itself is Nature Darwin, Domestic Varieties and the Mechanical Philosophy ! ! ! ! ! ! ! ! ! there is no reason art should gain the point of honor of our great and puissant mother Nature. We have so much by our inventions surcharged the beauties and riches of her works that we have altogether overchoked her; yet wherever her purity shineth, she makes our vain and frivolous enterprises wonderfully ashamed. Michel de Montaigne, 1580, Of the Cannibals !

Common to both the scientific and Darwinian revolutions were discussions impinging on the relationship between art and nature. Was art a part of nature? Could art be used as a model for nature? This intellectual congruence, however, is more than just nominal. Charles

Darwin and Asa Gray, for example, were well-aware of the 17th century debates which preceded them about the relation between art and nature through the works of such revered

English writers as William Shakespeare and Thomas Browne. Furthermore, they used their understandings of these debates to inform and express their own thinking about the relation between artificial and natural selection. ! !

!84 DOMESTIC VARIETIES AND HUMAN INTERFERENCE

In a well-known 1860 review of Darwin’s Origin of Species (1859), the Harvard botanist Asa Gray wrote the following about the art of breeding (what Darwin had termed

“artificial selection”): ! “the art itself is Nature,” since the whole art consists in allowing the most universal of all natural tendencies in organic things (inheritance) to operate uncontrolled by other and obviously incidental tendencies. No new power, no artificial force, is brought into play.192 ! Perhaps a little enigmatic, this quotation succinctly captures and defends a major theme in

Darwin’s work. In the first four chapters of the Origin Darwin developed an extended analogy between artificial and natural selection. Likening nature to a breeder, he compared the creation of domestic varieties through artificial selection to the creation of natural races in the “state of nature.” Just as human selective breeding produces new domestic varieties with desirable characteristics, he argued, selection in nature, a result of the “struggle for existence,” produces new varieties and even species, filling up cracks in the economy of nature. In a letter written prior to the publication of the Origin, he explained to Gray that

Nature is an “unerring” breeder, “which selects exclusively for the good of each organic being” (Darwin to Gray, Sept 5, 1857).193

192 Gray [1860] 1963, 27; Darwin 1859. 193 There is a voluminous literature about Darwin’s analogy, much of which is tangential. See Young 1971; Ruse 1975; Evans 1984; Waters 1986; Bartley 1992; Richards 1998; Gayon 1998; Theunissen 2012. Unless otherwise noted, Darwin letters were retrieved from the Darwin Correspondence Database: https:// www.darwinproject.ac.uk.

!85 In the eyes of his contemporaries, however, the value of Darwin’s comparison hinged on whether the creation of domestic varieties was merely a result of the artificial conditions in which they were raised and bred. A common sentiment in circulation prior to the publication of the Origin held that breeding was too unnatural to represent a natural process.

This sentiment was often used to buttress arguments against drawing an analogy between domestic varieties and wild species. Charles Lyell wrote in 1832 that domesticated varieties were “extreme cases brought about by human interference, and not [...] phenomena which indicate a capability of indefinite modification in the natural world.”194 The mathematician- geologist William Hopkins asserted in a review of the Origin, “we have no right whatever to assume that nature will necessarily produce such effects at all when left to her own unobstructed operations, as those which she produces under man's interference”; he maintained that we “commit an error” when we assert that under ordinary conditions nature is capable of producing modifications similar to those observed in domestic varieties.195 And

Gray’s colleague at Harvard, Louis Agassiz, wrote in his copy of the Origin that Darwin’s mistake “has been to study the origin of species among domesticated animals” rather than wild ones.196 Darwin disagreed with these assessments, responding to one commentator, “it is an error to speak of man ‘tampering with nature’.”197 In the opening quotation, Gray defends

194 Lyell 1832. 195 Hopkins 1860, 75. 196 Agassiz wrote in his copy—sent to him to review—of the first edition of Darwin’s Origin of Species, “The mistake of Darwin has been to study the origin of species among domesticated animals exclusively instead of wild ones; his results concerning species are founded not on an investigation of species but on an investigation of breeds” (Agassiz papers, Ernst Mayr Library, Harvard University). Agassiz repeated his opinion in a review: “this process of raising breeds by the selection of favorable subjects, is in no way similar to that which regulates specific differences. Nothing is more remote from the truth than the attempted parallelism between the breeds of domesticated animals and the species of wild ones.” Agassiz 1860, 147. 197 These comments were made by the French anatomist Georges Pouchet. See Darwin 1868, 2.

!86 Darwin: artificial selection should not be seen as an unnatural interference in nature’s otherwise normal development. Instead, he responds, “the art itself [that is, of breeding] is

Nature,” “no artificial force [...] is brought into play.”

Understood in this light, Gray’s comment should be recognized as part of a broader theme running through the Darwinian revolution—a theme referred to by historian John

Cornell as, “Darwin’s reinterpretation of the meaning of art and nature.”198 Darwin said, or implied, something significant—albeit highly contested—about the art-nature relationship which was not lost on his peers. This theme, we might recall, is central to another very well- known revolutionary period: the reconfiguration of 17th century natural philosophy known as the scientific revolution. Floris Cohen, author of the most comprehensive historiographical inquiry of the scientific revolution, writes, “Every interested thinker in the 17th century had to grapple with the distinction between ‘natural’ and ‘artificially produced’ nature and the root issue of how one was related to the other.”199 Lorraine Daston provides my favorite articulation of this theme: “It is a platitude among historians of the Scientific Revolution that the seminal thinkers of the seventeenth century, most notably Bacon and Descartes, abolished the nature/art distinction by subsuming the artificial under the natural. However, it would be just as, if not more, accurate to claim that the distinction was collapsed [...] by subsuming the natural under the artificial.”200 Regardless of the direction of subsumption, scholars of the early modern period generally agree that the relationship between these two categories was transformed in the 17th century, as we have seen above.

198 Cornell 1984, 308. See also, Burnett 2009. 199 Cohen 1994, 188. 200 Daston 1995, 41-2.

!87 If discussions impinging on the relationship between nature and art played a pivotal role in both the scientific and Darwinian revolutions, what similarities do they share and what can we learn from them? It is my contention that this intellectual congruence is more than just nominal. I will argue that Darwin and Gray were aware of 17th century debates about art and nature, and furthermore, used their understandings of these debates to inform and express their own thinking about the relation between artificial and natural selection. Let me begin with a typical 17th century discussion of art and nature. After which I shall return to the Darwinian revolution. !

NATURE AND ARTIFICE IN THE SEVENTEENTH CENTURY

Early modern discussions of the relationship between art and nature found expression in a variety of contexts.201 Here I will again take as an exemplar the mechanical philosophy of Robert Boyle, but I will elaborate beyond the discussion in Chapter 2, in a way that introduces the terms of 17th century debates about art and nature that are relevant for the present context.

Robert Boyle likened nature to an intricate work of art, or as he characteristically put it, an “engine.” In his view, clocks provided a commonplace and appropriate model for the natural world (Figure 4.1). !

201 See, Daston 1998. and Bensaude-Vincent and Newman 2007.

!88 Figure 4.1: An Elaborate Astronomical Clock in Strasbourg (c. 1875). An example of the type of artifice likened to nature by Robert Boyle. (© Wellcome Library, London) ! In likening nature to an clock, Boyle sought to distance himself from what he called the “vulgarly received notion of nature,” a confused and confusing notion, fabricated by the ancients, and passed down through the Medieval period and Renaissance.202 The vulgar notion of nature was problematic for a number of reasons, notably, it encouraged what Boyle deemed unintelligible anthropomorphisms that appropriated praise rightfully belonging to

God. A typical example was the oft-cited phrase, “nature abhors a vacuum.” Nature, Boyle

202 Boyle [1686] 1999.

!89 thought, could not abhor anything, nor could it serve as an active arbitrator in philosophical debates.

Although his primary motivation was theological, likening nature to artifice was consistent with Boyle’s natural philosophy as it helped justify experimentation as a legitimate form of knowledge production.203 According to the “vulgar,” Aristotelian-inspired philosophy

Boyle denounced, the proper object of study was nature’s due course. Natural processes, it was purported, were goal-directed and the aim of natural philosophy was to understand these goals. The “nature” of an acorn, for example, could be found by watching an acorn develop into an oak tree under typical, everyday conditions.204

As a consequence, and as we saw in Chapter 2, this “vulgar” philosophy advocated a distinction between artifice and nature—a point of view so entrenched throughout the renaissance that historians have referred to it as a “habit of the understanding” or “conceptual reflex.”205 Art was understood as an intervention upon or frustrating of nature’s ends. It was able to imitate or overcome nature, but it could not reveal nature, since through art, nature’s ends were subverted by human ends.206 The object of Aristotelian natural philosophy could not be apprehended through the contrived circumstances characteristic of experimentation.

In contrast, for Boyle nature was artificial because it was God’s work of art. This view of nature as a passive work of artifice cohered well with the rise of experimentalism and

203 Boyle thought that the vulgarly received notion was blasphemous as it often portrayed nature as an artisan who oversaw the natural world. This imbued nature with an autonomy that stole praise away from God—God needed no intermediary (see Boyle [1686] 1999; see also Daston 1995). To Boyle, nature was simply a machine and God the engineer that set the clockwork system in motion. 204 See, Dear 1995. 205 Park and Daston 2006, 265 and 276. See also, Tayler 1966 and Close 1969. 206 See, Dear 1990.

!90 with the rising belief that nature is regular, uniform, and can be described according to laws.

By likening nature to artifice in this way, Boyle provided a foundation for experimentation.207 Once nature was understood as machine-like, the usefulness of experiments—and in general methods that involved manipulating nature’s mechanisms— became obvious. Just as a clock is the product of a designer, so nature was seen as a great engineering feat created and set in motion by the hand of God (Figure 4.2). If nature was like a clock, one should approach it in the same way as a clockmaker approaches a foreign watch: determine the laws that govern the regular and uniform movement of its variously sized and shaped parts by taking it apart and fiddling with its pieces. Lambasting the vulgar practice of passive observation, Boyle wrote, “He must be a very dull Enquirer, who, demanding an

Account of the Phaenomena of a Watch, shall rest satisfied with being told, that ‘tis an

Engine made by a Watch-Maker; though nothing be thereby declar’d of the Structure and Co- aptation of the Spring, Wheels, Ballance, and other Parts of the Engine; and the manner, how they act on one another.”208 !

207 Shapin 1996, 97-8. 208 Boyle [1686] 1999, 558

!91 Figure 4.2: Clockmaker About to Fix a Clock. Published in The universal magazine (1748). (© Wellcome Library, London) ! The clock metaphor runs deep since it could be used to justify experimental- manipulative practices and the uses of instruments which aided the unmediated senses or created unnatural conditions. Joseph Glanvill, recall, wrote that anatomy was useful because it tended “mightily to the eviscerating of nature, and disclosure of the springs of its motion.”209 Given these colorful metaphors, it is unsurprising that Boyle wrote, “Proper comparisons do the imagination almost as much service as microscopes do the eye.”210 As

209 My emphasis. Glanvill 1668 [1958]. 210 Quoted in Mayr 1986, 82.

!92 the distinction between the artificial and the natural broke down, the methods appropriate for the study of machines and the mechanical arts, and the rigor these philosophers saw accompanying these methods, could be applied to the study of nature.

Likenings of nature to artifice throughout the early modern period were certainly not unique to Boyle and the Royal Society.211 An influx of artifacts, like clocks and automata, and artifactual techniques, like plant grafting, in practical, philosophical and leisurely affairs, sparked debates about how artifice related to nature which permeated the wider culture, and found expression in the works of playwrights and scholars, such as William Shakespeare and

Thomas Browne, both of whom we will encounter below (Figure 4.3).212 The conceptual coherence of the art-nature distinction was widely challenged on both practical and theoretical grounds in the 17th century, and was discussed in a variety of contexts. As we saw in Chapter 2, modern commentators often present the collapse of the art-nature distinction during this period as a prerequisite or ingredient for “modern science” because it helped underwrite experimentation as a legitimate source of natural knowledge.213 As we’ve seen, the art-nature relationship was far from resolved and returned for further disputation in the context of early evolutionary biology. !

211 See for the early modern Italian context: Grafton 2007. See also Daston and Park 1998. 212 See Mayr 1986 and Riskin 2007. 213 See Shapin 1996. For a dissenting view see Newman 2004 and 2006.

!93 Figure 4.3: A Mechanical Toy. Hydraulic power drives a musical box and works a skeleton puppet. From Mechanica hydraulico-pneumatica, by Gaspar Schott (1657). (© Wellcome Library, London) ! THE ART ITSELF IS NATURE

Just as Boyle’s clock metaphor was a product of the technological innovation and fascinations of its time—a fact Boyle exploited—so too was Darwin’s use of breeding.214 As urban populations increased dramatically throughout the Victorian era, agriculture, horticulture, animal husbandry, and so on, developed as never before (Figure 4.4).

214 See Young 1985.

!94 Periodicals such as the Gardener’s Chronicle disseminated information about these practices widely.215 Darwin’s father-in-law (and uncle) Josiah Wedgwood was himself a leading sheep breeder as well as a successful businessman.216 It is thus not surprising that Darwin noted in the margin of William Youatt’s famous volume on cattle, “As this simple principle [of selection] only lately discovered even in most reliable practice, no wonder not discovered as theory of species.” Darwin was perhaps aware that technological or practical innovation—in this case the art of breeding—often predates the discovery of a theory about the natural world.217

Figure 4.4: Victorian’s in Domesticated Nature, “Botanising.” From New illustration of the sexual system of Carolus von Linnaeus, by Robert John Thornton (1807) (© Wellcome Library, London)

215 Ruse, M. 1979. Secord 1985. See also Browne 1995. 216 Ruse1979, 178. 217 March, 1840. Quoted in Cornell 1984, 325.

!95 ! Although Darwin’s likening of Nature to a breeder may have been timely, there were notable obstacles to overcome. First there was a social obstacle: although breeding occupied a critical place in the wider cultural context, naturalists tended to distrust breeders and their art (and vice versa), creating—if not full-on animosity—then at least mutual ignorance through a divergence of interests.218 My goal in this chapter, however, is to address obstacles of a more intellectual nature, as exemplified by Lyell’s and Hopkins’ remarks about the

Origin: should the art of breeding be seen as an unnatural interference? how did this impinge on debates about the proper object of study in natural history?

In asking these questions, my discussion parts ways with previous histories of the relation between artificial and natural selection. Such histories have focused on, for example, the use of analogies in science, the use of scientific rhetoric, the genesis of Darwin’s own thinking, Darwin’s debt to the philosophy of science of his day, and whether such an analogy was or is justified.219 I’m interested in a different issue that Darwin had to worry about, namely, how to justify the study of domesticated varieties as a way to learn about natural species, when the former (and the practices giving rise to them) were traditionally looked at as being outside the “state of nature” and thus improper objects of study. In their efforts to deal with this issue, Darwin and Gray gained insight from the 17th century literature they were reading. This worry was especially germane because Darwin’s first-hand experience

218 Secord 1985, 522. But see, Ritvo, H. 1987, on “aristocratic” breeding as an exception to the trend Secord outlines. 219 See the citations above.

!96 with the natural species he wrote about diminished significantly following his return from the

Beagle voyage in 1837.

Allow me then to return to Asa Gray’s review of the Origin. His comment, “the art itself is Nature,” is taken from Shakespeare’s 1623 play, The Winter’s Tale. It should not be surprising that Gray quotes Shakespeare, since early modern writers like Shakespeare,

Michel de Montaigne, John Milton and Thomas Browne were widely read and beloved in the

Victorian era, often even more so than in their own day.220 In The Winter’s Tale, Shakespeare contrasts two perspectives on the relation between art and nature through a dialogue between the King Polixenes and the shepherdess Perdita. The dialogue represents early modern debates and gives a sense of the broader intellectual atmosphere in which men like Boyle were working. Perdita says she has no “streaked gillyvors [carnations]” in her garden, referring to them disdainfully as “Nature’s bastards,” since they are a product of both art

(techniques of grafting and breeding) and of nature, rather than “great creating Nature” alone.221 King Polixenes objects that manipulated nature should not be treated with scorn and points out that the art of producing hybrid flowers is itself a part of nature and therefore worthy of appreciation.222

Shakespeare’s intention in this section was to signal to his audience that the already well-worn topic of the relations between art and nature could speak to the broader social issues discussed in the play.223 Similarly, Gray’s intention in quoting Shakespeare is to signal

220 Beer 1983 and 1985. 221 Shakespeare [1623] 1998, 68. 222 For a 17th century discussion of the horticultural arts see Austen 1653. See pp. 28-9 for grafting. For a broad discussion of how art and nature were compared in 17th century literature on gardening, see Tayler 1966, 16ff. 223 See the Introduction by Frank Kermode, in Shakespeare The Winter’s Tale. For an extended discussion, see Tayler 1966, 121ff.

!97 to his audience that the playwright’s well-known dialogue is relevant to his scientific discussion of the relation between domestication and natural history. Gray sides Darwin with

King Polixenes: artificial selection is itself a part of nature. Domesticated organisms are no less worthy of study because they are the products of a human art; in fact, they are particularly valuable for this very reason. Gray laments that domesticated races have “been generally neglected by naturalists, because these races are not in a state of nature; whereas they deserve particular attention on this account, as experiments [...] ready to our hand.”224 It is worth noting that, as with Boyle, Gray’s endeavor to dissolve the art-nature distinction also ends up underwriting an experimentalist agenda.

Seventeenth century literature was in the immediate background of Darwin’s own thinking when he first likened the art of breeding to nature. Many Darwin scholars locate his earliest use of the analogy between artificial and natural selection to November 1838.225 It is during this period that Darwin starts to employ this familiar analogy in his notebooks; in early December he records, “if nature had had the picking she would make such a variety far more easily than man” and “It is a beautiful part of my theory, that domesticated races of organics [sic] are made by precisely the same means as species.”226 It is suggestive that a few months earlier, in late August, Darwin had been rereading the 1643 Religio Medici, by

Thomas Browne.227 Browne was a physician and natural philosopher, influenced by Bacon

224 Gray [1860] 1963, 21. 225 See Hodge 2009. 226 Notebook E, 63 [after Dec 4th] and Notebook E, 71 [after December 16th]. Unless otherwise noted, Darwin’s notebooks were retrieved from Darwin Notebooks Online: http://darwin-online.org.uk/ reproductions.html. 227 For the Darwin enthusiasts: this reading occurred after Darwin’s reading of the Sebright and Wilkinson breeding pamphlets in early 1838, just following his reading of Brewster’s review of Comte in early August, just before he starts to refer to domesticated organisms as adapted—rather than ill-adapted—, and before his reading of Malthus in late September.

!98 and admired by Boyle. Darwin’s notes on Browne’s text—crammed in a transmutation notebook between thoughts on varieties versus species, natural arrangements of animal groups, and domestication—make it clear what sections interested him most.228 In these sections Browne dealt directly with the relation between art and nature in a characteristically

17th century way.229 He writes, ! now nature is not at variance with art, nor art with nature, they being both the servants of his providence: art is the perfection of nature [...] In brief, all things are artificial; for nature is the art of God.230 ! Like King Polixenes, Browne suggests that the categories of art and nature should not be opposed in the classic, Aristotelian way. He describes nature, similarly to Boyle, as being composed of “narrow engines” and “contrived parts” with God as its “excellent Artist.” His conclusion is unequivocal: “all things are artificial.”231 Browne sought to subsume the natural under the artificial, undermining a clear distinction between the two by rendering Nature

God’s work of art. Darwin records the last line in his notebook, “p. 23 for Nature is the art of

God.”232

Browne’s discussion is instructive for other reasons as well. The passages Darwin read consider also the beauty of God’s design. Browne wrote, “I hold there is a general

228 Kohn 1987, 350. 229 Daston 1995, 40. Daston and Park 2006, 300; Preston 2005, 34. 230 Browne, T. [1643] 1862, 35. 231 Browne [1643] 1862, 30-5. 232 Notebook D, 54e; Kohn “Notebook D,” p. 350. There is a discrepancy between the online and print transcription of Darwin’s notebooks. Online the Browne passage is transcribed as “for Nature is the act of God,” which is incorrect and fails to capture the meaning of Browne’s discussion and why Darwin would care about it. The print version of Darwin’s notebooks is transcribed correctly and is thus preferable.

!99 beauty in the works of God, and therefore no deformity in any kind or species of creature whatsoever.”233 This included creatures such as toads, bears, or elephants, but also those labeled monstrosities, “wherein notwithstanding there is a kind of beauty, nature so ingeniously contriving the irregular parts, that they become sometimes more remarkable than the principal fabric.”234 In the writings of Darwin’s contemporaries domesticated varieties were derogatorily referred to as monstrosities, and often neglected by naturalists for that reason.235 Lyell, for instance, objected that species hybrids could not be used as examples to corroborate theories of transmutation in nature since a significant “number of such monsters” could only be “obtained by art.”236

Darwin’s notes demonstrate that through the first half of 1838 he would have largely agreed with Lyell’s assessment. Until at least July 1838 he frequently contrasted the formation of wild “natural” species with the making of domesticated varieties or

“monstrosities” by humans.237 His famous analogy—a comparison of wild, natural species formation and the creation of domestic varieties—was thus a complete reversal of his earlier thoughts on the matter.238 Darwin’s reading of Browne in August 1838 thus fell at a time when his thinking about the relation between the art of selective breeding and wild species

233 Browne [1643] 1862, 34-5. 234 Ibid., 35. 235 See Secord 1985. Again, see Ritvo 1987 on aristocratic breeding. As Ritvo shows, not all domesticated species were judged in such a way. My interest is not in how domesticated varieties were viewed tout court, but how they were viewed by the naturalists with which Darwin would have engaged and held in high esteem. In this regard, Darwin and Gray both lament the lack of “scientific” treatment of domesticated varieties. This argument is consistent with Ritvo’s claim that animal-human relations changed throughout the early-18th and late-19th centuries such that animals became significant primarily as objects of human manipulation. See Ritvo 1987, 2-3. 236 Lyell 1932, 51. 237 See Hodge 2009, 66. For further discussions of Darwin’s changing views during this time, See Hodge and Kohn 1985, 197-200; Ospovat 1981. 238 I’d like to thank Jon Hodge for pointing this out to me.

!100 formation was fruitfully transforming from contrast to comparison. It should thus not be overlooked that Browne’s discussion put considerable tension on the coherence of the label

“monstrosity” itself, and again like King Polixenes, recommended that manipulated nature not be treated with scorn. In terms of Darwin’s changing point of view, such a discussion was fuel for the fire. After August 1838, we should note, Darwin’s entries on artificial selection become considerably more frequent.239

Perhaps we should also keep this literature in mind when considering Darwin’s later reflections on monstrosity and abnormality, as when he warns that “We should [...] be cautious in deciding what deviations ought to be called monstrous.”240 After all, “the great plume of feathers on the head of the Polish cock has been thus designated, though plumes are common with many kinds of birds” (Figure 4.5).241 !

239 See Schweber 1977, 257. 240 Darwin, 1868: 2, 413. 241 Ibid.

!101 Figure 4.5: The Polish Pigeon or Polish Cock. From Darwin’s 1868 The Variation of Animals and Plants under Domestication. Often the Polish Pigeon was referred to as a monstrosity because of the great plume of feathers on its head. Darwin was wary of such attributions. (© Wellcome Library, London) ! 17th century discussions of art and nature permeated Darwin’s intellectual milieu when he was thinking about the relation between domestication and nature. When Darwin described the history of domestication as “an experiment on a gigantic scale” and protested that “It is an error to speak of man ‘tampering with nature’,” it is hard to imagine that he

!102 wasn’t aware of the similarities between his own reliance on the art of breeding and well- known 17th century debates about art and nature.242 Similarly, when Darwin wrote in conclusion to his 1842 Essay, “We must look at every complicated mechanism and instinct, as the summary of a long history, of useful contrivances, much like a work of art,” and then in 1844 changed “work of art” to “great mechanical invention,” we likewise see his continuity with the 17th century as we find him modifying an early modern turn-of-phrase.

His use of manipulated nature to produce knowledge applicable to unmanipulated nature brought art under the purview of natural history, a bold move which could gain legitimacy from comparisons with the works of Shakespeare, Browne and Boyle that undermined the distinction between art and nature.243

Darwin’s later, more systematic writings also undermined that distinction. Brief but profound examples can be found throughout his discussions of unconscious selection and sexual selection, other types of selection that have traditionally received less scholarly attention. In the next chapter we will consider these cases in a different context. !

242 Darwin 1868:1, 2-3. 243 There are a number of complexities which, although slightly tangential, are worth mentioning. First, there is a large literature about what Darwin was using studies of domesticated varieties for prior to the publication of the origin. Was he looking at them as examples of natural selection? Was he looking more specifically for laws of inheritance? See Evans 1984 and Bartley 1992. For the philosophical context of experiments when Darwin was performing them see, Schweber 1977. In the text I have left this unspecified on purpose, as my concerns are more general than this. Second, one of the problems with the artificial-natural selection analogy was that natural species were infertile with one another, whereas domestic varieties, no matter how extreme their trait differences, continued to interbreed. There is another literature about the extent to which Darwin saw this as a problem, and following from this, whether he employed the analogy solely didactically. See, e.g., Kottler 1985. I believe, as hinted at in the below text, there are good independent reasons to think Darwin saw artificial selection as a type of natural selection, especially later in his life. I believe he possibly held this belief as early as November 1838, when he started to see domestic varieties as adapted to their particular circumstances (See Hodge and Kohn, 1985, 198). Even in the Origin, he didn’t refer to the category of “artificial selection” until page 109—prior to that referring to “selection by man” which was a category including a range of types of selection from unconscious to methodical.

!103 ! !

NATURE’S EXPERIMENTS

Needless to say, Darwin’s and Gray’s primary interlocutors were not 17th century natural philosophers, but their own contemporaries. Darwin’s reliance on breeding located him amidst 19th century debates over the proper place of experiment in generating knowledge, and sets him at the crossroads between two strongly supported traditions. The first tradition was more closely aligned with natural history and dismissed the value of human manipulation in the study of nature. This tradition had a rich lineage, itself tracing back to the 17th century. Georges Cuvier was a notable endorser; his work was tremendously influential in early-19th century England and pervaded Darwin’s intellectual environment.244

Late in life Darwin referred to Cuvier as one of his “two gods” (the other being Linnaeus)— an expression of his general admiration for the French naturalist.245

Cuvier’s authoritative discussion of domestication in the Discours Preliminaire

(1812; translated 1813) argued that variation under domestication is superficial, a result of humans tampering with nature’s order and nothing more.246 “Nature is [...] careful to prevent the alteration of species that could result from their mixture,” he writes, “It requires all kinds of ruse and human constraint to achieve these unions.”247 Through domestication, “man develops all the variations of which the type of each species is susceptible, and draws out

244 Ospovat 1981. 245 Darwin to William Ogle, 12 Feb 1882. 246 The English translation of the Discours Preliminaire was commissioned by none other than Robert Jameson, Darwin’s dull-mannered geology professor at Edinburgh; Rudwick 1997, xi. Outram 1984, 141ff. 247 Cuvier 1812, translated in Rudwick 1997, 227.

!104 forms that the species, left to themselves, would never have produced.”248 The degree of variation under domestication is unnatural and proportional to its cause, which he referred to as “slavery.”249 Cuvier’s view influenced Charles Lyell’s thought, as should be evident in the above quotation from the Principles of Geology, where Lyell dismissed the epistemological value of domesticated varieties.

Cuvier’s view of domestication was consistent with the suspicious outlook towards experimentation and manipulated nature he inherited from French physiologists like Xavier

Bichat.250 In Le Règne Animal ([1817] 1840), Cuvier explained that there were three general scientific methods: ! Calculation, so to speak, commands Nature; it determines phenomena more exactly than observation can make them known: experiment forces her to unveil; while observation watches her when deviating from her normal course, and seeks to surprise her.251 ! He concluded, however, that because of the holistic, complex nature of living phenomena,

“Natural History will long remain [...] one of pure observation.”252 The most effectual mode of observation in natural history, he argued, was non-manipulative and comparative. These represent “kinds of experiments ready prepared by Nature, who adds to or subtracts from

248 Ibid. 249 Ibid. The 18th-century French naturalist and polymath, Georges-Louis Leclerc Comte de Buffon, similarly referred to domesticated animals as carrying the “stigma of slavery” in his Histoire naturelle. It is the duty of the naturalist, he wrote, “to separate artifice from Nature; and never confound the animal with the slave, the beast of burden with the creature of God.” See Buffon 1780, 301-2. Also see, Roger 1997, 303. 250 Albury 1977; Outram 1986. Bichat himself was in this regard influenced by earlier physiologists of the Montpellier school who were reacting to the type of mechanical philosophy advanced by natural philosophers like Boyle and Browne. See, Williams 2003. 251 Cuvier, G. [1817] 1840, 14. 252 Ibid.

!105 each of them different parts, just as we might wish to do in our laboratories.”253 By successively studying the same bodies in “the different positions in which Nature places them” we are able to recognize constant relations that indicate underlying causal relationships.254 For example, Cuvier found that the relative size of the liver was inversely proportional to the relative size of the respiratory apparatus across many animal species.

Insects lacked a liver and underwent cutaneous respiration over the entire outer surface of their bodies. Mammals, on the other hand, had a liver and breathed only through a single bodily system, the lungs. He concluded from this comparison, or “natural” experiment, that in mammals the liver assists the action of the lungs.255

The Cuvier outlook permeated Darwin’s early life, especially the time he spent studying medicine in Edinburgh (1825-1827). Robert Grant, a teacher at Robert Knox’s famous extramural school of medicine and later Professor of Comparative Anatomy at

University College London, allowed the young Darwin to assist him collecting specimens near Edinburgh.256 Grant, like Knox, was familiar with Cuvier’s work and even spent time in

France studying under Cuvier himself. Given that Darwin and Grant were close and conversed over many topics while collecting, it would be strange if Cuvier’s work did not arise for discussion.

At this time in Edinburgh “vitalistic” theories were also a commonplace, and while the content of these theories is perhaps less relevant to Darwin, the methodological

253 Ibid. 254 Ibid., 15. 255 Example from Outram 1984, 331. 256 Browne 1995.

!106 prescriptions they espoused often had direct ties to Cuvier’s work.257 Darwin’s readings in

Edinburgh, for instance, included works by John Fleming and John Abernathy, both of whom sympathized with Cuvier and espoused a form of vitalism.258 In his Philosophy of Zoology

(1822), for instance, Fleming cites Cuvier specifically, referring to Le Règne Animal as “the most valuable of modern synthetic arrangements.”259 Methodologically speaking, Fleming’s work paralleled Cuvier’s in many ways. Particularly relevant for the types of allegations lobbed against Darwin’s work, Fleming referred to domestic breeds as “monstrosities” and discussed the “deranging influence of domestication” on animals species.260 Darwin was thus presented with Cuvier’s views on domesticated varieties through works such as these.

Standing at the crossroads and looking in the opposite direction, Darwin would have encountered another tradition, more closely aligned with the philosophy of the physical sciences and one that had less scruples with experimentation. For example, in early August

1838, shortly before his reading of Thomas Browne, Darwin closely studied a review of

Auguste Comte’s Cours de philosophie positive—so closely he developed an intense headache.261 The review was an overview of Comte’s philosophy in general, but particularly relevant was an outline of the role of experiment which stressed the usefulness of studying phenomena under artificial or unnatural conditions. Through this powerful instrument of investigation, the reviewer commented, “we observe bodies out of their natural state; by

257 For a summary of vitalism in Edinburgh in the early 19th-century, see Jacyna 1983. 258 Browne 1995, 85. Darwin re-read Flemings book on December 15th, 1840, when he wrote “—well read— references at end.—,” and potentially also between June 1st, 1838 and December 1839. 259 Fleming 1822, xiv. 260 Ibid., 36. 261 See Schweber 1977, 241. Brewster 1838. The review is published anonymously. On the implications of this reading, see: Schweber 1977 and Schweber 1978.

!107 placing them in artificial aspects and conditions contrived for the purpose of exhibiting to us, under the most favourable circumstances, their phenomena and their properties.”262 As shown by historian Silvan Schweber, such quotations bear a close resemblance to the text of

Darwin’s earliest sketches of his theory.263 Darwin’s reading of 17th century literature which muddled the distinction between “the artificial” and “the natural” was carried out alongside this literature which exalted the study of unnatural nature. The 17th century debates would have thus appeared quite germane, even timely, given the philosophical context in which

Darwin was situated.

While certainly opposed to dismissals of the experimental value of domesticated varieties, Darwin was nonetheless in a position to appreciate the concerns of Cuvier’s following. Despite his reliance on domestication as a gigantic experiment, Darwin did at times express his own reservations towards experimentation. For instance, in the early 1860s he was preoccupied with experiments on the fertilization of orchids by insects.264 One experiment had him artificially remove the nectaries of flowers and observe whether insects still visited the flowers. After performing these experiments he wrote to the English Botanist

George Bentham: ! I have for years been attending to insect fertilisation of Orchids, & I shd. very much from several curious reasons like to see what effect no nectary will have produced on

262 Brewster 1838. 263 He writes, likely in 1842, “The most favourable circumstances for variation seems to be propagation for many generations in domesticity [...] the influence of domesticity seems to resolve itself into conditions different from those under which nature placed the organisms.” See Schweber, 1977, 256. Vorzimmer 1975. This early “Pencil Sketch” of Darwin’s theory was likely written in 1842, not 1839. The “<>” indicate corrections Darwin made at the time of writing. 264 See Darwin 1862.

!108 the visits of insects. I once tried cutting off the nectaries; but nature's cutting off would be much better.265 ! Bentham had recently discovered a variety of Orchis pyramidalis that was naturally lacking a nectary. In this letter Darwin was asking for a few samples so that he may repeat his experiment with them. But, we might inquire, why would Darwin consider “nature’s cutting off” to be preferable to his own experimental intervention? Here I think we can see his attempting to preempt objections to the use of experiments in natural history. The flowers

Bentham discovered were a perfect opportunity for a “natural” experiment in accordance with Cuvier’s prescriptive reflections on method, indicating that Darwin took such prescriptions to heart. !

CONCLUSION ! Up to the age of thirty, or beyond it, poetry of many kinds, such as the works of [John] Milton, [Thomas] Gray, [Lord] Byron, [William] Wordsworth, [Samuel Taylor] Coleridge, and [Percy Bysshe or Mary] Shelley, gave me great pleasure, and even as a schoolboy I took intense delight in Shakespeare, especially in the historical plays.266 Charles Darwin, 1876, Autobiography ! As these autobiographical reflections reveal, Darwin, like many 19th century intellectuals, was well-versed in 17th and 18th century literature and its 19th century heirs—his reading notebooks are full of references to this body of work. Much of this literature was ripe with discussions of art and nature, as was, it’s worth mentioning, a 19th century disagreement

265 Darwin to Bentham, June 17th, [1861]. 266 Barlow 1958, 138.

!109 between Coleridge and Wordsworth over the proper interpretation of art and nature in

Shakespeare’s work.267 It is unsurprising, then, that Darwin, Gray, and their intellectual peers, would have been particularly well prepared to understand and debate the relation between art and nature. Darwin’s use of this dichotomy to categorize different types of selection should be approached with this literature in mind.

My aim in this chapter was to assess an intellectual congruence between the scientific and Darwinian revolutions: the role played by a likening of nature to artifice. The brief history I have provided indicates that this congruence is more than just nominal. Darwin and

Gray were well-aware of important 17th century debates about the relation between art and nature through the works of such revered and widely-read English writers as William

Shakespeare and Thomas Browne. These 17th century debates played a pivotal role in the ideology of the scientific revolution, as demonstrated through the philosophical reflections of

Robert Boyle. I have argued that Darwin and Gray used these 17th century debates to inform their own discussions, particularly those concerning the relation between artificial and natural selection. Darwin’s reliance on the art of breeding as a central element in his theory of natural selection cohered well with 17th century texts that argued for a dissolution of the art- nature distinction by claiming that Nature was simply the art of God. Darwin was aware of incompatible understandings of natural history that themselves trace back to 17th century reactions to a mechanical worldview. That he did not fully accept these methodological

267 On Coleridge and Wordsworth, see Abrams 1958, 122. Their disagreement, maybe unsurprisingly, is expressed through Shakespeare’s dialogue between King Polixenes and Perdita, the same dialogue used by Gray to defend Darwin above.

!110 prescriptions against human-manipulated nature show him to be taking a stand on a 19th century debate that could be illuminated by one that occurred throughout the 17th century.

!111 ! ! CHAPTER 5 ! Selection in the State of Nature Darwin and Wallace on Domestication ! ! ! ! ! ! ! ! I find myself having wondered several times at what time—how late you had to be born or at least professionalized in evolutionary and populations terms before the word selection lost all vestiges of its everyday meaning. It was many years before it occurred to me that that word was the same as the word selection [Lewontin pauses to spell it out: s-e-l-e-c-t-i-o-n] in English. When I realized that, then I reunderstood the whole plan of writing of Darwin’s Origin of Species. Richard Lewontin, 1974, Amidst a Conference on the Evolutionary Synthesis !

Although there are many similarities between their theories of evolution by natural selection, Alfred Russel Wallace and Charles Darwin continued throughout their lives to disagree about the significance of domestication. Was it the differences between artificial and natural selection which mattered or the similarities? What were the purported differences?

Building on the analysis in the preceding chapter, I will demonstrate how different ways of conceiving of the artificial-natural distinction were used to think through and express deep seated differences of opinion about whether domesticated organisms could be used to shed light on natural species change. ! !

!112 DOMESTICATION, REVERSION, AND NATURAL SELECTION

One of the now famous differences between Darwin’s and Wallace’s theories of evolution is the role played by domestication, or what Darwin called “artificial selection.”268

This difference persisted throughout their careers, onwards from their joint papers delivered to the Linnaean Society in 1858—the first time the learned public heard of evolution by natural selection.

In his 1858 contribution, Darwin drew repeatedly on an analogy between domestic varieties and natural races to argue for natural selection. He writes, “Let this work of selection on the one hand, and death on the other, go on for a thousand generations, who will pretend to affirm that it would produce no effect, when we remember what, in a few years,

Bakewell effected in cattle, and Western in sheep, by this identical principle of selection?”269

He repeats, “I can see no more reason to doubt that these causes in a thousand generations would produce a marked effect, and adapt the form of the fox or dog to the catching of hares instead of rabbits, than that greyhounds can be improved by selection and careful breeding.”270 At which point, he introduces natural selection, ! Man, by this power of accumulating variations, adapts living beings to his wants [...] Now suppose there were a being who did not judge by mere external appearances, but who could study the whole internal organization, who was never capricious, and should go on selecting for one object during millions of generations; who will say

268 See Young 1971; Bowler 1984; Cornell 1984 Rheinberger & McLaughlin 1984; Beddall 1988; Bartley 1992; Gayon 1998, etc. 269 Darwin and Wallace 1858, 49 270 Ibid., 49.

!113 what he might not effect? [...] I think it can be shown that there is such an unerring power at work in Natural Selection271 ! Darwin had accepted the importance of this comparison many years before, writing to his friend J. D. Hooker in 1844, “I believe all these absurd views [older views of immutability and mutability of species alike], arise, from no one having, as far as I know, approached the subject on the side of variation under domestication, & having studied all that is known about domestication.”272 When the Origin of Species was published in 1859, the analogy between domestication and natural selection only seemed to gain in significance, extending throughout the first four chapters of the book. His 1868 book, The Variation of

Animals and Plants under Domestication, lay in the same vein, as the name implies.

In his 1858 paper, Wallace, on the other hand, argued for the same conclusion— species mutability and common descent—by drawing on the differences between domesticated varieties and “wild animals in a state of nature.” Unaware of Darwin’s thinking but aware that an analogy with domestication had recently been drawn to support the opposite conclusion, species immutability, Wallace opted to argue against the analogy. He concludes, “no inferences as to varieties in a state of nature can be deduced from the observation of those occurring among domestic animals. The two are so much opposed to each other in every circumstance of their existence, that what applies to the one is almost sure not to apply to the other.”273

271 Ibid., 51. 272 Darwin to J D Hooker, 10 Nov 1844, No789. 273 Darwin and Wallace 1858, 61.

!114 While we might expect this difference to simply reflect their ignorance of each other’s theories in 1858, this was not the case. As time went on Darwin remained committed to the analogy between artificial and natural selection, and Wallace continued to approach the topic with extreme caution and noted reservation. When Wallace wrote Darwinism (1889) after Darwin’s death—a textbook of evolution by natural selection laid out in a structure intentionally similar to the Origin—he tellingly professed in the Preface: “It has always been considered a weakness in Darwin’s work that he based his theory, primarily, on the evidence of variation in domesticated animals and cultivated plants,” and thus, he, “endeavoured to secure a firm foundation for the theory in the variations of organisms in a state of nature.”274

This longstanding difference in their views is often explained by drawing attention to how each of them dealt with the phenomenon of “reversion”: whether varieties—natural or domesticated—have a tendency to physically revert over time back to their ancestral type.275

This common view creates the impression that Wallace thought reversion undermined any attempt at drawing an analogy between domestication and natural selection, whereas Darwin

274 Wallace [1889] 1897, vi. 275 Rheinberger and McLaughlin 1984; Beddall 1988; Richards 1998, 2005; Gayon 1998. Two other ways touch on differences between Darwin and Wallace with regard to (i) the importance of selection on permanent varieties, rather than individuals (see Bowler 1976, Gayon 1998; and see Kottler 1985 and Fagan 2008 for dissenting views) and (ii) relatedly, the fact that domesticated varieties could diverge from one another a great deal physically, but still interbreed (T. H. Huxley didn’t like this; see Huxley 1906, 256). The first is tied somewhat to reversion and I will return to it below. This view can explain why Darwin and Wallace differed in 1858, but not by itself why the difference maintained itself despite Wallace later accepting individual variation as important. The second is much more complicated. Although Darwin and Wallace had a fairly extensive discussion of hybrid sterility and whether species sterility could be brought about through natural selection (Wallace thought it could, Darwin thought it couldn’t), Wallace never raises the objection that domestic sterility makes them different from wild species. In Darwinism (1889, 184-5) he says that the sterility of domestic varieties can be explained away in terms of our preserving their sterility (because it is important to us) through domestication. Unlike Huxley, he did not think this was a problem for natural selection because he didn’t draw the artificial selection analogy in the first place. Darwin and Wallace’s discussions are further complicated by an underlying ambiguity over the meaning of “species” (Beatty 1982, 1986; Rheinberger & McLaughlin 1984).

!115 saw things differently. It creates the further impression that the connection between accepting reversion and denying an analogy with domestication was unproblematic.

This explanation is supported by what Wallace says in his 1858 paper and how

Darwin responds in Chapter 1 of the Origin.276 Although somewhat scattered, Darwin’s discussion is worth quoting in full. ! Having alluded to the subject of reversion, I may here refer to a statement often made by naturalists—namely, that our domestic varieties, when run wild, gradually but certainly revert in character to their aboriginal stocks. Hence it has been argued that no deductions can be drawn from domestic races to species in a state of nature. I have in vain endeavoured to discover on what decisive facts the above statement has so often and so boldly been made. There would be great difficulty in proving its truth: we may safely conclude that very many of the most strongly-marked domestic varieties could not possibly live in a wild state. In many cases we do not know what the aboriginal stock was, and so could not tell whether or not nearly perfect reversion had ensued. It would be quite necessary, in order to prevent the effects of intercrossing, that only a single variety should be turned loose in its new home. Nevertheless, as our varieties certainly do occasionally revert in some of their characters to ancestral forms, it seems to me not improbable, that if we could succeed in naturalising, or were to cultivate, during many generations, the several races, for instance, of the cabbage, in very poor soil [...], that they would to a large extent, or even wholly, revert to the wild aboriginal stock. Whether or not the experiment would succeed, is not of great importance for our line of argument; for by the experiment itself the conditions of life are changed. If it could be shown that our domestic varieties manifested a strong tendency to reversion,—that is, to lose their acquired characters, whilst kept under unchanged conditions, and whilst kept in a

276 It is assumed that Darwin is responding to a Wallacean-type argument when he discusses reversion at the beginning of the Origin. See Rheinberger & McLaughlin 1984 and Gayon 1998. This puzzle is also further complicated by confusing things Darwin sometimes says in letters, for example, he writes to Wallace in one of their earliest exchanges, “I have acted already in accordance with your advice of keeping domestic varieties & those appearing in a state of nature, distinct; but I have sometimes doubted of the wisdom of this, & therefore I am glad to be backed by your opinion” (May 1, 1857).

!116 considerable body, so that free intercrossing might check, by blending together, any slight deviations of structure, in such case, I grant that we could deduce nothing from domestic varieties in regard to species. But there is not a shadow of evidence in favour of this view: to assert that we could not breed our cart and race-horses, long and short-horned cattle, and poultry of various breeds, and esculent vegetables, for an almost infinite number of generations, would be opposed to all experience.277 ! This quotation suggests that Darwin sees nothing compelling in an argument from

“reversion” and also that he disagrees with how Wallace employed reversion in his 1858 paper. Here I want to complicate this account and, in doing so, clear up a number of common confusions surrounding the phenomenon of “reversion” as it relates to Darwin and Wallace.

Part 1 of this chapter is largely deflationary. I argue, against this common view, that whatever disagreements Darwin and Wallace might have had over the nature of “reversion,” these cannot by themselves (or in any straight-forward way) explain their differing views towards the importance of domestication. It will be shown that Darwin and Wallace largely agreed with regard to reversion and (paradoxically, given the above quotation) Darwin was— or at least became—more permissible when it came to tolerating reversion. Yet they continued to disagree about domestication.

In Part 2 I want to approach their disagreement in a different light, by looking at how each of them reflected more broadly on the artificiality of domestication and on how these reflections interacted with the conclusions they drew about whether the process of domestication was representative of natural selection in the “state of nature.” At stake was the ability of domestic varieties to serve as objects for the study of nature.

277 Darwin 1859, 14-5.

!117 !

PART 1. A MISCELLANY OF REVERSIONS

Charles Lyell’s Principles of Geology (1832) presents numerous examples of reversion as it was characteristically portrayed in the early-19th century: ! The black cattle that have run wild in America, where there were many particularities in the climate not to be found, perhaps, in any part of the old world, and where scarcely a single plant on which they fed was of precisely of the same species, instead of altering their form and habits, have actually reverted to the exact likeness of the aboriginal wild cattle of Europe.278 ! It seems now admitted by horticulturists, that none of our garden varieties of fruit are entitled to be considered strictly permanent, but that they wear out after a time; and we are thus compelled to resort again to seeds; in which case, there is so decided a tendency in the seedlings to revert to the original type, that our utmost skill is sometimes baffled in attempting to recover the desired variety.279 ! Lyell’s Principles was widely read, it was influential on the views of both Darwin and

Wallace. It is also presumably the work Wallace is responding to in his 1858 paper.280

Published in 1832, Lyell’s text represents commonly held views about the relation between organisms under domestication and those that exist in the “state of nature.”

In these passages, Lyell—largely occupied with discrediting Lamarck’s progressive theory of evolution—draws an analogy between domestic varieties and natural species in order to argue for the immutability of species. As he says, “The best authenticated examples

278 Lyell 1832:2, 30. My Emphasis. 279 Lyell 1832:2, 33. Second emphasis mine. 280 See Ruse 1979 for Darwin; see Beddall 1968 and McKinney 1972 for Wallace.

!118 of the extent to which species can be made to vary, may be looked for in the history of domesticated animals and cultivated plants.”281 But alas, when human interference is alleviated domesticated varieties, and thus by analogy also natural varieties, have a tendency to revert to the “likeness” of their original type.282 All variation is, according to this view, ephemeral. Domesticated varieties “may be regarded as extreme cases brought about by human interference, and not as phenomena which indicate a capability of indefinite modification in the natural world.”283 They could “never be perpetuated in a wild state for many generations, under any imaginable combination of accidents.”284

Two things should be said about early descriptions of reversion. First, commonly cited examples are vague and are for the most part hearsay; this is not a phenomenon that had undergone systematic study (leaving aside the fact that it had been documented by frustrated breeders for a long time).285

Second, they are used to argue for species immutability by analogy. The reversion of domesticated species to their original type only reinforces the idea of species stability and the permanence of species types. The “best authenticated examples of the extent to which species can be made to vary” still indicate an inherent principle towards type. Though Lyell’s discussion of animal breeding and horticulture is sporadic, it is understandable why Wallace

—who wrote his 1858 paper while in the Malay Archipelago taking Lyell’s Principles as his authoritative text—felt the need to argue against drawing an analogy between domesticated

281 Lyell 1832:2, 53. 282 As the examples above attest, this is true for both animals and plants. 283 Lyell 1832:2, 32. 284 Lyell 1832:2, 32. 285 See Müller-Wille & Rheinberger 2012, chapters 3 and 4.

!119 varieties and natural species: it was commonly held that such an analogy (if anything) only proved the immutability of species.286 As one historian summarizes, “the belief had grown

[from the 17th century onwards] that domesticated varieties were quite unlike wild species, being much more variable as a result of better nutrition and care and liable to revert in its absence.”287

Lyell’s discussion highlights one type of 19th century “reversion,” (i) Reversion by an inherent principle towards an ideal type. According to this view, when species reversion occurs, it is the result of an inherent tendency to return to an ancestral species type.

Variations are accordingly monstrosities; luckily by way of reversion Nature has a means for removing them.288

By 1858 Darwin and Wallace both deny the credibility of this type of reversion and they both have means for explaining it away. In the Origin, as we saw above, Darwin refers to this type or reversion as not having a “shadow of evidence” in its favor. His argument is a thought experiment: in order to prove the existence of this type of reversion it would have to be shown that varieties demonstrated reversion under unchanged external conditions.

Otherwise it would be ambivalent whether reversion was caused by the change of conditions or, as purported, an inherent tendency towards type. But, Darwin argues, this experiment has never been performed in reality, and furthermore, seems to be in contradiction to all evidence from the literature on breeding.

286 In the last chapter we saw many other examples of this from people like William Hopkins and Louis Agassiz. Darwin himself held this position in the late-1930s. 287 Evans 1984, 133. 288 In his Sketch of 1842, Darwin called attention to this type of reversion as an analogue of vis medicatrix, a principle of Hippocratic medicine referring to the “healing powers of nature” (Darwin 1909, p. 3). This is an apt characterization because it emphasizes an inherent tendency towards a fixed and ideal type. In full it refers to vis medicatrix naturae. Darwin had a copy of a book containing a discussion of this principle in his library.

!120 In later work, Darwin allows for the other type of reversion mentioned in his thought experiment, (ii) Reversion by changed external conditions. For instance, in the Variation, when discussing the “best known case of reversion, and that on which the widely-spread belief in its universality apparently rests,” Darwin writes “with feral pigs, exposure to the weather will probably favour the growth of the bristles, as is known to be the case with the hair of other domesticated animals, and through correlation the tusks will tend to be redeveloped.”289 Domesticated varieties may be caused to revert back to the likeness of the wild individuals because of similar external conditions. This is a far cry from the first type of reversion.

In 1858, Wallace provided a different argument against the first type of reversion. He writes, “Domestic varieties, when turned wild, must return to something near the type of the original wild stock, or become altogether extinct.”290 Since he was not given the chance to emend the 1858 paper before it was published, Wallace later clarified the passage: “That is, they will vary, and the variations which tend to adapt them to the wild state, and therefore approximate them to wild animals, will be preserved. Those individuals which do not vary sufficiently will perish.”291 By “will be preserved” he is making direct reference to natural selection. He continues to argue throughout his career that reversion is to be expected given the theory of natural selection. When domesticated varieties are released back into the state of nature they will either (i) perish altogether and be replaced by their wild counterparts or

(ii) gradually evolve by natural selection to the likeness of the wild species, since the wild

289 Darwin 1868:2, 33 and 47 290 Darwin and Wallace 1858, 60. 291 Wallace 1895, 31.

!121 species exemplify adaptation to local conditions.292 The wild species “must be that in which the various powers and faculties are so proportioned to each other as to be best adapted to procure food and secure safety.”293 In either case, an observant onlooker would notice that the domesticated variety had ceased to exist (and here Wallace means ceased to exist in the same form as it had under domestication). This influential argument against the first type of reversion gives rise to another, (iii) Reversion by natural selection. Wallace’s argument was taken up by Darwin and even Lyell in later work.294

The most straight-forward explanation for reversion was through blending inheritance. When domesticated varieties are released, they breed with their wild counterparts, producing offspring that have inherited some of the “wild” characteristics. As

Darwin puts it, when domesticated varieties are released into the state of nature “crossing alone would tend to obliterate their proper character.”295 Over time the differences between domesticated and wild varieties will tend to dissolve, resulting in a type of reversion to the wild form (especially effective in combination with the previous type of reversion), (iv)

Reversion by crossing with wild varieties.

292 Keep in mind here that Wallace was a proto-adaptationist. 293 Darwin and Wallace 1858, 60. 294 Darwin writes, “It has often been argued that no light is thrown, from the admitted changes of domestic races, on the changes which natural species are believed to undergo, as the former are said to be mere temporary productions, always reverting, as soon as they become feral, to their pristine form,” however, he continues, “This argument has been well combated by Mr. Wallace” (Darwin 1868:2, 425). Similarly, in the Antiquity of Man, Lyell writes, “It had usually been supposed by the advocates of the immutability of species that domesticated races, if allowed to run wild, always revert to their parent type. Mr. Wallace had said in reply that a domesticated species, if it loses the protection of Man, can only stand its ground in a wild state by resuming those habits and recovering those attributes which it may have lost under domestication” (Lyell 1863, 327). I thus strongly disagree with Gayon (1998, 40) that Darwin is making “Wallace say the exact opposite of what he did in fact say!” here. Darwin and Wallace saw eye-to-eye that this type of reversion is explained by natural selection. I do agree that Darwin and Wallace differed with regard to the implications of this reversion, as explained in the next section. 295 Darwin 1868: 2, 32.

!122 By the time he published the Variation in 1868, Darwin’s fascination with “reversion” had clearly intensified. He now referred to the phenomena as a “great principle of heredity” and recognized that any theory of inheritance would have to explain it.296 His theory of pangenesis did just this. And in his discussion of it, he also highlights another type of reversion, (v) Reversion through expression of latent characters. This type of reversion occurs regardless of changing an animal’s external conditions. As opposed to its closest relative, reversion type (i), reversion by latent characters could be explained without reference to an inherent tendency towards an ideal type.

Darwin repeatedly drew on two examples to demonstrate this type of reversion: first,

“that in every horse of every generation there should be a latent capacity and tendency to produce stripes, though these may not appear once in a thousand generations” and second,

“that in every white, black, or other coloured pigeon, which may have transmitted its proper colour during centuries, there should be a latent capacity in the plumage to become blue and to be marked with certain characteristic bars.”297

The latent capacity of pigeon plumage to become marked with certain characteristic bars through breeding may be encouraged by changes in external living conditions or outbreeding, but neither is necessary. There is something of an inherent tendency towards latent characters and Darwin used pangenesis to explain it.298 When introducing his

“provisional hypothesis” he thus mentions reversion as one of its explananda. Wallace also

296 See Bartley 1992, pp. 323-7. 297 Darwin 1868:2, 56. 298 Darwin writes, “For instance, there is a latent tendency in all pigeons to become blue, and, when a blue pigeon is crossed with one of any other colour, the blue tint is generally prepotent. When we consider latent characters, the explanation of this form of prepotency will be obvious.” Darwin 1868:2, 386.

!123 accepted this type of reversion. Referring directly to Darwin’s example, he writes, “This is considered to be a case of reversion to the ancestral type, just as the slaty colours and banded wings of the wild rock-pigeon sometimes reappear in our fancy breeds of domestic pigeons.”299

What this section has demonstrated is, firstly, that reversion was not a simple, unambiguous phenomenon, and secondly, contrary to the common view, differences with regard to reversion cannot by themselves explain Darwin’s and Wallace’s views towards the importance of domestication. Far from being at odds, they largely agreed about the facts of reversion and were united in their rejection of type (i).300 And yet still they disagreed with regard to domestication.301 ! ! !

299 Wallace [1889] 1897, 107. 300 Although Wallace without doubt saw (iii) as the most important explanation for reversion and played down the importance of (ii). 301 Not only did Darwin allow for reversion, he also put the phenomenon to work. In the Variation Darwin used reversion to deal with one of the stickiest problems for his theory: the fact that “transitional varieties” are seldom found and thus could not provide evidence of the slow process of continuous modification predicted by the theory of natural selection. As Darwin states the problem: “why, if species have descended from other species by insensibly fine gradations, do we not everywhere see innumerable transitional forms? Why is not all nature in confusion instead of the species being, as we see them, well defined?” (Darwin 1859, 171). Reversion-type (v) was exploited on many occasions to fill in the gaps. Darwin used the fact that varieties often expressed clearly defined latent characters through breeding to infer their ancestral form. He writes, “From these facts we see that the crossing of the several equine species tends in a marked manner to cause stripes to appear on various parts of the body, especially on the legs. As we do not know whether the primordial parent of the genus was striped, the appearance of the stripes can only hypothetically be attributed to reversion. But most persons, after considering the many undoubted cases of variously coloured marks reappearing by reversion in crossed pigeons, fowls, ducks, &c., will come to the same conclusion with respect to the horse-genus; and in this case we must admit that the progenitor of the group was striped on the legs, shoulders, face, and probably over the whole body, like a zebra” (Darwin 1868:2, 43; emphasis mine). Rather than rejecting “reversion” to support an analogy between domestication and natural selection, Darwin actually puts the concept to work within an evolutionary framework. Using reversion in this way continued into the 20th century. A very interesting discussion of this can be found in Wang 2012 about the political motivations for attempting to bring back the Aurochs, the supposed wild ancestor of domesticated cattle, in 1920-40s Germany.

!124 PART 2. ARE DOMESTICATED VARIETIES ARTIFICIAL AND WHAT DOES THIS ENTAIL?

WALLACE’S STATE OF NATURE AND STATE OF DOMESTICITY

While differing with Darwin as to its significance, and continuing to express a level of uneasiness, Wallace did not discount all knowledge gained from the study of domestication, especially later in his career. This is best demonstrated by how Wallace approaches domestication in his book Darwinism (1889). In the Preface, Wallace

(in)famously states, ! It has always been considered a weakness in Darwin's work that he based his theory, primarily, on the evidence of variation in domesticated animals and cultivated plants. I have endeavoured to secure a firm foundation for the theory in the variations of organisms in a state of nature; and as the exact amount and precise character of these variations is of paramount importance in the numerous problems that arise when we apply the theory to explain the facts of nature, I have endeavoured, by means of a series of diagrams, to exhibit to the eye the actual variations as they are found to exist in a sufficient number of species. By doing this, not only does the reader obtain a better and more precise idea of variation than can be given by any number of tabular statements or cases of extreme individual variation, but we obtain a basis of fact by which to test the statements and objections usually put forth on the subject of specific variability; and it will be found that, throughout the work, I have frequently to appeal to these diagrams and the facts they illustrate, just as Darwin was accustomed to appeal to the facts of variation among dogs and pigeons.302 ! This quotation is often taken as an illustration of his aversion to domestication. It says much more than this however. Firstly, Wallace is primarily interested in explaining evolution among organisms in the state of nature, not in the “state of domesticity.”303 As we saw in the

302 Wallace [1889] 1897, vi. 303 So-called in his 1858 paper.

!125 last chapter, this was a common distinction drawn by naturalists in the 19th century. Darwin’s work was founded on domesticated varieties and will thus always involve a further inference from principles that hold within the state of domesticity to principles that hold within the state of nature. A common objection was that the variation of domesticated varieties was artificial, a product of human interference, and thus not representative of the natural world.304

Since many people saw this inference as problematic, Wallace attempted to remove it by going straight to the source: variation in the state of nature.

Secondly, Wallace, paying lip service to Darwin’s Variation, does not argue that domestication is epistemically useless. He even includes a chapter in Darwinism about variation under domestication. This chapter demonstrates that domestication provides us with information about the origin of variation and about heredity. It demonstrates various general hereditary “facts” necessary for the action of natural selection: “the occurrence of incessant slight variations in all parts of an organism, with the transmission to the offspring of the special characteristics of the parents; and also, that all such slight variations are capable of being accumulated by selection till they present very large and important divergencies from the ancestral stock.”305 These are the “facts” of variation upon which is founded Darwin’s theory.306

304 See citations in last chapter. 305 Wallace [1889] 1897, 100. Many scholars have pointed out that Darwin saw domestication as an “experiment on a gigantic scale” (1868:1, 3; e.g., Schweber 1978; Rheinberger & McLaughlin 1984; Bartley 1992; Richards 1998). As we see here, this cannot straightforwardly explain differences between Darwin and Wallace with regard to their valuations of domestication. It might be argued that Wallace sees domestication as an artificial experiment, whereas Darwin as a natural experiment. This would be in accordance with what I claim below, but would require much more argumentation. 306 He also draws an analogy with domestic varieties and natural species to defend a point: add this here.

!126 But, thirdly, Wallace supplements (and supplants) this discussion with one about “the variations of organisms in a state of nature.” This chapter precedes the one about domestication and prepares the reader to appreciate that what is demonstrated through variation and heredity in domesticated organisms is not altogether inconsistent with what is found in the state of nature. This discussion makes the analysis of domestication largely superfluous, and acts to preempt objections that were leveled at Darwin’s theory. Wallace uses diagrams demonstrating variation in the state of nature in the same way that he understood Darwin to be appealing to variation in domesticated varieties: to demonstrate to the reader that variation is adequately supplied for natural selection to be effective (Figure

5.1 and 5.2). Wallace’s diagrams were based on measurements taken in the wild and thus directly represented wild variation.

!127

! Figure 5.1: Wallace’s Variation in the State of Nature. The left image represents measurements of the length of various traits for a number of species of the lizard genus Lacerta. The right image represents measurements of the length of various traits for twenty males of Icterus baltimore, Baltimore Oriole. (Wallace 1889, Darwinism) (© John van Wyhe, Darwin Online) !

!128 ! ! Figure 5.2: Darwin’s Variation in the State of Domesticity. The left image is of a Pouter pigeon. The right image is of a number of domestic pigeon skulls (from top to bottom: Wild Rock-pigeon, Columba livia; Short-faced Tumbler; English Carrier; Bagadotten Carrier). (from: Darwin 1868. Variation of Animals and Plants under Domestication) (© John van Wyhe, Darwin Online) ! Illustrating variation in the state of nature for the reader was certainly important. Most natural history texts at this time focused on the stable, characteristic traits of species in the state of nature, playing down any variation within that might upset systematic description.

The influential 17th century botanist John Ray, for example, famously argued that characters that could be externally manipulated (e.g., by humans) or that were for other reasons too variable, were insufficient to distinguish species and thus should be ousted from taxonomy.307

Such views advocated what historian and biologist Ernst Mayr famously called the

307 See Müller-Wille & Rheinberger 2012, p. 60, and also chapt. 2-3.

!129 “typological” view of species: that members of a species are all instances of the specific type, and that the perpetuation of the species consists in the production of ever more instances of the type.308 According to this view, all variation was error; variation was the result of disturbances that interfered with the natural tendency of organisms of a particular type to produce offspring of that type. This was obviously a view hostile to evolutionary change.

What Wallace and Darwin wanted to argue, using among other things the above diagrams, was that variation was natural. (Although as we will see below, Wallace never could shake the idea that artificial selection, rather than variation, was unnatural and a disturbance.)

Without having spent time in the state of nature and observed the variations first-hand

—as Wallace had done—the common 19th century reader would have no way of adjudicating whether natural selection could work at all. Wallace thus says of the Duke of Argyll’s reading of the Origin, “The noble author represents the feelings and expresses the ideas of that large class of persons who take a keen interest in the progress of science in general, and especially that of Natural History, but have never themselves studied nature in detail, or acquired that personal knowledge of the structure of closely allied forms [...] which is absolutely necessary for a full appreciation of the facts and reasonings.”309 By “nature” Wallace means “state of nature,” as opposed to the domesticated nature. His diagrams provided a solution to this problem.

Although domestic varieties were not epistemically useless, Wallace continued from

1858 onwards to avoid invoking Darwin’s analogy between domestication and natural selection. He held his ground in this regard for two reasons, both having to do with

308 Mayr 1982. 309 Wallace 1868 [1895], 141.

!130 differences between the state of nature and the civilized state—and both expressed views that were deep-seated among Victorians.

First, Wallace worried that the analogy with domestication suggested that natural selection required an intentional selector and thus imbued the natural world with a problematic level of consciousness.310 It is easy to forget today that Darwin brought the term

“selection” into common biological parlance, and changed its meaning considerably (as

Lewontin points out in the epigraph). For instance, in his 1860 review of Darwin’s Origin, the Harvard professor and natural historian Louis Agassiz wrote, “The fallacy of Mr.

Darwin's theory of the origin of species by means of natural selection, may be traced in the first few pages of his book, where he overlooks the difference between the voluntary and deliberate acts of selection applied methodically by man [...] and the chance influence which may effect animals and plants in the state of nature. To call these influences ‘natural selection,’ is a misnomer.”311 Wallace addressed such issues in a paper section entitled, “Mr.

Darwin’s Metaphors Liable to Misconception,” where he argued that Darwin was merely speaking metaphorically and did not mean to anthropomorphize “nature.” But even before this in-print defense, Wallace worried about Darwin’s analogy. He wrote to Darwin in 1866 that he had been struck by the “utter inability of numbers of intelligent persons to see clearly or at all” that natural selection was “self-acting” and did not involve an intelligent chooser behind the scenes.312 Such misunderstandings, Wallace thought, could be avoided by removing the terminology of selection itself. He suggested that “survival of the fittest” was

310 Wallace 1868 [1895], 144-5 311 Agassiz 1860, 147. 312 Wallace to Darwin, July 2, 1866, Wallace to Darwin.

!131 more apt. “Selection” was, after all, a civilized human activity, preformed by a selector with a high degree of consciousness and agency. Selective breeding occurred, he said, in the “state of domesticity,” not the state of nature. To say that nature selects individuals best fitted to their environments, just as humans might select the swiftest greyhounds, is simply misleading.313 Eventually Wallace gave in and accepted “natural selection,” but he never fully accepted the analogy with domestication.314

Wallace also held his ground for a different reason. Although the origin of variation and the heredity principles were essentially the same for domestic varieties and natural species, he believed that the accumulation of that variation was entirely different, in effect, it was diametrically opposed. In the state of nature, natural selection resulted in organisms that become progressively well fitted to their environments (in the mid-19th century sense of fulfilling the requirements of a particular environment or way of life). Selection in the state of nature is for fitness. In the state of domesticity, however, artificial selection led to organisms that were maladapted. Selection under domestication was for useless, fanciful or superficial characters and thus often indirectly at odds with fitness. Consider a few of

Wallace’s comparisons between domestication and natural selection:

313 Although Darwin recognized the possibility that the phrase would be misunderstood he remained reluctant to replace it. He added a section to the 3rd edition of the Origin (1861, 84-5) explaining how the phrase should be understood. Even by the 2nd edition (1860) Darwin had begun to worry about his use of “selection,” adding the word “metaphorically” to the following sentence on page 84: “It may metaphorically be said that natural selection is daily and hourly scrutinising, throughout the world, every variation, even the slightest” (my emphasis). See also Thomas Henry Huxley's paper “Criticisms on 'The Origin of Species'” (1864) which addresses M. Flourens' concerns about Darwin's use of “selection.” For an historical treatment of the Darwin- Wallace disagreement about Darwin's use of “selection” see Rheinberger & McLaughlin (1984) and Gayon (1998, chapter 1). See Young (1971, 1993) for excellent discussions of the reception more generally of natural selection. 314 And furthermore he continued to speak of natural selection as a passive “weeding out” process, rather than an active “selection” process. For more detail and the broader context see discussion in Young 1985, Chapter 4. For Wallace’s use of “weeding out”, see for example, Wallace 1890, 1892.

!132 ! The essential difference in the condition of wild and domestic animals is this,—that among the former, their well-being and very existence depend upon the full exercise and healthy condition of all their senses and physical powers, whereas, among the latter, these are only partially exercised, and in some cases are absolutely unused. A wild animal has to search, and often to labour, for every mouthful of food [...] There is no muscle of its body that is not called into daily and hourly activity [...] The domestic animal, on the other hand, has food provided for it, is sheltered, [etc.] [...] Half of its senses and faculties are quite useless [...] in the domesticated animal all variations have an equal chance of continuance; and those which would decidedly render a wild animal unable to compete with its fellows and continue its existence are no disadvantage whatever in a state of domesticity. Our quickly fattening pigs, short- legged sheep, pouter pigeons, and poodle dogs could never have come into existence in a state of nature [...] The great speed but slight endurance of the race horse, the unwieldy strength of the ploughman's team, would both be useless in a state of nature [...] Domestic animals are abnormal, irregular, artificial; they are subject to varieties which never occur and never can occur in a state of nature: their very existence depends altogether on human care; so far are many of them removed from that just proportion of faculties, that true balance of organization, by means of which alone an animal left to its own resources can preserve its existence and continue its race.315 ! As man has considered only utility to himself, or the satisfaction of his love of beauty, of novelty, or merely of something strange or amusing, the variations he has thus produced have something of the character of monstrosities. Not only are they often of no use to the animals or plants themselves, but they are not unfrequently injurious to them. In the Tumbler pigeons, for instance, the habit of tumbling is sometimes so excessive as to injure or kill the bird [...] This peculiar character of domestic productions distinguishes them broadly from wild species and varieties, which, as will be seen by and by, are necessarily adapted in every part of their organisation to the conditions under which they have to live.316 !

315 Darwin and Wallace 1858, 59-61. 316 Wallace [1889] 1897, 100.

!133 Wallace’s use of “artificial” to disparage domesticated varieties has a multidimensional meaning. It means, firstly, created by human interference and existing in a state of domesticity. Secondly, it means ephemeral, capricious, or useless and associated largely with changing fashion. The artificiality of domestic varieties is contrasted with natural varieties which exhibit a “true balance of organization” and are “necessarily adapted in every part of their organization to the conditions under which they live.” And third,

“artificiality” has an air of “unreality” about it. Artificial, domesticated varieties are not as real as natural varieties; they are fleeting examples of human interference, not the products of natural law.317

The fact that the accumulation of variation in the state of nature and under domestication were at odds indicated to Wallace that the analogy was a poor one. His argument involves the amalgamation of two ideas constitutive of his thinking. The first is that

Wallace, like Darwin, was strongly influenced by Lyell’s “uniformitarianism,” which held that explanations of former changes in the history of life must only avail themselves of natural causes now in action.318 If a cause cannot be demonstrated to be acting at present, then we cannot simply invent it to explain events that happened in the past.319

317 This was a commonly held view among naturalists; in the last chapter we considered Asa Gray’s lament that the study of domestication “has been generally neglected by naturalists, because these races are not in a state of nature” (Gray [1860] 1963, 20). 318 McKinney 1972, 32-33. Wallace brought Lyell’s book with him on his first trip to South America. See also Beddall 1968, Wilson 1970 and Berry 2002, p. 156. Lyell’s famous Principles of Geology was aptly subtitled “Being an Attempt to Explain the Former Changes of the Earth’s Surface by Reference to Causes Now in Operation.” 319 To explain how species had evolved in the state of nature, Wallace had to demonstrate how causes now in action in the state of nature could by analogy have led to species evolution in the past. As Wallace wrote in an early notebook, “While the inorganic world has been strictly shown to be the result of a series of changes from the earliest periods produced by causes still acting, it would be most unphilosophical to conclude without the strongest evidence that the organic world so intimately connected with it had been subject to other laws which have now ceased to act.” Wallace 1855, quoted in McKinney 1972, 23.

!134 Second, Wallace’s presentation of evolution often gives the impression of a two-level process.320 The first is the creation of “permanent varieties” which differ from the natural species because of (something like) selection on individual differences through individual competition.321 The second, and the more important level in Wallace’s early writings, is the struggle between “permanent varieties.” Wallace writes, if “any species should produce a variety having slightly increased powers of preserving existence, that variety must inevitably in time acquire a superiority in numbers,” relative to the parent species. When conditions become unfavorable the parent species may be replaced by the variety. “The superior variety would then alone remain, and on a return to favourable circumstances would rapidly increase in numbers and occupy the place of the extinct species and variety.” In such a case, “The variety would now have replaced the species.”322

A species’ departure from its original type thus occurs as a permanent variety supplants the natural species.323 The permanent variety is, in return, supplanted by a better

320 See Bowler 1976 and Gayon 1998, chapter 1. Wallace’s use of “variation” and “variety” in his 1858 paper is quite perplexing and while much literature has abounded which addresses this problem, there is no obvious answer as to what Wallace meant by these terms in 1858 (see the early disagreement between H. F. Osborn and E. B. Poulton in 1896 quoted in Kottler 1985; also see for a summary of recent literature). While I have here presented Wallace’s theory of evolution as a two-step process (like Bowler 1976, 2013) because I believe this to be the most sensible, my discussion does not hinge on this view and can equally be interpreted in terms of individual selection only: in this case, Wallace treats domesticated varieties as still unfit and unable to outcompete or eventually replace the parent species, and the second step is instead understood in terms of individual selection. 321 “[A]s the individual existence of each animal depends upon itself, those that die must be the weakest,” writes Wallace, “while those that prolong their existence can only be the most perfect in health and vigour—those who are best able to obtain food regularly, and avoid their numerous enemies. It is, as we commenced by remarking, ‘a struggle for existence,’ in which the weakest and least perfectly organized must always succumb” (Darwin and Wallace 1858, 56-7). See Bowler 2013, 63ff. Some historians argue that Wallace did not mean to present evolution as a two-level process. I agree with Peter Bowler, Janet Browne, and Michael Ruse that at least in 1858 he did. 322 Ibid., 1858, 58. 323 The latter really just another permanent variety, since Wallace admits there is “generally no means of determining” which is the natural species and which the permanent variety (Darwin and Wallace 1858, 53).

!135 adapted permanent variety and so on. This leads to the “tendency of varieties to depart indefinitely from the original type,” the title of Wallace’s 1858 paper.

Thus, being a uniformitarian, Wallace was looking for a theory of evolution that could be demonstrated to be currently in action and that could by analogy explain species evolution in the past. Domestication did not provide a model of this process since the varieties it created could never outcompete the parent species in the state of nature. Why was this?

Because domestication gave rise to varieties that were unfit. The human disturbance which created domestic varieties was not the same as the process which caused species evolution in the past—the latter consisting of better fitted varieties replacing those that are less well fitted.

Domestication was ephemeral, capricious, and useless and associated largely with changing fashion. It created monsters with lopsided characters, not organisms perfectly adapted to their surrounding environment and displaying a “true balance of organization.”

Wallace’s strategy when it came to explaining natural selection was thus to deduce natural selection from a number of laws or principles or facts which held true in the state of nature itself, without recourse to domestication.324 This strategy is lucidly expressed in his

1868 paper “Creation by Law” where Wallace presents a “series of facts or laws” that “are mere statements of what is the condition of nature.”325 These laws include: (1) law of multiplication in geometrical progression; (2) law of limited population; (3) law of likeness of offspring to their parents; (4) law of variation; (5) law of unceasing change of physical conditions upon the surface of the earth; (6) equilibrium or harmony of nature (i.e., well-

324 This was more consistent with strategies employed in the physical sciences. See for example William Hopkins’ review of Darwin’s Origin. Hopkins 1860. 325 Wallace 1868 [1895], 143.

!136 adapted species flourish). Wallace saw evolution by natural selection to be straight-forwardly entailed by these facts and needing no support by way of analogy to domestication.326

As radical as Wallace was in other respects, his views reflect the teachings of traditional natural history. The artificiality of domestic varieties had always entailed a kind of unreality. They were products of humans interfering with natural laws, and not stand ins for natural processes themselves. As products of interference, domestic varieties were not outside of nature per se, but they were definitely outside of “natural nature,” to use Sorel’s terms.327 Wallace’s discussions were therefore compatible with mainstream accounts of domestic varieties—as not themselves the objects of speculation or study. The objects of study were well-adapted species in the state of nature. What happened when organisms left the state of nature? To Wallace, and many others, they became unfit.

Wallace’s belief that the accumulation of variation through domestication was at odds with fitness extended even to domesticated humans. In an uncharacteristic moment for an otherwise self-effacing man, Wallace summarized his intellectual contributions, emphasizing two papers on “human selection,”

326 More accurately Wallace used the phrase “survival of the fittest,” rather than “natural selection” in this paper. Natural selection, he thought, reminded readers of human selection, and he wanted to distance himself from that view. Darwin also at times appears to be emphasizing that natural selection is a consequence of certain laws, e.g., “These laws, taken in the largest sense, being Growth with Reproduction; Inheritance which is almost implied by reproduction; Variability from the indirect and direct action of the external conditions of life, and from use and disuse; a Ratio of Increase so high as to lead to a Struggle for Life, and as a consequence to Natural Selection, entailing Divergence of Character and the Extinction of less-improved forms” (1859, 489-90). 327 I quoted George Sorel’s distinction between “natural nature” and “artificial nature” in Chapter 1. Domestic varieties were either seen as having themselves triumphantly emerged from the state of nature to serve humans (as civilized domestication practices replaced those of savage domestication practices) or having been enslaved and degenerated to appease human fancy, depending on the author’s point of view. (For former see Ritvo 1987 and Crawford 1863; for latter see quotes and footnotes in last chapter. The latter seems to have been predominantly the view of the naturalists to which Darwin and Wallace were indebted. See Secord 1985.) Either way, since they were not part of the state of nature, it was argued that domesticated animals and the processes involved in domesticating them could not be used to shed light on the state of nature—and often times such an inference was strongly advised against. There are many examples of this which were cited in the last chapter. See especially Louis Agassiz’s notes on Darwin’s Origin as well as subsequent publications.

!137 ! These deal with different aspects of the same great problem—the gradual improvement of the race by natural process; and they were also written partly for the purpose of opposing the various artificial processes of selection advocated by several English and American writers. I showed that the only method of advance for us, as for the lower animals, is in some form of natural selection, and that the only mode of natural selection that can act alike on physical, mental, and moral qualities will come into play under a social system which gives equal opportunities of culture, training, leisure, and happiness to every individual. This extension of the principle of natural selection as it acts in the animal world generally is, I believe, quite new, and is by far the most important of the new ideas I have given to the world.328 ! As with animals, artificial selection on humans is ineffective if the goal is the general

“improvement” of the race. Only natural selection can accumulate variation in a direction that increases fitness. The civilized human will is too susceptible to fancy, caprice, and most importantly, wealth.

Wallace’s views also reflect a much discussed popular understanding of the relation between the state of nature and the state of domesticity as a recovery narrative, from

“savagery” to “civilization.”329 The language of the “state of nature” comes from 17th century political texts that endorsed the human flight from nature by way of social contracts.

This progressive narrative permeated the imaginative consciousness of 19th century intellectuals. “With every step of this progress in civilization,” T. H. Huxley famously writes

328 Wallace 1905:2, 389. By “natural selection,” Wallace has in mind sexual selection. While he argued against Darwin’s theory of sexual selection as applied to animals, he promoted it as a form of selection in the state of domesticity. 329 For a brief history of the recovery narrative see Merchant 1995. See Ruse 1996 for a discussion of “progress” more broadly speaking, and its relation to evolutionary theories.

!138 near the end of the century, “the colonists would become more and more independent of the state of nature; more and more, their lives would be conditioned by a state of art.”330

At times, Wallace promoted these views himself. Man is, “a being apart,” Wallace agrees, “since he is not influenced by the great laws which irresistibly modify all other organic beings,” including most importantly natural selection. Rather than existing in a state of nature, man has become “fitted for the social state.” Not only had man escaped natural selection, but “he is actually able to take away some of that power from nature.” We can anticipate the time, for better or worse, “when the earth will produce only cultivated plants and domestic animals; when man's selection shall have supplanted ‘natural selection.’”331

And to Darwin’s great chagrin, Wallace in 1869 went one step further, claiming that natural selection could not account for the mental qualities of modern man—some higher force must be invoked.332

In summary, according to Wallace, domestication was superfluous once “facts” about the origin of variation and heredity could be proven to exist in the state of nature. In addition, the analogy between domestication and natural selection was misleading because the accumulation of variation that resulted from each process was at odds—domestication created maladapted monsters, whereas natural selection created organisms fitted for survival in nature. Domestication and domestic varieties were, for Wallace, unnatural. That is,

Wallace invoked the artificial-natural distinction to argue against the domestication analogy. !

330 Huxley 1894, 19. 331 Wallace [1864] 1870, 325-8. 332 Wallace 1869. He was an outspoken proponent of spiritualism for the rest of his life.

!139 CHARLES DARWIN ON ART AND NATURE

Darwin’s discussions, in contrast, collapse the artificial-natural distinction itself.333

Many of the properties characteristic of Wallace’s “state of domesticity”—most uniquely tied to civilization—are shown by Darwin to have counterparts in the state of nature.334 I will focus on three cases that demonstrate how strongly Darwin’s discussions encouraged a merging of the artificial world of domestication and civilization with the wild and savage world of the state of nature (all of the emphasized phrases being loaded with cultural significance that Darwin exploited repetitiously). The first case focuses on the artificiality of the state of nature itself, the second on the naturalness of domestication, and the third undermines attributions of “monstrous.” Darwin’s strategy is to render unanswerable questions such as, “is this a product of art or nature?”335 !

THE ART OF SEXUAL SELECTION

Initially, Darwin’s theory of sexual selection had the appearance of an afterthought.336

It was called on to explain what he believed natural selection could not: the interesting and

333 I am indebted in this section to papers by Cornell 1984, Rheinberger & McLaughlin 1984, Alter 2007, Burnett 2009, and Müller-Wille 2009. 334 Unsurprisingly many of these ideas were also strongly contested. 335 Since Darwin’s reasoning with regard to the analogy from artificial to natural selection has attracted much attention (and was briefly dealt with in the last chapter), I will spend less time explaining his views and instead just point out a few interesting, yet often overlooked, cases. The discussions I indicate here are in contrast to a different way in which Darwin exploits the artificial-natural distinction. For instance, “We have seen that man by selection can certainly produce great results, and can adapt organic beings to his own uses, through the accumulation of slight but useful variations, given to him by the hand of Nature. But Natural Selection, as we shall hereafter see, is a power incessantly ready for action, and is as immeasurably superior to man’s feeble efforts, as the works of Nature are to those of Art.” (Darwin 1859, 61) Here he is being, what we might call, strategically inconsistent. By emphasizing the differences between Art and Nature he is trying to get around the problem that domestic varieties freely interbreed. For a discussion see Burnett 2009. 336 In the Origin Darwin introduces sexual selection, but it doesn’t take centre stage until Darwin's (1871) The Descent of Man, and Selection in Relation to Sex.

!140 unusual sexual differences in ornamentation and behavior found within many groups of animals, for example, the male peacock’s large and conspicuous tail. Its significance, however, increased over the course of his life and it came to sit interestingly between the processes of domestication and natural selection: a kind of choice-based selection occurring in the state of nature.

Sexual selection differed from natural selection in that the selective force was not a result of the “struggle for existence,” but was instead a result of the struggle between individuals of one sex to find and secure mates. It was a weaker type of selection, which resulted not in “death to the unsuccessful competitor, but few or no offspring.”337 Even still, sexual selection was invoked to explain the most anomalous and beautiful aspects of the state of nature—for example, the argus pheasant’s plumage and the bower of the bower bird.

This struggle for mates took one of two forms. It either took the form of a contest between the males, where the victor secured a mate, or was more peaceful and involved elaborate displays or songs. In its more peaceful guise, sexual selection depended on choice, not unlike in domestication.338 Consequently, in many of the examples illustrating sexual selection Darwin relies on an analogy between these two types of selection to substantiate his case. The result is surprising and fascinating. Just as Wallace observed above, choices can be quite whimsical, often based on a love of beauty or just novelty and tied to current fashion

337 Darwin 1859, 88. 338 Here I am considering how Darwin compares sexual selection and artificial selection. The parallels between sexual selection and natural selection, in terms of the degree of conscious planning behind the “choice” made, can be brought out by considering the other form of sexual selection, what might be called the less peaceful “contest” form. In this form, sexual selection does not involve choice at all; it is, rather, the result of the contest that determines who is able to find a mate. In this sense, the “struggle between the males for possession of the females” seems to involve no more consciousness than the “struggle for existence” between individuals in nature. The “contest” form of sexual selection, therefore, would seem to fit in at the low end of the spectrum, next to natural selection.

!141 rather than fitness.339 For Wallace, this entailed that domesticated varieties were unrepresentative of their counterparts in the state of nature. For Darwin, the theory of sexual selection highlights the capriciousness of the state of nature itself.340

In the Origin, Darwin describes sexual selection as follows, ! successive males display their gorgeous plumage and perform strange antics before the females, which standing by as spectators, at last choose the most attractive partner. Those who have closely attended to birds in confinement well know that they often take individual preferences and dislikes [...] It may appear childish to attribute any effect to such apparently weak means: I cannot here enter on the details necessary to support this view; but if man can in a short time give elegant carriage and beauty to his bantams, according to his standard of beauty, I can see no good reason to doubt that female birds, by selecting, during thousands of generations, the most melodious or beautiful males, according to their standard of beauty, might produce a marked effect.341 ! As can be seen, he is not at all shy in allowing for the female bird to “choose” a mate based on her “preferences and dislikes,” drawing the reader's attention to the similarity between the female bird (i.e., an instance of sexual selection) and the human breeder (i.e., an instance of domestication) to argue for the efficacy of sexual selection. Furthermore, it is by a standard of beauty—and not of utility—that the female bird makes consistent choices. “In the same manner as man can give beauty, according to his standard of taste, to his male poultry,” he writes, “so it appears that in a state of nature female birds, by having long selected the more

339 Again, “fitness” here is meant in the mid-19th century sense of fulfilling the requirements of a particular environment or way of life. 340 See Cronin 1991 and Milam 2010b for reflections on this theory and its history. See also Müller-Wille 2009. 341 Darwin 1859, 87.

!142 attractive males, have added to their beauty.”342 Darwin thus suggests that it is not just in the state of domesticity that beauty, rather than utility, may affect the production of varieties.

But, Wallace had asked, what of changing fashion? What of capriciousness? Are these not characteristics of domestication alone? Darwin responded, just as “any fleeting fashion in dress comes to be admired by man, so with birds a change of almost any kind in the structure or coloring of the feathers in the male appears to have been admired by the female.”343

Darwin was even prepared to go “a short distance” with the Duke of Argyll—not someone he would normally go any distance with—when the he wrote, “that variety, mere variety, must be admitted to be an object and an aim in Nature.”344 “It would even appear that mere novelty, or change for the sake of change,” Darwin agreed, “has sometimes acted like a charm on female birds, in the same manner as changes of fashion with us.”345 And against one writer, who “oddly fixed on Caprice ‘as on of the most remarkable and typical differences between savages and brutes’,” Darwin writes, “lower animals are, as we shall hereafter see, capricious in their affections, aversions, and sense of beauty.”346 The male

Argus pheasant, Darwin concluded, is “more like a work of art than of nature” (Figure

5.3).347

342 Darwin 1871:1, 259. My emphasis. 343 Darwin 1871:2, 74. 344 Darwin 1871:2. 230. 345 Darwin 1871:2, 230. Darwin is so confident that birds have a similar, although less developed, sense of aesthetic appreciation governing their selection of mates that he says in the Origin, “the tuft of hair on the breast of the turkey-cock [...] can hardly be either useful or ornamental to this bird;—indeed, had the tuft appeared under domestication, it would have been called a monstrosity” (Darwin 1859, 90). In case we might misunderstand, in later additions of the Origin Darwin replaced “ornamental to this bird” with “ornamental in the eyes of the female bird” (Darwin 1872, 70; my emphasis). In other words, we should be leery of attributing all sexual differences to sexual selection because we find traits, like this tuft of hair, which cannot be favoured by the female bird because it is ugly in the eyes of human breeders! 346 Darwin 1871:1, 64-5. 347 Darwin 1871:2, 92.

!143 !

Figure 5.3: An Argus Pheasant Sitting on a Branch of a Tree. Etching by John Le Keux, 1783-1846. (© Wellcome Library, London) ! To make such claims seem plausible—that birds choose at all would strike many

Victorians as problematically anthropomorphic—Darwin draws comparisons between the abilities of the female bird and “the savage,” what many in the 19th century saw as a middle- ground between the human-animal dichotomy and the object much of speculation. When it comes to selection based on aesthetic discrimination, he says, female birds may even out- rank savages. “The taste for the beautiful, at least as far as female beauty is concerned, is not of a special nature in the human mind,” he writes. And judging “from the hideous ornaments

!144 and the equally hideous music admired by most savages, it might be urged that their aesthetic faculty was not so highly developed as in certain animals, for instance, in birds.” Of course,

“no animal would be capable of admiring such scenes as the heavens at night, a beautiful landscape, or refined music,” but then again, such “high tastes” are “not enjoyed by barbarians or by uneducated persons.”348

By selecting certain types of individuals over others—pigeons with shorter beaks, feathers that run backwards or carunculated skin—human breeders create long-term changes in their stocks. These choices are often based on an assessment of what is most beautiful, an assessment that is whimsical, certainly liable to caprice, and based on current fashion.

Breeders are guided in their selection by external “circumstances” and “fashions,” or even idiosyncrasies associated with their individual “frame of mind” or “character.”349 What

Darwin draws our attention to when discussing sexual selection is that the taste for the beautiful and liability to caprice or fashion can also be found in nature. Darwin is not here denying the artificiality of domestication, but is instead playing up the artificiality of the state of nature. !

UNCONSCIOUS SELECTION IN THE STATE OF NATURE

Darwin undermines the artificial-natural distinction in a different way by exploiting the concept of “unconscious selection.” The scholarly focus on Darwin’s analogy between artificial and natural selection has tended to hide the fact that “artificial selection” is graded.

Darwin spills a lot of ink—especially in the Variation—explaining and exploiting types of

348 Darwin 1871:1, 64. 349 Darwin 1868:2; 212, 208-210, 214, 214.

!145 artificial selection. Most notably methodical and unconscious selection, though Darwin is aware that “Unconscious selection so blends into methodical selection that it is scarcely possible to separate them.”350

Methodical selection is the most artificial. It requires that the selector be consciously aware of their favourite animals, be consciously aware of the modifying power of selecting them over many generations, and work towards the goal of producing new varieties. This type of selection “is the magician's wand, by means of which [the breeder] may summon into life whatever form and mould he pleases.”351 It is this type of selection which, in characteristic portrayals, figures in Darwin’s argument by analogy.

But Darwin’s other type of selection, unconscious selection, should not be overlooked. This type of selection, as the name implies, occurs when breeders unintentionally alter their breeds by unconsciously selecting the best or most fancied individuals to mate.352

As Darwin writes in the Origin, “a man who intends keeping pointers naturally tries to get as good dogs as he can, and afterwards breeds from his own best dogs, but he has no wish or expectation of permanently altering the breed.” He continues, “I cannot doubt that this process, continued during centuries, would improve and modify any breed, in the same way as Bakewell, Collins, &c., by this very same process, only carried on more methodically, did greatly modify, even during their own lifetimes, the forms and qualities of their cattle.”353

Darwin believed that this form of selection led to far more significant results than

350 Darwin 1868:2, 211. 351 Darwin 1859, 31. 352 See Darwin 1859, 34-6. 353 Ibid, 34-5.

!146 methodological selection, and moreover, was theoretically interesting because it closely resembled natural selection.354

This resemblance is two-fold. Unconscious selection resembles natural selection because of its “unconsciousness”—Darwin demonstrated that breeds can be significantly altered despite the “selectors” having no intention to do so. This was one way in which he evaded the criticism that natural selection required intention to be efficacious. Unconscious selection also resembles natural selection in terms of its physical locality; unconscious selection occurs not just on the estates of those who keep pointers, but also within the state of nature itself.355 To make the second point, Darwin again exploited a 19th century category that captured the imagination of Victorians and was the topic of much discussion: the

“savage” or “barbarian.” Living within the state of nature, indigenous peoples, many

Victorians suspected, were those that the progressive narrative failed to capture; they were uncivilized, Godless and lawless. To some they were a link between humans and the state of nature, to others they were a different species entirely, more akin to the “brutes” with which they cohabited.356 Darwin drew repeatedly on the “primitive” domestication practices he observed in the Tierra del Fuego region while on the Beagle voyage. This type of domestication was essentially occurring in the state of nature.

These dimensions of unconscious selection are brought out in the following story that

Darwin returned to on many occasions and in different works:

354 Darwin writes in the Variation, “This form of selection has probably led to far more important results than methodical selection, and is likewise more important under a theoretical point of view from closely resembling natural selection.” Darwin 1868:2, 424; my emphasis. 355 See Burnett 2009. 356 See Browne 1995, Chapter 10. See also Marx 2005 for a later 19th century and American perspective.

!147 ! If there exist savages so barbarous as never to think of the inherited character of the offspring of their domestic animals, yet any one animal particularly useful to them, for any special purpose, would be carefully preserved during famines and other accidents, to which savages are so liable, and such choice animals would thus generally leave more offspring than the inferior ones; so that in this case there would be a kind of unconscious selection going on. We see the value set on animals even by the barbarians of Tierra del Fuego, by their killing and devouring their old women, in times of dearth, as of less value than their dogs.357 ! The barbarians of Tierra del Fuego do not consciously attempt to create new strains from their domesticated animals, nor do they even consciously breed the best individuals. They may not even be conscious of their own preference for certain animals or make a distinction between species, as the purported fate of their old women suggests.358 In times of famine, these savages simply kill off and devour those least preferred by them (including their own elders!) leaving the rest to reproduce, in a “kind of unconscious selection.” Darwin is drawing attention to the fact that selective breeding activities do not require much consciousness at all, which is significant if we remember that natural selection is supposed to be an unconscious analogue of human selection.

This “unconscious selection” is also occurring in the state of nature. As I said, many of Darwin’s readers would not have considered the Fuegians to be of the same species as

357 Darwin 1859, 36. 358 Darwin elaborates further on this example in the Variation, “When the Fuegians are hard pressed by want, they kill their old women for food rather than their dogs; for, as we were assured, 'old women no use—dogs catch otters.' The same sound sense would surely lead them to preserve their more useful dogs when still harder pressed by famine.” (1858:1, 214-5) Darwin also adds that the same process of unconscious selection occurs when “aborigines of Australia” are given European kangaroo dogs. 359 Burnett 2009.

!148 themselves, let alone civilized.359 With examples like these, Darwin sought to bring his readers closer to the border with natural selection.

Darwin took advantage of the resemblances between unconscious selection and natural selection to underwrite the naturalness of domestication, subverting the claim that breeding is merely unnatural interference. For example he writes, “But the gardeners of the classical period, who cultivated the best pear they could procure never thought what splendid fruit we should eat; though we owe our excellent fruit, in some small degree, to their having naturally chosen and preserved the best varieties they could anywhere find.”360 These

“gardeners” did not consciously breed the best fruit, they just naturally preserved the ones that tasted best, in the same way as bees might be said to naturally choose to pollinate those flowers most accessible by flight.361 The art of human breeding, in this light, appears quite natural.362 Or, as Gray observed in the last chapter, “the Art itself is Nature.”

In light of this discussion, I find Darwin’s remark about domestication at the beginning of the Descent of Man striking. He writes, ! It has been asserted that man alone is capable of progressive improvement; that he alone makes use of tools or fire, domesticates other animals, possesses property, or employs language; that no other animal is self-conscious, comprehends itself, has the power of abstraction, or possesses general ideas; that man alone has a sense of

360 Darwin 1859, 37; my emphasis. 361 As two historians put it, “Man’s natural historical past shows him to be a part of nature himself. Considered as a species, man has even in a state of nature always been an instance of selection for the nature which surrounds him; he has always functioned as an element of natural selection” (Rheinberger and McLaughlin 1984, 355). 362 See also Burnett 2009 and Rheinberger and McLaughlin 1984.

!149 beauty, is liable to caprice, has the feeling of gratitude, mystery, &c.; believes in God, or is endowed with a conscience.363 ! When considering a set of commonly ascribed differences between humans and nature—ones he will ultimately attempt to debunk—he includes among this list domestication. As we have seen, the way Darwin puts the concept of unconscious selection to work acts to undermine a distinction, founded on conscious intent and physical locality, between selection as it occurs in the state of nature and domestication as it is done by humans.

Darwin at times entertained an even more extreme view, the logical conclusion of the last quotation. In his unpublished manuscript Natural Selection, Darwin muses over an interesting relationship between a species of ant and aphid.364 Though certainly unconscious, these ants have modified a species of aphid for their own benefit to such an extent that,

Darwin says, we might call the aphids “their domesticated cattle!”365 Darwin ends the passage with the following striking remark: “Hence we see that man is not the only animal which keeps domesticated animals.”366 Such examples, we should note, are not the same as his classic examples of natural selection, which involve Nature selecting, but hint at particular animals doing the selecting just as humans select. These provide an indication that

Darwin toyed with applying “domestication” outside of even the most unconscious of human

363 Darwin 1871:1, 49; my emphasis. 364 Natural Selection was the book he was writing when he received Wallace’s letter in 1858 that prompted their joint publication and the subsequent publication of the Origin of Species. 365 Stauffer 1975, 511-2. 366 Ibid. Furthermore, in an early paper published in The Gardeners' Chronicle and Agricultural Gazette Darwin considers the relationship between “insect-agents” and the orchids they pollinate over many generations (especially pertinent given the above “gardeners” quotation) (Darwin 1860, 528). See also the Origin where Darwin draws the readers attention to the relationship between natural selection and insect selection: “When our plant, by this process of the continued preservation or natural selection of more and more attractive flowers, had been rendered highly attractive to insects, they would, unintentionally on their part, regularly carry pollen from flower to flower; and that they can most effectually do this, I could easily show by many striking instances.” (93)

!150 breeders.367 Domestication crossed the divide between art and nature, making plausible arguments by natural selection. !

ARTIFICIAL MONSTERS

Domestic varieties were at this time often described by naturalists as “monstrosities,” unfit because they were created and maintained by man, not by nature.368 Purportedly human interference created awkward organisms with lopsided characters, not organisms perfectly adapted to their surrounding conditions and displaying a “true balance of organization.” For many this was enough to warrant—or could be rhetorically used to defend—their choice of a different object of study, natural species. Darwin was critical of such a perspective.

Darwin argued at times that what is considered a “monstrosity” is largely arbitrary and does not reflect a tangible difference between the state of nature and the state of domesticity. We should be “cautious in deciding what deviations ought to be called monstrous,” he writes. Although “there can hardly be a doubt that, if the brush of horse-like hair on the breast of the turkey-cock had first appeared on the domesticated bird, it would have been considered a monstrosity.” And while, “we might call the wattle or corrugated skin round the base of the beak of the English carrier-pigeon a monstrosity, [...] we do not thus

367 Another example of this can be gleaned from outside Darwin's work. Dr. Hermann Müller and Darwin corresponded from about 1867 till Darwin's death in 1882. Müller wrote a series of articles published in Kosmos in 1878 entitled, “Die Insekten als unbewusste Blumenzüchter” or “Insects as unconscious selectors of flowers.” In these articles he—according to a 1879 issue of American Naturalist—splits the evolution of flowers into peculiarities due to natural selection and peculiarities due to insect-selection. Müller seems to have run to the logical conclusion with Darwin's concept of unconscious selection, and sees no problem attributing this to insects. See Trelease 1879. Darwin also corresponded (more extensively) with Hermann Müller's brother Johann Friedrich Theodor (Fritz) Müller. 368 See also Secord 1986, 523.

!151 speak of the globular fleshy excrescence at the base of the beak of the male Carpophaga oceanica.”369

In other words, if we are consistent with our attributions of “monstrous,” it would have to apply to many things that exist throughout the natural world as well, he argues.

Lopsidedness is as much a property of creatures in the state of nature, as it is domestic productions. Such attributions are therefore not a good way to divide domestic varieties from natural species: many of our domestic varieties “cannot be said to have an unnatural appearance; and no distinct line, as it seems to me, can be drawn between natural and artificial races.”370 Although in “extreme cases the distinction is plain, in many other cases an arbitrary line has to be drawn.”371

Moreover, Darwin attacked the two most common reasons why domestic varieties were labeled monstrosities: they lacked a true balance of organization and they were maladapted. To argue that species in nature, such as the giraffe, could have gained their harmoniously coordinated characters through the natural selection of small, continuous variations, Darwin takes the opportunity to draw attention to the harmoniously coordinated characters of domesticated varieties. “If the short-faced tumbler-pigeon, with its small conical beak, globular head, rounded body, short wings, and small feet—characters which appear all in harmony—had been a natural species,” he writes, “its whole structure would have been viewed as well fitted for its life.”372 Not the way many naturalists would have seen things. He continues, “Look at the greyhound, that perfect image of grace, symmetry, and

369 Darwin 1868:2, 413. 370 Ibid., 245. 371 Ibid., 413. 372 Ibid., 221.

!152 vigour; no natural species can boast of a more admirably co-ordinated structure, with its tapering head, slim body.”373

Darwin also repeatedly drew attention to the adaptedness of domestic varieties. This, however, required seeing “adaptation” in a more context-specific or relative light, and in this way differently from Wallace and many others. The reason Wallace considered domestic varieties to be maladapted was because they were not adapted to conditions in the state of nature; they could never outcompete a wild species. Adaptation to man’s purposes is, for

Wallace, to be maladapted. Darwin on the contrary seems not to restrict the application of the term to conditions in the state of nature. Of the 18th century naturalist Johann Blumenbach’s remark that “many dogs, such as the badger-dog, have a build so marked and so appropriate for particular purposes, that I should find it very difficult to persuade myself that this astonishing figure was an accidental consequence of degeneration,” Darwin retorts, “had

Blumenbach reflected on the great principle of selection, he would not have used the term degeneration, and he would not have been astonished that dogs and other animals should become excellently adapted for the service of man.”374

The idea that domestic varieties were adapted, rather than maladapted, appears also to be a fairly early discovery for Darwin. In March 1839, he notes, “Get instances of adaptation in varieties — greyhound to hare. — waterdog, hair to water — bulldog to bulls. — primrose to banks — cowslip to fields.” These being “adaptations just as much as Woodpecker. —

373 Ibid. 374 Darwin 1868:2, 220. By the “great principle of selection” is Darwin here referring just to artificial selection or a more broad category? 375 Quoted in Hodge and Kohn 1985, 200.

!153 only here we see means.”375 What is fascinating about this quote is how freely he passes from adaptation under domestication to adaptation in nature.

Darwin’s multi-angled criticisms of the concept of a “monstrosity” put considerable stress on the distinction between domestication and natural selection. Again, as the boundary between this distinction is dissolved, domestication starts to appear less like a disturbance in nature’s “normal development,” and more like a part of nature. !

CONCLUSION: HUMAN DISTURBANCE AND ARTIFICIALITY FOR DARWIN AND WALLACE

In the first part of this chapter I argued against the common view that disagreements between Darwin and Wallace over the nature of reversion can explain their differing views about the significance of artificial selection. In the second part I argued that a better way to make sense of their differing views is by taking into account the broader context in which their discussions were embedded. Their disagreement over the appropriateness of the analogy between artificial and natural selection took place in the broader context of Victorian speculations about the state of nature (“the natural”) and the state of domesticity (“the artificial”). Each supported their view by drawing on a common stock of normatively loaded, tropes and perceptions (consciousness, monstrosity, caprice, etc.). Wallace argued against the analogy between artificial and natural selection and in the process reinforced a distinction between the state of domesticity and the state of nature. This fit well with Wallace’s beliefs regarding the unique status of humans: they alone were endowed with a mental and moral capacity that required a supernatural explanation. Darwin argued for the analogy and

!154 collapsed the distinction.376 This fit well with Darwin’s larger project to reduce the gap between human beings and the rest of nature (which often consisted in displaying standard human abilities, activities, creations, and so on, and showing that they differ only in degree from those found in nature). To Wallace, Darwin’s analogy was too anthropomorphic: human domestication was unlike, in fact it was in direct opposition to, what happened in wild nature.

To Darwin, Wallace’s anthropomorphic injunctions were simply anthropocentric.

This disagreement between Darwin and Wallace is, among other things, about the status of domestic varieties and has a bearing on my discussion of artifacts in Chapter 3. Are domestic varieties artifacts caused by human disturbance or are they natural phenomena in their own right? Wallace, as was typical of 19th century naturalists, treats domestication as human disturbance that results in artifacts, not natural phenomena, that is, well-adapted organisms in the state of nature. Human actions are treated as factors that must be excluded from our analysis as they are outside the proper object of study. This is reflected in the judgment that domestic varieties are maladapted—to be properly adapted is to be adapted to conditions characterizing the state of nature. That is the type of adaptation that natural selection is supposed to explain.

Wallace pays lip service to Darwin and uses domestication to illustrate the origin of variation and the principles of heredity, directing the reader to Darwin’s “remarkable”

Variation.377 But he intends this discussion only to supplement one about the origin of

376 I have not looked into the more historical question of what might have caused them to hold different views about the artificiality of the state of nature (or the naturalness of domestication). Certainly it had something to do with where they developed their theories and what sort of organisms there abounded: Darwin crossing the boundary between the world of breeding and the world of natural history, Wallace working to classify species in the field (see Secord 1981 and 1986 for Darwin; see Fagan 2008 for Wallace). 377 Morgan and Morrison 1999.

!155 variation and the principles of heredity in the state of nature. And, importantly, human selection is not a snap-shot of natural selection in action for Wallace. Domestication is like the production of seedless watermelons; not supernatural creation, that is, within the bounds of physical law, but also not something that could stand for or explain how the present constitution of species has been created from past causes that are still now in action. The existence of domesticated varieties is ephemeral and tied too closely and contingently to human disturbance. Wallace would therefore rather just show how natural selection follows from a set of descriptive laws that he then demonstrates to hold in the state of nature.

Domestication is useful, but it is not natural selection.

Darwin’s view is less straightforward; he is often frustratingly, but strategically, inconsistent. It helps to think in terms of Darwin’s uses, instead of his use, of domestication.

Here I have called attention to cases where it appears that Darwin’s strategy is to make questions such as, “is this a product of art or nature?” unanswerable, maybe even to collapse the distinction altogether. Of course fancy pigeons are artificial, but what about sheep modified by completely unconscious selection, dogs modified by the Fuegians, male birds with their uselessly fashionable colors, or ants and their domesticated cattle? In these cases a distinction between the artificial and the natural is put under considerable strain. And with it the distinction between the state of domesticity and the state of nature. We should not forget that these distinctions themselves were central to common disparagings of domestic varieties as proper objects of study.

Darwin’s strategy, in the face of opposition such as that expressed by Wallace, could have been to just deny the “artificiality” of domestication altogether—to play down the

!156 capriciousness, ephemerality, consciousness, or whimsical nature of domestication. In places he does do this, for instance, he tries to show that the conscious intent of the selector is not a prerequisite for the selection to be efficacious. But he also does the inverse: he plays up the artificiality of the state of nature. He concentrates on concepts classically associated with civilized society and demonstrates that they can be not only found in the state of nature but are constitutive of the selection therein.

For Darwin, domesticated varieties are not created through the human disturbance of a natural process, they are not unnatural, but are instead the result of a natural process. They are adapted to a particular set of circumstances outside the state of nature—and we shouldn’t forget that the domestic dog was certainly doing better than the wolf in England. Darwin’s

Variation, in effect, is as much a natural history of domesticated organisms as it is about the specific facts or laws of variation.378 Through this work Darwin actively made domestication a part of natural history by writing the authoritative text, thus further corroborating his prior uses of domestication in the Origin.379

These differences between Darwin and Wallace are not simply about finding artifacts and removing them—as is common in all scientific practice—but about whether a purported artifact is actually a natural object. They could not simply be resolved by appealing to technical or practical factors having to do with the production of domestic varieties. There is a more fundamental difference in their outlooks that compels us to appeal to wider concerns about the artificial-natural distinction itself. Are domestic species artifacts (Wallace’s view) or are they natural phenomena (Darwin’s view)? Are human actions outside of the state of

378 Gayon 1998, 47. 379 See Secord 1986, 536, who originally makes this point.

!157 nature? These questions can only be answered through attempts to navigate the boundary between artificial and natural.

As we have seen in this chapter, this boundary is navigated through classification.

Depending on which way domestic varieties are classified makes a difference to whether they are considered valid objects for the study of nature, including whether or not they can tell us about natural selection. As the artificial-natural distinction is defined and redefined so too is what counts as valid for biological study and whether humans count as disturbances.

As one final consideration, in Chapter 1 I hypothesized that the concepts of “nature” and “natural” were not about physical place. In the discussions of the last two chapters I have demonstrated that what is meant by “nature” is “state of nature”—the term inherited from

17th century political philosophy. This term is used 39 times in Darwin’s Origin of Species alone, and was a standard category that stood for the proper object of study in natural history, that is, things as they exist in their uncivilized or undomesticated state. The “state of nature,” however, does not refer to a specific physical place—it is an ideal involving no civilized human disturbance. Darwin exploits this fact: there were people there, the Fuegians, and they practiced a “natural” domestication. ! !

!158 ! ! CHAPTER 6 ! Searching for What Nature has Wrought Dobzhansky and the “Natural” Experimental Fruit Fly380 ! ! ! the work on natural selection is of necessity confined mainly to experiments in which the environment of the organism is modified artificially [...] Skeptics may contend that if the change in the environment is wrought directly or indirectly by man, the resulting selection is no longer “natural.” ! Theodosius Dobzhansky, 1937, Genetics and the Origin of Species It is advisable to avoid collecting in orchards or too close to human habitations because in such places the Drosophila populations consist chiefly of cosmopolitan introduced species [...] Such “garbage” species are relatively rare in woods or other “natural” habitats. ! Theodosius Dobzhansky to J. George Harrar, 1950 The standpoint which I for one find extremely difficult to understand is that any such thing is contrary to nature. What is contrary to nature and what isn’t, and who is there to tell? [...] Well, again, it is perfectly reasonable to ask, is Muller’s “Bravest New World” project contrary to Nature? ! Theodosius Dobzhansky, 1962, Reminiscences quite authoritative biologists in the 1920s and 1930s [contended that ...] Drosophila mutants described by the Morgan school were monstrosities, and they were all found in laboratory bottles, not in natural populations, products of some kind of disruption of hereditary materials in highly artificial environments. Theodosius Dobzhansky, 1980, The Birth of the Genetic Theory of Evolution ! Of the many biologists involved in the evolutionary synthesis of the 1930s and 40s,

Theodosius Dobzhansky is often singled out as its instigator. The research program endorsed in his Genetics and the Origin of Species (1937) fruitfully blurred the line between the

380 A significant portion of the research for this chapter was completed at the American Philosophical Society. Throughout this chapter, I will use the abbreviation APS.

!159 outdoor, “natural” world of naturalists and the indoor, “artificial” world of experimentalists— a line that had impeded the synthesis of evolutionary theory and genetics. Although

Dobzhansky was perhaps concerned with “the natural” in a way that his immediate laboratory predecessors were not, his experimental practice was fundamentally similar to theirs and manipulative laboratory work continued to play a dominant and necessary role throughout his career. What then was so “natural” about Dobzhansky’s research and why might this have mattered? !

THE ARTIFICIAL AND THE NATURAL IN THE EVOLUTIONARY SYNTHESIS PERIOD

As the founding of the Journal of the History of Biology in 1968 testifies, the 1960s and 1970s saw an unprecedented interest in the history of the life sciences. Many members of this younger generation had trained, worked or were influenced by the renowned evolutionary biologist turned historian Ernst Mayr. Mayr was never afraid to tell these younger historians when he thought they had overlooked details that were of crucial importance for the historical actors themselves, and when it came to the 20th-century, Mayr often exercised the authority of being an historical actor as well as an historian.

In 1973, Mayr sent the geneticist Michael Lerner—also an historian and historical actor—a draft of an essay review of recent books on the historical origins of genetics. “It is obviously a rather personal and subjective interpretation but I am trying to point out things that are always ignored,” Mayr wrote. “I am trying to make future students of the subject more critical.”381 He was “greatly relieved” when Lerner did not object.382 What “things” did

381 Ernst Mayr to I. Michael Lerner, 19 Oct 1972, APS. 382 Mayr to Lerner, 6 Nov 1972, APS.

!160 he think this younger generation had missed? Mayr’s published article shows how closely related they are to the central topic of this dissertation: ! One major development of the 1915-1930 period is entirely ignored [...] I am referring to the work on the genetics of geographic races. [...] studies dealing with natural populations did far more to convince the “Darwinians” of the Mendelian nature of selectively important “natural” variation than either the “artificial” (as they called them) Drosophila mutations or mathematical calculations [the traditional historical foci]. [...] This work was a crucial factor in making the “new [evolutionary] synthesis” of the 1930s and 1940s possible.383 ! Mayr’s allusion to “natural” populations, “natural” variation, and “artificial” mutations illustrates a way of classifying the objects of biological study that became increasingly prevalent during the 20th century. The natural-artificial dichotomy was used to classify races, hybrids, populations, species, mutations, variations, processes, environments, methods of study, you name it. These classifications had complex descriptive-normative dimensions; they often concealed underlying concerns about which sorts of objects and methods one should employ to produce valid knowledge—which were the right ones for testing evolutionary theory and which were not. As I mentioned in Chapter 1, although historians and philosophers widely recognize that “natural” (as well as its antitheses) carries normative weight, it is surprising that such accounts rarely extend this analysis to the above uses in biology, or to epistemic rather than moral concerns in general.384

383 Mayr 1973, 152-3. My emphasis. 384 But see Gooday 1991 and Schiebinger 2004. For “nature” as a moral authority in science, see: Daston and Vidal 2003; for similar issues regarding non-native species, see: Chew and Laubichler 2003, Chew and Hamilton 2011; for similar issues in environmental philosophy, see Ereshefsky 2007

!161 In this chapter I will demonstrate that concerns about the relation between “the artificial” and “the natural” had an important part to play in the historical period referred to by Mayr as the evolutionary synthesis. It is necessary to say a brief word about the synthesis itself. The phrase “modern synthesis” (later “evolutionary synthesis”) was introduced by biologist Julian Huxley in 1942 to mark a recent general consensus amongst biologists that

Mendelian genetics (the study of the principles of inheritance and mutation) was compatible with the theory of Darwinian selection (that evolution is gradual and the result of natural selection acting on continuously varying characters).385 Since this time the “synthesis” has taken on a much more complicated meaning; in fact, depending on who and what you read

(of both historians and historical actors), you will get very different impressions of what intellectual or material “pieces” were synthesized and what made the synthesis possible or even significant.386 This chapter is not about the synthesis; it is about the time period associated with the synthesis (roughly 1930-60) and more specifically about the thought and practice of someone who is often referred to as a synthesis “architect,” Theodosius

Dobzhansky.

Dobzhansky makes for a particularly interesting case study. Not only did concepts of

“the natural” and “the artificial” play important roles throughout his career, as is suggested by the epigraph quotations, but his uses of the distinction were quite influential. First of all, the synthesis of which Dobzhansky is often said to have instigated is associated with the breakdown of a previous divide between naturalists and experimentalists, a divide that itself

385 Huxley 1942. 386 See especially Cain 2009 for a discussion of uses of “evolutionary synthesis.” Cain argues that we abandon the unit concept of “evolutionary synthesis” altogether as this assumed, unitary narrative tends to hide, rather than illuminate, many of the interesting changes occurring in the life sciences throughout this time period.

!162 has a lot to do with drawing a distinction between the artificial and the natural (see Appendix

1).387 His scientific work of the late-1930s is heralded as productively blurring the line between the outdoor, “natural” world of natural historians and the indoor, “artificial” world of experimental biologists. His friend, the geneticist L. C. Dunn, prefaced Dobzhansky’s seminal Genetics and the Origin of Species (1937) with the comment that Dobzhansky epitomizes a new “Back to Nature” movement in biology: “The methods learned in the laboratory are good enough now to be put to the test in the open and applied in that ultimate laboratory of biology, free nature itself.”388 The book is described by one geneticist and editor of Dobzhansky’s work as “the first book that had consequences in the biological world of the twentieth century comparable to the effects of [Charles Darwin’s] The Origin of

Species by means of Natural Selection in the nineteenth century.”389

Dobzhansky’s movement called for a new object of study, the “natural population,” which was distinguished from the “artificial” or laboratory population. The natural population was to be found in the field, rather than created in the laboratory.390 This movement also required a switch of research organism: from the—in Dobzhansky’s words

—“domestic,” “artificial,” “cosmopolitan,” or “garbage” fruit fly, Drosophila melanogaster, to its “wild,” “natural,” or “noble” cousin, Drosophila pseudoobscura. (Note the normativity implicit in these “thick” concepts.) In fact, Dobzhansky’s studies of “natural populations”

387 Following a 1974 conference on the evolutionary synthesis, the historian William Provine wrote to conference attendee Theodosius Dobzhansky asking him how it felt to be singled out as “the perpetrator of the great synthesis of the 1930’s and 40’s.” Provine to Dobzhansky, , APS. See Allen 1979, Mayr 1980. 388 Dobzhansky 1937, xii. 389 Glass 1980, 2. 390 His most famous series of papers, the “Genetics of Natural Populations” (43 papers; 1938-1976), focused on the genetics and evolution of natural populations and, as the editors of its bounded collection write, “taken as a whole, the [series] is the most important single corpus in modern evolutionary biology” (Lewontin et al. 1981, xi). Through this venuw his thoughts on the proper object of study reached a wide audience.

!163 were, in all but a few cases, simply studies of populations of D. pseudoobscura. It might not be so surprising that, when asked what made Dobzhansky different from his laboratory-based precursors, one student, Hampton Carson, recalled, “Dobzhansky felt strongly that laboratory methods should be employed to reflect exactly what natural selection in nature had wrought.

This is the reason for his emphasis on analyses of specimens taken directly from nature.”391

While these brief examples go a way towards establishing that Dobzhansky thought the artificial-natural distinction was worth invoking, they also open a number of further alleys for investigation: Why was “the natural” meaningful and useful for Dobzhansky? Why did Dobzhansky believe that a study of evolution based on natural populations would provide different (and better) results than one on artificial populations? Given that his practice remained fundamentally laboratory-based, in what way did he think his experimental practice met this “natural” goal? The partial answers to these questions given below will turn out to be complicated and patchy; rather than a single narrative I will flag a variety of scientific, socio-political, and personal factors that interact with one another.

I will argue for two theses in this chapter: (i) that assumptions about what was artificial and what natural played a role in structuring Dobzhansky’s research practice and his evaluations of others and their work; (ii) Dobzhansky tended to be skeptical of studies that were too artificial, and this skepticism was reinforced throughout his career, because he believed they promoted both misrepresentative and undesirable views of the nature and evolutionary consequences of genetic variation.

391 Quoted in Provine 1981, 52. Emphasis in original.

!164 In the last two chapters we saw that in 19th century Britain the artificial-natural distinction corresponded to a distinction between “civilized” or “domesticated” and the “state of nature.” Darwin drew on these tropes in his work on sexual and unconscious selection. In this chapter—on the other side of what many have seen as an “experimental turn” in biology

—the distinction means something different: it corresponds to a distinction between things made in the laboratory by humans and things made in the world outside of the laboratory by natural processes.392 !

A BRIEF ACCOUNT OF DOBZHANSKY’S INTELLECTUAL LIFE IN RUSSIA

Although this is a chapter about American evolutionary biology, it would be impossible to reconcile Dobzhansky’s later attitudes without a brief look at his Russian roots.

There are four points to be made here and they come from a wealth of scholarship on

Dobzhansky’s early life.393 The first is that in Russia the distinction between “naturalists” and

“experimentalists” was less distinct. When Dobzhansky was born in 1900 in the Ukraine,

American biology, across the Atlantic ocean, was in the midst of considerable disarray: ! The more descriptive naturalists and the more analytical experimentalists had been at war in a number of fields [...] the dichotomy was real, persisted into the twentieth century, and had profound ramifications on the ways in which members of each tradition viewed the same biological problem. [...]

392 For the experimental turn, see below, and Allen (1975) and Coleman (1977). 393 This very brief discussion of Dobzhansky’s Russian roots owes a lot to four sources: Mark Adams’ work (see Adams 1968, 1980, 1994), Provine 1981, Dobzhansky’s own recollections especially his Reminiscences, and the Russian scholars whose work is provided in Adams 1994 (Alexandrov 1994, Konashev 1994, Krementsov 1994).

!165 While such different perspectives appeared to exist in a number of areas of biology, probably nowhere were the battle lines drawn more sharply than in the study of evolution and heredity. Here, the effect of each tradition conditioned the types of questions asked and the kinds of answers considered acceptable. In fact, each tradition determined in some ways the very problems considered worth studying.394 ! If this accurately describes the disunified state of academic affairs in American biology during the late-19th and early-20th centuries, then Dobzhansky’s early training in

Russia put him in a unique position compared to his American counterparts. In Russia,

Darwinism had enjoyed a more favorable reception than in many other countries.

Dobzhansky had read the Origin of Species by the age of 15, for example.395 The prevalence of Darwin was not attributable to purely scientific reasons, Darwinism was thought to be well suited to Marxist materialism (and as Dobzhansky later wrote, “The criterion of validity of theories of evolution [at this time] was their congruity with dialectical materialism”).396

Futhermore, Russian politics at this time favored research institutes that were “synthetic,” that brought together many biologists from diverse backgrounds, all of whom were expected to have training in natural history as well as experimental methods.397

Dobzhansky’s career in natural history began at the age of 16, collecting butterflies, but within a year he had switched to more serious entomological studies of ladybird beetles

(family Coccinella).398 The importance of these “naturalist” studies cannot be

394 Allen 1979, 179. 395 Provine 1981, 7. 396 Adams 1980, 246; Dobzhansky 1980, 230. 397 See Adams 1980. 398 As Dobzhansky recollected in 1974, “I was, if you please, an entomologist specializing in taxonomy of Coccinellidae, lady beetles. Seeing a lady beetle still produces in me a flow of a love hormone. The first love is not easily forgotten.” Quoted in Gould 1982, xxxix.

!166 overemphasized. While his later career was dominated by laboratory work with fruit flies,

Dobzhansky wrote more than 30 descriptive papers on ladybird beetles, the majority in

Russian or German, and continued to draw on this research as examples in later work.399 A principle focus of Dobzhansky’s research at this time was evolutionary, and he was concerned with many of the same problems that would later make him famous across the

Atlantic: the variation among and between geographically distinct populations, and the causal sequence that leads from mutations arising in a population to the origins of species differences. Thus the seeds of Dobzhansky’s later work on the ubiquity of variation and speciation were already sown during his early years in Russia.

As a result of political instability in Russia between 1905 and the end of the Civil War

(1917-1922), the scientific community had been largely cut off from the recent advances in classical genetics made in the Western world, particularly those made by Thomas Hunt

Morgan’s laboratory at Columbia University. As the war abated, this research came in all at once and was widely disseminated. By early 1922, after reading a review of Morgan’s work published by the Russian geneticist Iurii Filipchenko, Dobzhansky was decided on a future in genetics. In 1923, he was invited by Filipchecko himself to join the Department of Genetics at the University of Leningrad.400 At Leningrad Dobzhansky worked on the genetics of the famous fruit fly Drosophila melanogaster—which had been brought to Russia by Hermann J.

Muller, a former Morgan lab member, in August 1922—but continued to remain committed to entomology and to studies of the ladybird beetle. For Dobzhansky, and many of his

Russian contemporaries, the divide between experimentalist and naturalist camps was

399 See Dobzhansky 1937, 51ff. 400 It became Leningrad (formerly Petrograd) the day after Dobzhansky arrived in January 1924.

!167 indistinct. But, as the historian William Provine put it, by 1926 Dobzhansky had reached a frustrating plateau in his research efforts, “He knew a great deal about the systematics of

Coccinellidae, but almost nothing about its genetics [which turned out to be very complicated]. He knew some Drosophila melanogaster genetics, but nothing about natural populations of Drosophila.”401 His later studies using a different fruit fly, D. pseudoobscura, would fill this gap.

Besides allowing Dobzhansky the freedom to move unimpeded between experimentalist and naturalist camps, foreshadowing his later “Back to Nature” move,

Dobzhansky’s early Russian training was significant for three other reasons. The first is that

Dobzhansky was able to keep up with advancements in Russian genetics while his American colleagues could not. The other two reasons, which Dobzhansky considered constitutive aspects of Russian biology, were summarized in a later recollection, ! Russian biologists habitually viewed [evolution] in two contexts: that of philosophical implications, and in the light of life as it exists in pristine nature. To be acceptable, a paradigm had to pass scrutiny from these points of view, in addition to being valid on purely factual grounds.402 ! Although it would be naive to take Dobzhansky’s account at face value, it at least sheds light on important aspects of his own career. The first he labels “philosophical implications” and by this he has primarily in mind the meaning of evolution for humans and human societies.

Although Americans tended to be less inclined to discuss such philosophical topics, Russian

401 Provine 1981, 13. 402 Dobzhansky 1980, 241.

!168 biologists debated these topics vigorously and openly. Dobzhansky was surprised when he arrived in America in 1927 that so little of the wider implications of research was discussed by his lab mates Alfred Sturtevant and Calvin Bridges.403 The first university classes

Dobzhansky taught included those on the cultural implications of biology.404 He wrote in

1962, ! Evolution is the part of biology which has the highest and most direct implications, the most reflections in fields not immediately connected with biology, such as sociology and philosophy [...] This sociological-philosophical angle was really the aspect which interested me most in the whole field of biology, from the earliest days, I think really from my first reading of Darwin at about age fifteen. It is hardly surprising that both during the pre-Revolutionary days in Russia and during the post- Revolutionary days, to biologists, these philosophical-humanistic implications of evolutionism were in the center of attention. I think it is not an exaggeration to say that probably this interest is what made me, if not a biologist, at least an evolutionist.405 ! The second aspect Dobzhansky draws attention to is perhaps more cryptic but no less significant. He says that a “paradigm” must pass scrutiny in the context of “life as it exists in pristine nature.” I take him to be making the point that a theory, to be acceptable, must be able to explain phenomena as they exist in nature, rather than just in the laboratory.406 This is, in my view, equivalent to what Dunn said above, that the “ultimate laboratory of biology” was “free nature itself.”

403 See Dobzhansky 1962a. See also Kohler 1994 256-7. 404 Provine 1981, 9. 405 Dobzhansky 1962a, 351-2. 406 My reading is informed by what Dobzhansky says about Russian biologists in other contexts, and also the particular scientific disputes he was involved in throughout his life.

!169 In summary, there are four points to be made about Dobzhansky’s Russian roots: in

Russia, as compared to America, the distinction between naturalist and experimentalist was less distinct; Dobzhansky could read and keep up with Russian scientific literature; he was always interested in evolution, especially human evolution, and had personal experience with the ubiquity of variation in nature in ladybird beetles; and he acknowledged that a theory must account for, or be representative of, “pristine nature” (for reasons we have yet to consider). Each of these points will play a role in what follows. !

THE MORGAN LAB AND CLASSICAL GENETICS

After emigrating from Russia in 1927 to take up a Rockefeller fellowship,

Dobzhansky spent his early American career as a member of T. H. Morgan’s laboratory at

Columbia University and then CalTech—famously known as the “fly group.” Dobzhansky largely gave up his descriptive studies of Coccinella, as well as field work in general, after this move; as with the other members of Morgan’s lab, he was expected to work on the fruit fly D. melanogaster and because there were plenty of laboratory stocks of the fruit fly, field collection was unnecessary. The Morgan lab, both before Dobzhansky arrived as well as after, addressed problems of classical genetics through the study of chromosomal mechanics in D. melanogaster.407 The activities of the fly group included, among other things, breeding

407 Members of Morgan’s laboratory started to branch out into other fields starting in the 1930s, see Kohler 1994. The best historical studies of the Morgan laboratory are Allen 1978 (especially 164ff) and Kohler 1993. The fly group began in 1909 when Morgan offered two then-undergraduates in his introduction to biology seminar jobs in his laboratory. These were Alfred Sturtevant and Calvin Bridges. (Bridges was the younger student and started out simply washing out fly bottles. When Morgan realized how perceptive he was at recognizing mutations, his position was upgraded.) Two years later another undergraduate, H. J. Muller, also joined the group. Muller, Bridges and Sturtevant all finished undergraduate and graduate degrees with Morgan. The group was so productive that by 1915 they published the first comprehensive statement of the chromosome theory of heredity and of chromosome maps, Morgan et al. 1915.

!170 flies, crossing fly strains, and mapping genes (in the physiological form of “mutant types,” such as flies with black bodies, white eyes or no eyes) (Figure 6.1). !

! Figure 6.1: The Mutant Type: “eyeless.” The mutant is characterized by small eyes but in extreme cases may lack one or both eyes entirely. The two upper images (a and a’) are of a normal Drosophila for comparison. (From Morgan et al 1915, 14.) (© Biodiversity Heritage Library, MBLWHOI Library) ! The fly group’s day-to-day procedure went something like this. They would breed flies until a mutant arose in a laboratory strain (such as “white eye”); they would then create a population of flies homozygous for that allele; then, by backcrossing different combinations

!171 of mutant strains with normal—or “wild-type”—flies and counting the progeny, they would SEX-LINKED FACTORS IN DROSOPHILA 49 construct genetic linkage maps: maps that represented the order of and relative distances THE LINEAR ARRANGEMENT OF THE FACTORS between mutant genes on chromosomes (Figure 6.2).408 This research was largely concerned Table 2 shows the proportion of cross-overs in those cases withwhich structure-function have been relations worked (“proximate out. causes”The detailed in recent terminology): results of whatthe genescrosses are physiologicallyinvolved are important, given how at traits the are end passed of from this parent paper. to offspring, The etc. 16287409 cases for B and CO are from Dexter (’12). Inasmuch as C and 0 are Dobzhansky’scompletely early linked role was I largely have in addedmicroscopy: the he numbers had more training for C, than for the other0, and for C and 0 taken together, giving the total results in the lines membersbeginning of Morgan’s (C, 0)lab in P, microscopic B (C, 0), dissection,etc., butand was have much usedless familiar these with figures, recent geneticsinstead literature, of the so thisindividual role was only C, natural. 0, or410 CO The results,Morgan lab’s in researchmy calculations. was, at this The fractions in the column marked ‘proportion of cross-overs’ time,represent very exciting the because number it provided of cross-overs the first simple (numerator)scientific system into which total the avail- able gametes (denominator). mechanisms of heredity could be studied with relative ease.411 As will be explained later, one is more likely to obtain accurate !figures for distances if those distances are short, i.e., if the asso-

0 BC PR M WlD sa7 SSJ 6’1.8 ! Diagram 1 Figure 6.2: The First Linear Linkage Map. It represents the ordering of and relative distances betweenciation a set isof factorsstrong. (or mutant For traits). thi,s The reason map distance I shall, represents in percentage so far oras rate possible, of crossover;use the one percent map unit equalsof cross-overs one percent crossover. between The procedure adjacent assumes points that (i) in the mapping rate of crossoverout the is constant distances along the between length of the the chromosome, various (ii) factors. that rate of crossoverThus, depends B (C, only 0), on distance,(C, 0) (iii) P, there PR, are noand double PM crossovers. form (Fromthe basisSturtevant of 1913,diagram 49.) (© Biodiversity1. The Heritage figures Library,on the Gerstein diagram - University represent of Toronto) calculated distances from B. ! Of course there is no knowing whether or not these distances as drawn When the represent fly group began the their actual work inrelative the 1910s, spacial their fruit distances fly of choice, apart D. of the factors. Thus the distance CP may in reality be shorter melanogasterthan the, neededdistance to be BC,made laboratory-ready.but what we In doorder know to be experimentally is that a breakuseful, it is far more likely to come between C and P than between B and C. 408Hence, For a nice breakdowneither ofCP this processis a seelong Kohler space, 1994, 65 ffor. See else also Morganit is etfor al. 1915.some reason a 409weak Gannett andone. Griesemer The 2004, point 73ff. I wish to make here is that we have no 410means Provine 1981, of 15-6.knowing that the chromosomes are of uniform strength, 411 Compared, for instance, to geneticist William Ernest Castle’s work at Harvard with guinea pigs and rats, in whichand the ifgenetic there systems are were strongmuch more orcomplex. weak See Castleplaces, 1913. Seethen also Allenthat 1978, will 145ff .prevent our diagram from representing actual relative distances-but, I think, will not detract from its value as a diagram. !172 Just how far our theory stands the test is shown by table 3, giving observed per cent of cross-overs, and distances as calcu-

TEE JOURNAL OF ZIXPERIMENTAL ZOBLOQY, VOL. 14, NO. 1 had to be standardized, that is, bred in a way that removed the genetic variation that existed between individuals.412 The fly group focused on creating mutant strains that were, besides the mutant gene, as uniform as possible. Although necessary, the focus on standardization had the effect of jettisoning evolutionary questions.413 The variation that could be acted upon by natural selection and that held the clue to differences between races and full-fledged species outside the laboratory—the variation Dobzhansky had studied in his early work with

Coccinella—was removed from laboratory strains of D. melanogaster.414

The fly group’s work of the 1910s and 20s had thus caused much excitement over mutations and their role in heredity, but it had done so at the expense of evolution.415 It wasn’t because Morgan was uninterested in evolution—evolution may even have been his

412 See Kohler 1994, chapter 3. 413 The relationship between what the Morgan laboratory was doing with Drosophila and traditional evolutionary problems is complicated and is partially addressed in the next section (more complicated, I think, than portrayed by Kohler 1993). Certainly the Morgan laboratory was addressing evolution from an experimentalist perspective, rather than a morphological-naturalist one, and this was apparent to those at the time (see the criticisms outlined in the next section). This obviously does not mean that what they were doing had no bearing whatsoever on evolution—that would be a silly inference given the significance of the genetics approach they designed and advocated with regard to the subsequent trajectory of evolutionary biology. This relationship is complicated partially because Morgan’s relationship with evolutionary theory itself was complicated (see Allen 1978, 106ff). Morgan disliked anything speculative and this included much of evolutionary theory as it was presented by idealist morphologists like Ernst Haeckel and Morgan’s supervisor at Johns Hopkins, W. K. Brooks. He was also skeptical of Darwinian selection as an explanation for evolution. Morgan held this position for a number of different methodological and evidential reasons, summarized in Allen 1978, 109ff. Morgan was, however, always interested in adaptation and evolution (like many others, for a short while between 1900 and 1910 he accepted Hugo de Vries’ Mutation Theory as a superior alternative to Darwinian selection), and it was his interest in evolution and adaptation that caused him to be interested in mutations and that ultimately led to his work on Drosophila (Allen 1978, 125). 414 Naturalists pointed out that the manipulations involved in transforming D. melanogaster into an experimentally useful object ensured that it was quite unlike its natural counterparts. And the naturalists were right. Morgan and many early geneticists assumed that because flies in natural varied little morphologically, they were genetically homogenous as well. Hence the term, “wild-type.” As Dobzhansky would later demonstrate, there was an awful lot of hidden variation within wild populations in the form of recessive alleles. See next section for examples. 415 Kohler 1993, 1994.

!173 reason for starting work with D. melanogaster.416 However, by the time 1911 rolled around and the fly group had found over 15 extreme mutations in D. melanogaster they turned to the readily available, and arguably more important, task of performing experiments for genetic mapping which would preoccupy them for the next 20 years (and win Morgan a Nobel prize).

417The Morgan group’s disregard for evolutionary problems was, furthermore, consistent with a wider American scientific environment that was suspicious of anything deemed

“speculative,” including evolution.418 Dobzhansky’s friend, botanist Ledyard Stebbins, recalled this attitude during his student years at Harvard in the 1920s: “Evolution is a good topic for the Sunday supplements,” a Professor remarked, “but is not science; --you can’t experiment with twenty million years.”419 By moving to America, Dobzhansky had thus

416 When Morgan set out in 1909 to induce mutations in D. melanogaster by altering its physical environment (e.g., heat, cold, radiation) and by artificial selection he was looking for an evolutionary, De Vriesian-style “mutating period” (see Allen 1978, 147-8). Inspired by Hugo De Vries—and contra Darwin—Morgan believed that new species were not created through the selection of small discontinuous mutations, but instead by the accumulation of mutations during natural episodes called “mutating periods,” caused by changes in a species natural environment. See Kohler 1993, 299ff. Morgan caused a significant confusion surrounding the term “mutation.” 417 Op. cit. footnote 407. 418 Morgan himself repeatedly attacked work that he saw as idealistic and speculative. This was especially so after the Drosophila work had become established in the mid-1910s. See Allen 1978, 318ff. The historian Garland Allen traces this attitude back to Morgan’s own revolt from the idealist morphological tradition of Ernst Haeckel and William Keith Brooks (the latter being one of Morgan’s supervisors at Johns Hopkins—see Allen 1978, 35ff). To Morgan, and many others, the work of the idealist morphologist (or worse, the vitalist) was based less on concrete observations of the natural world and more on an order and pattern imposed on nature by the mind of the morphologist. It took a later generation of evolutionary biologists to distance evolution from this kind of idealism and from overly speculative theorizing (Haeckel, we should recall—along with others in the same German tradition according to Morgan—was both idealistic and evolutionary, thus suggesting to many an intimate relation between the two). Although Morgan attacked speculative work, his relationship to evolution is not straightforward. Op. cit. footnote 413. 419 Stebbins Unpublished Essay written in 1942 and sent to Mayr in 1970s; APS. See Gregory 1917. Similarly, in 1922 the famous English geneticist William Bateson wrote that “Less and less was heard about evolution in genetical circles and now the topic is dropped” (Quoted in Gould 1982, xviii).

!174 entered a foreign intellectual atmosphere in which evolutionary studies and field work were not standard practice amongst his geneticist colleagues.420 !

THE STATE OF AMERICAN BIOLOGY: FODDER FOR THE RHETORICAL VALUE OF “NATURAL”

The Morgan lab was representative of one side—the experimental side—of a larger rift in American biology that I flagged above: that occurring between “naturalists” and

“experimentalists.” As many historians have pointed out, throughout the late-19th and early-20th centuries in America and Britain there was a general feeling of uncertainty about how approaches that studied nature on its own terms—such as natural history and taxonomy

—related to approaches which studied nature “constrained” in an experimental laboratory— such as physiology, embryology, or genetics.421 To put this crudely and in a way that is oversimplified—but in language that is consistent with the circulating banter of the time—to the laboratory physiologist or embryologist, natural history was unanalytical, imprecise, and

420 I should add a qualification here pointed out to me by my external examiner Garland Allen. Although this is a true statement as regards the Morgan laboratory circa 1920-30s, that is after Dobzhansky arrived, it is not true that field work and evolutionary questions were not on the minds of (at least some) members of the Morgan laboratory. Morgan had been trained at Johns Hopkins by W. K. Brooks as a traditional field naturalist (he wrote a dissertation on embryology and phylogeny—at the time these categories fell largely under the academic discipline of morphology). See Allen 1978, 46ff. Morgan was also interested in evolution, although for a period of time in the early 1900s he rejected Darwinism and Mendelism (in his dissertation he dealt directly with Haeckel’s biogenetic law, and although he later rejected this on the grounds that it was idealist and speculative, he never lost an interest in evolution—see Allen 1978, 97ff). Alfred Sturtevant too was a qualified naturalist. He wrote a natural history text about the North American species of Drosophila in 1921 (Sturtevant 1921). He also showed his interest in phylogeny and evolutionary problems in the late-1930s when he and Dobzhansky started to plan the “Genetics of Natural Populations Series” together (this became Dobzhansky’s project as their academic and personal relationships fell apart) (see Provine 1981). 421 See for examples Kohler 2002, 43ff.

!175 old-fashioned; to the naturalist, the laboratory simply created phenomena that were too artificial to teach one about natural processes, including most importantly evolution.422

To fully appreciate the rhetorical power and foundational character with which “the natural” is endowed in Dobzhansky’s biological program of the late-1930s, we need to return briefly to the historical sources of its authority. As I said in Chapter 3, rhetoric is effective only insofar as it takes advantage of a common framework for understanding and communicating. This common framework was forged, or at least enforced, in America through debates between naturalists and experimentalists.

While it is certainly true that many naturalists were wary of laboratory results, they were not skeptical of experiments per se—after all, some performed field experiments and many performed natural experiments, the same type of experiments used by Cuvier and

Darwin in Chapter 4. For example, the zoologist Francis B. Sumner extolled the virtues of

“natural” experiments for his studies of mice populations in nature. “I think that I have had enough to do with the experimental [i.e., laboratory] method in zoology to make me realize its rigid limitations,” he wrote. “I am therefore disposed to attach considerable importance to what have been called ‘Nature’s experiments’.”423 The ecologist Frederic Clements as well

422 Of course there were those that tried to combine the two traditions. Such was the basis for many proposals for experimental evolution field stations. Often however these “combinations” also accented the differences by pointing out the shortcomings of a truly laboratory approach Kohler 2002, 44ff.). See Allen 1979 for the definitive piece on naturalists and experimentalists. See also Allen 1994 on Dobzhansky’s role in dissolving this distinction. See William Bateson’s 1914 address to the British Association for the Advancement of Science as well as his reflections in his 1894, Materials for the Study of Variation. See Allen 1979. That naturalists continued to worry about whether their discipline was old-fashioned well into the mid-20th century, see Dobzhansky 1966. Dobzhansky began his 1966 paper with the assertion, “Naturalists study nature.” 423 Sumner 1915, 696. Why Sumner’s application of genetics to evolution in mice was not as influential as Dobzhansky’s is an interesting question but outside of the boundaries of this dissertation. Another example: The paleontologist William Gregory, a colleague of Osborn’s at the American Museum of Natural History, was of a similar opinion. In an article aptly titled “Genetics versus Paleontology,” he argued against the geneticist’s claim that there were no experiments in paleontology. “Nature herself often provides control experiments,” he wrote, as “when she takes a primitive type of skull and dentition and molds them into a wide variety of adaptive types, meanwhile preserving the original pattern as a ‘control’ (Gregory 1917, 627).”30

!176 argued for a “scientific natural history” which would utilize “natural” experiments.424 In these experiments one could gain an understanding of natural ecosystem succession without the disturbing and unpredictable influence of human interference that characterized the laboratory.425

It wasn’t experiments per se that worried the naturalists. Rather, it was human disturbance: that studies which dealt with “disturbed” objects would be misrepresentative of their natural counterparts, or that laboratory studies provided a misrepresentative picture of the evolutionary process. Naturalists expressed concerns that laboratory experimental methods did not teach one about “undisturbed nature,” that is, nonhuman nature. And, in return, experimentalists chided them for their “reverence for natural or ‘normal’ phenomena.”426

Consider one of Dobzhansky’s recollections. When interviewed in 1966 at

Rockefeller University, Dobzhansky drew attention to what he saw as “violent opposition” to

Morgan’s work.: ! Dobzhansky. [...] there was opposition—violent opposition—as you probably know from Henry Fairfield Osborn, who at that time held forth as president of the

424 We saw this type of experiment used by Cuvier and Darwin in Chapter 3. See Kohler 2002 and Kingsland 2005. 425 The paleontologist William Gregory, was of a similar opinion. In an article aptly titled “Genetics versus Paleontology,” he argued against the geneticist William Bateson’s claim that there were no experiments in paleontology. “Nature herself often provides control experiments,” he wrote, as “when she takes a primitive type of skull and dentition and molds them into a wide variety of adaptive types, meanwhile preserving the original pattern as a ‘control’” (Gregory 1917, 627). Jacques Loeb, unsurprisingly, mocked the rigor of natural experiments in paleontology. Paleontologists dated fossils by measuring their geological depths and used this information to compare, as Gregory says, adaptive types to a more primitive control. “Of two animals abroad on the same day,” Loeb mocked, “one might fall into a deeper hole than the other without being a more primitive species.” (Translated in Fleming 1964.) 426 MacDougal 1909, 121; Kohler 2002, 88; see Kingsland 1991. Livingston 1917; quoted in Kohler 2002, 93-4.

!177 American Museum [of Natural History]. Now, Osborn referred to genetics as DeVriesianism. Allen. Oh he did? He outwardly opposed genetics? Dobzhansky. You can read in the American Naturalist, I think, as late as 1930 Osborn published an article where he said that all this Drosophila business is nonsense, and has nothing to do with nature, it's just pathological changes.427 ! The article referred to was published by Henry Fairfield Osborn in 1927 and was a naturalist’s assessment of evolution in nature. Osborn, a colleague of Morgan’s at Columbia and curator of the American Museum of Natural History, argued that the problem of the origin of species “can best be studied under natural conditions of past and present time.”

“Speciation is a normal and continuous process,” he argued, “it governs the greater part of the origin of species; it is apparently always adaptive. Mutation [what the fly group investigated] is an abnormal and irregular mode of origin, which while not infrequently occurring in nature is not essentially an adaptive process; it is, rather, a disturbance of the regular course of speciation.”428 Dobzhansky wrote in 1937 that these statements had been made into “professions of faith by a number of writers.”429

Osborn’s comment, with its strong juxtaposition of mutation and selection, sounds strange to us now, but the idea that mutations were “abnormal,” “pathological,” or

427 1966 interview by Garland Allen at Rockefeller University. Edited by Allen. Square brackets are Allen’s. Dobzhansky continues, “You have instead DeVriesianism, and aristogenesis [his own invention] and of course in good old Harvard University there was a gentleman by the name of [Edward Charles] Jeffrey. Well now Professor Jeffrey, who lived to be quite old—when he died I'm not quite sure [it was 1952]—but he was violently opposed to Morgan and genetics. He believed everything was due to hybridization. He published an article in 1924 or '25 in the American Naturalist, an article in which he claimed that Drosophila was a hybrid—a between what and what was not quite clear—but also the chromosomes are completely unreliable, their number is variable; and while that made some impression some places, most people just laughed at him. He was still active in at least 1930.” Jeffrey’s paper “Polyploidy and the Origin of Species” is also interesting, but to explain it’s criticism of Morgan’s work would bring us away from the current topic (see Jeffrey 1925). 428 Emphasis in first quote mine. Osborn 1927, 40-1. The second quote has emphasis throughout in the original which I have removed. 429 Dobzhansky 1937b, 39.

!178 “disturbances” whose perceived importance was an artifact of laboratory experimentalism, was a commonly held view among those whose work was primarily done in the field.430

Natural selection and mutation were often treated as distinct and competing hypotheses about the origin of evolutionary change.431

The idea that in the laboratory one works with nature’s monstrosities—the

“unnatural”—was a dilemma inherited from before Darwin’s time. There are many examples of biologists framing their concerns in this way. The influential physiologist and friend of

Morgan’s, Jacques Loeb wrote to the physicist Ernst Mach, ! I now have animals that have heads on each end of their bodies; I thus have animals with bilateral symmetry that in nature have different oral and aboral poles. The idea is now hovering before me that man himself can act as a creator even in living nature, forming it eventually according to his will. [...] Biologists label that the production of monstrosities; railroads, telegraphs, and the rest of the achievements of technology of inanimate nature are accordingly monstrosities. In any case they are not produced by nature; man has never encountered them.432 ! Loeb’s “engineering” conception of experimental biology which created “monstrosities” did not fit well with many naturalists who were concerned to understand undisturbed nature.433

Loeb’s comment testifies to the recurrence of this theme among biologists of his generation; to many of them, “monsters” were nothing but an imitation of nature’s mistakes.434

430 See also Kohler’s various presentations of the distinct world of laboratory and field biologists (Kohler 2002a). Osborn allows that mutations, in restricted circumstances, may arise in nature, but argues against their evolutionarily significance. 431 The question, going back to Darwin, was often put, “which was the creative agent?” For an early but nice summary see Provine 1971. 432 Jacques Loeb to Ernst Mach, 26 February 1890; cited in Pauly (1987) 433 See Pauly 1987. 434 See for instance, Pauly 1987, 93ff. This was a common theme in the renaissance as well, see Daston and Park 1998.

!179 In later recollections, Dobzhansky, Ernst Mayr and, Dobzhansky’s close friend, the geneticist Curt Stern, all remembered these kinds of reactions to laboratory work.435 Mayr wrote that Osborn’s comments “well expressed the feelings of the naturalists,” including

Mayr himself at that time.436 Dobzhansky remembered that “To most senior biologists

Drosophila mutations were a collection of monstrosities of no significance for evolution.” “It seems hardly believable now,” he wrote in 1980, “that in the 1920s and even 1930s influential biologists declared ‘Morgan’s mutations’ to be products of abnormal laboratory environments; these mutations did not occur in nature at all.”437 Even as late as 1939,

Dobzhansky prefaces discussions of mutation by addressing these types of concerns.438

The skepticism of many naturalists was, at the time, warranted. What Morgan and the fly group were creating in the laboratory appeared to be incompatible with what was typically found in nature. Early laboratory studies with Drosophila had created two common misconceptions.

First, many naturalists believed that mutations were created by the laboratory’s artificial conditions, rather than simply in the laboratory, because such variation was not to be found in nature. There seemed to be a contradiction between the high variability found in

435 See Stern’s mailed in answers to Mayr evolutionary synthesis questionaire, APS. These are presented below when he speaks about Chetverikov. 436 Keynote address for Evolutionary Synthesis Conference, p. 13, 33, APS. Although Osborn certainly disagreed with Morgan, Dobzhansky’s comments may be slightly overstated, since Osborn helped hire Morgan (and experimental zoologist E. B. Wilson) at Columbia, it would be strange if he thought this research was complete nonsense. 437 Dobzhansky 1980, 449. Mayr wrote to Dobzhansky shortly after the conference, that this reflected “accurately the sentiment encountered by me in Germany in the 1920’s.” (Evolutionary Synthesis - Mayr’s questions answered by Dobzhansky, APS.) 438 “In the minds of some biologists a false impression has been created, as though a very high mutability under experimental conditions coexists in Drosophila with a striking constancy under natural ones. Mutations have been branded ‘monstrosities’, ‘domestication products’ and the like, and their existence in free-living population was doubted or even denied outright.” (Dobzhansky 1939, 340.)

!180 laboratory strains and the lack of variability found in natural populations: it was widely known that when free-living flies caught in the field were examined under the microscope, they were by-and-large morphologically uniform. Morgan named these free-living flies

“wild-type,” and assumed that they were genetically as well as morphologically uniform.

This uniformity was contrasted with the mutations which kept springing up, by some good fortune, in the laboratory. Not even the members of the Morgan lab really knew where the mutations were coming from.439

Second, laboratory studies also suggested that, when they did arise, gene mutations and chromosomal variations (inversions in chromosome segments) were detrimental. Given that mutational changes were apparently random, this was to be expected and was often explained a priori by drawing an analogy with a watch, ! Suppose you prod the innards of a watch at random—bring about some alteration in ignorance of the effect it may have. Are you likely to make it a better-running watch? A change, purely accidental in this sense, wrought in any complicated organization is more likely to injure or wreck than to improve that organization [...] which it subserves.440 ! How was a mutant fly with no eyes or no wings supposed to outcompete its counterparts in the state of nature? Moreover, many mutations seemed to affect viability—so-called lethal genes were very common. Dobzhansky summarized this concern in 1938, !

439 In the 1920s they found that they could be induced by x-rays. 440 Muller 1929, 288. See also Beatty 1994a, .

!181 Indeed, most mutations obtained in the laboratory and found in wild populations are deleterious to the organism. In fact, many of them are lethal. It would seem, then, that the presence of such mutations in nature presages an eventual catastrophe to the organisms rather than a progressive evolution. It is no idle phrase to say that the situation is an extremely challenging one.441 ! To many naturalists, who were well aware of the importance of variation in nature and drew on natural selection as an explanation for species differences, this result seemed contradictory and confusing. If it were true, how could species become progressively adapted? Some assumed, like Osborn, that natural species evolution was a different process from the selection against these mutations. Others, drawing a different conclusion, saw this as proof that the accumulation of mutations and variation led only to degeneration or deterioration of a species, and if left uncontested by natural selection, forecasted the eventual extinction of the species.442

Rather than address these puzzling concerns, the members of experimental genetics laboratories, such as Morgan’s, seemed to ignore them. They were not interested in studying evolution of flies in nature anyway, and thus didn’t need to address them as part of their daily scientific affairs. We will see below how Dobzhansky’s ability to follow Russian literature put him in a unique position to clear these puzzles up.

These two misconceptions provided fodder for the authority of “the natural.” That is,

“the natural” was already a loaded concept—and thus also a useful concept—when

Dobzhansky started to use it repeatedly in later work. When the rhetoric was effective this

441 Dobzhansky 1938, 448. 442 Dobzhansky 1937, 126. Here by selection I have in mind a particular type of variation reducing selection, see Beatty 1987. Dobzhansky always opposed this type of selection. Later he advocated different types of variation maintaining selection, such as selection for heterozygotes over homozygotes.

!182 was because it took advantage of a common framework in which “nature” and “the natural” were already loaded with evaluative content. !

D. PSEUDOOBSCURA, GENETICS AND THE ORIGIN OF SPECIES, AND SORTING OUT THE

MISCONCEPTIONS

A switch of research organism was the precondition for Dobzhansky’s posing and investigating evolutionary problems. Until 1932 Dobzhansky had predominantly worked in

America on D. melanogaster. In 1932, following the lead of his lab mate Alfred Sturtevant, he decided to investigate the genetic basis of hybrid sterility in the hopes that this would provide a clue to species differences. By this time members of the Morgan lab had begun to move beyond classical gene mapping and into other, applied areas of research.443 Sturtevant had been investigating the reproductive sterility between two fruit flies, D. melanogaster and

D. simulans, but this research had stalled for technical reasons: while these two fruit flies did produce hybrid progeny—which is better than many species combinations—the hybrids were all infertile.444

D. pseudoobscura provided the solution. Two races of D. pseudoobscura (Race A and

B) were discovered in the 1920s and these races coexisted throughout the Pacific

Northwest.445 This was geographically convenient because the Morgan lab had migrated in

1928 to Cal Tech in Pasadena. These two races were “physiologically isolated” in that there

443 See Kohler 1994, Part Two. 444 This meant that the most potent technique for genetic analysis could not be used. Thus little could be learned about the genetic basis of hybrid sterility. 445 Later, as a result of Dobzhansky’s research, Race B was given the designation of a separate species and renamed D. persimilis, see Dobzhansky and Epling 1944.

!183 was only partial genetic isolation between them—as opposed to the complete genetic isolation between D. melanogaster and D. simulans. While the hybrid males were sterile, hybrid females were perfectly fertile and could be backcrossed with males from either parent species. This meant that the genetic factors giving rise to the sterility could be investigated in the laboratory. Here was a chance to study the genetic mechanisms that gave rise to sterility between races and thus to investigate the type of isolation that develops between species. In short, here was a chance to provide results which had a direct bearing on evolution;

Dobzhansky gladly took up this study.

While the work on hybrid sterility was important for Dobzhansky, it was collecting D. pseudoobscura in the field in 1933 that is most crucial for our purposes. Although

Dobzhansky had collected wild Coccinella in the field, he had not collected wild Drosophila until the summer of 1933.446 Before that time he had used laboratory strains. What he discovered when collecting wild strains in the mid-1930s was that different strains of the races of D. pseudoobscura showed considerable “concealed genetic variation.” Much more so than he expected. This was a turning point for both Dobzhansky and for genetics.

The study of concealed genetic variation had been pioneered by the Russian geneticist

Sergei Chetverikov and was carried out by his students after his mysterious arrest.447 The discovery that genetic variation was “concealed” provided an answer to the first laboratory- caused misconception given above: it solved the problem of why there was high variability in laboratory strains but low variability in natural populations. Chetverikov argued in 1926 that despite their external uniformity, wild and laboratory Drosophila carry a great wealth of

446 Provine 1981, 32. 447 See Adams 1980 for a summary of Russian genetics and Chetverikov’s fate.

!184 recessive mutations, that is, mutations that result in observable phenotypic alterations (such as flies having no eyes) only when in a homozygous state; when they are in a heterozygous state and paired with a dominant “wild-type” allele, no phenotypic changes will be observable. Since most mutations were recessive, wild flies inspected under the microscope gave no clue as to their existence. As Chetverikov put it, and Dobzhansky repeated on many occasions, populations tend to soak up these recessive mutations “like a sponge” while remaining by-and-large morphologically uniform.

The reason why these concealed mutations tended to show up more often in laboratory strains was that the strains had small population sizes and were thus highly inbred.

In such strains it was only a matter of time before two flies heterozygous for a particular mutant allele mated and produced progeny homozygous for, and thus phenotypically expressing, the mutant allele. By inbreeding D. melanogaster collected from the field and demonstrating that the mutations were found in these flies as well, Chetverikov was able to substantiate the claim that mutations in the laboratory and in nature were the same.448 He concluded, “And thus, we do not have any basis for seeing in the process of origin of mutations the result of the artificial influence of man [...] this process goes on just as regularly under natural conditions.”449 This research was carried on throughout the 1930s by

Chetverikov’s students (particularly Nikolay Dubinin and Nikolay Timofeev-Ressovsky) in

Russia and Germany. While Dobzhansky followed this research, many of his American peers

448 A simple summary can be found in Dobzhansky 1960. 449 Chetverikov 1926, 174.

!185 could not because it was often not published in english. Until Dobzhansky’s popularization of this research program in 1937, many in America had not heard of Chetverikov.450

This concealed genetic variation in Russian D. melanogaster piqued Dobzhansky’s interest. His finding variation in natural populations of D. pseudoobscura in the summer of

1933 only increased his fascination. Of this time period, he writes, ! In the early thirties, I was busy with chromosomal aberrations. In the middle thirties, however, this matter of concealed genetic variability, what at present is called genetic load, began to look more and more interesting, especially in connection with the [sic] evolutionary problems. Really, it was the Jessup Lectures and the book resulting from them which helped to to [sic] direct my attention to that as a field of investigation.451 ! Dobzhansky’s move to D. pseudoobscura and to the investigation of concealed genetic variation was, however, more complicated than he makes it sound, particularly from a social-political standpoint. He recognized that, at least from the point of view of the

Morgan lab, he was heading against the grain of current research questions and organisms.452

His decision to study wild and genetically messy populations of D. pseudoobscura was seen, in Dobzhansky’s own eyes as well as many of his colleagues’, as a throwback to older days; evolutionary problems that had occupied the less critical naturalists of the 19th century, not the new rigorous experimental crowd.453 He wrote to his friend, the German geneticist Curt

450 His paper was not translated until 1961, two years after his death. 451 Dobzhansky 1962a, 410-11. 452 He may have upset the Morgan’s group “moral economy” that was so productive for the twenty years prior (Kohler 1994). Initially Sturtevant seems to have shared this interest with him but they had a infamous falling out in the late 1930s. The history of their relationship has been analyzed extensively by Provine 1981 and Kohler 1994. 453 Field research only became old-fashioned with the rise of experimentalism and laboratories in biology in general throughout the last half of the 19th-century (Kohler 2002b, 3ff). See also De Chadarevian 1996 this changing environment in the case of Charles Darwin’s versus Julius Sachs experiments.

!186 Stern, in 1936, “I have definitely left the track of the classical Drosophilaforschung, and becoming [sic] more and more engaged in subjects that are somewhat abhorrent (or at least not very interesting) for the drosophila fellows here [in Morgan’s laboratory].”454 And in the following year, “it seems I got firmly on this field of ‘genetics of populations’, which some of our colleagues consider old-fashioned, not enough advanced, and akin to such terrible things as taxonomy and morphology.” But, he continued, “I happened to be interested in just this old fashioned stuff, and perhaps after all it will give some bits of information interesting even for the modern biologists.”455

Dobzhansky felt comfortable confiding in Stern because he knew he was preaching to the choir. Stern replied, “I am glad that your research goes along evolution lines. You are exceptionally well equipped for this work and you know that I do not share the horror of taxonomy and morphology. ‘Genetics of Populations’ seems to me one of the few really fruitful fields left to the pure Geneticist and thus should be regarded quite ‘modern’.”456

Stern, it turns out, was right. Despite Dobzhansky’s initial uneasiness, his research on

D. pseudoobscura and evolution brought him quick fame. Although unforeseen by

Dobzhansky, it seems biologists of many traditions were ready to engage his work on evolution, whether they agreed with his conclusions or not. He wrote to the geneticist L. C.

Dunn in late 1938, just over a year after expressing his concerns to Stern, “At present there is nobody here [in Morgan’s laboratory] working on melanogaster - a situation which would be unthinkable” only a short while before.457

454 Theodosius Dobzhansky to Curt Stern, 26 February 1936, APS. 455 Theodosius Dobzhansky to Curt Stern, 16 September 1937, APS. 456 Curt Stern to Theodosius Dobzhansky, 29 October 1937, APS. 457 Theodosius Dobzhansky to L. C. Dunn, late 1938, APS.

!187 As many have pointed out, Dobzhansky’s work with wild D. pseudoobscura was especially successful at getting naturalists to see the worth of experimental genetics. To some historians this is what makes Dobzhansky the instigator of the evolutionary synthesis—he helped merge the field of evolutionary biology, principally relegated to naturalists, with genetics.458

Mayr—a naturalist by training—wrote to Dobzhansky late in life about the first time he read Dobzhansky’s papers: Mayr exclaimed, “Finally a geneticist who talks sense!”459

Mayr thought that Dobzhansky “talked sense” because his work on natural populations proved that he had an acquaintance with “nature”—something Mayr saw as necessary for the study of evolution, yet lacking amongst geneticists.460 The early reviews of Dobzhansky’s seminal Genetics and the Origin of Species (1937), demonstrate that Mayr’s reaction was widespread. One reviewer wrote, ! we feel that Professor Dobzhansky’s introduction to the field of general biology by way of his early experiences in entomology has given him his interest in genetics in the field, which has put the higher values in this work. It has given him an interest in species as they actually exist in nature rather than an interest in theories of speciation as based on laboratory techniques. The laboratory in his hands plays its proper part in checking natural events.461 !

458 See Gould 1982, xxi. 459 Ernst Mayr to Theodosius Dobzhansky, 25 February 1974, APS. 460 Similarly, the naturalist and paleontologist George G. Simpson, wrote that Dobzhansky “profoundly changed my whole outlook and started me thinking more definitely along the lines of explanatory (causal) synthesis and less exclusively along lines more nearly traditional in paleontology.” (Simpson’s answers to Mayr’s questionnaire, [1970], APS.) 461 Anonymous review, December 1937, Annals of the Entomological Society of America, APS. My emphasis.

!188 Dobzhansky’s Genetics and the Origin of Species, as he said above, began as the

Jessup Lectures at Columbia in 1936 and was published as a book in 1937. This book provided a synthesis of research on genetics with that on evolution. It offered a research program that set genetic variation in natural populations at the center. Dobzhansky provided a novel explanation for the origin of species, that is perhaps best summarized in his own words, ! Race formation begins with the frequency with which a certain gene or genes becomes slightly different in one part of a population from what it is in other parts. If the differentiation is allowed to proceed unimpeded, most or all of the individuals of one race may come to possess certain genes which those of the other race do not. Finally, mechanisms preventing the interbreeding of races may develop, splitting what used to be a single collective genotype into two or more separate ones. When such mechanisms have developed and the prevention of interbreeding is more or less complete, we are dealing with separate species.462 ! Following from this understanding of speciation was a research program which had two major goals. One was to describe variation in natural populations. Dobzhansky had only just begun his research on D. pseudoobscura at this time and had to largely rely on Chetverikov’s student, Dubinin’s, data on D. melanogaster. The second goal was an analysis or explanation of this variation.463 Using the mathematical theories of evolution offered by J. B. S. Haldane,

Ronald Fisher, and Sewall Wright, Dobzhansky projected that one could determine which evolutionary agents (natural selection, mutation, migration, or genetic drift) had led to the

462 Dobzhansky 1937, 62-3. 463 A third goal was the analysis of isolating mechanisms that prevented interbreeding between races, but I will not deal with this research here.

!189 current patterns of genetic variation among populations. One could thus determine which agents had been important for the origin of species. Without playing down the novelty of

Dobzhansky’s contribution, it is safe to say that this program in evolutionary genetics was strongly influenced by his Russian counterparts who were little known in America.

The third and forth chapters of Dobzhansky’s book dealt directly with solving the first laboratory-caused misconception. First he drew attention to the problem:

Variation-producing agents (gene mutation, chromosomal changes) are known from laboratory experiments. But it is not a priori certain that these agents observed under laboratory conditions have been effective in nature and are responsible for the organic diversity empirically observable out-of-doors. We assume this to be the case because no other equally satisfactory working hypothesis has been proposed. Nevertheless, the validity of the working hypothesis must be rigorously tested by examining whether the differences between forms encountered in nature can be resolved into the elements whose origin is known in experiments.464 ! Relying heavily on the research and procedures of the Chetverikov school, Dobzhansky demonstrated that the genetic variation found in the laboratory was consistent with that found in nature.

Dobzhansky also offered a solution to the second misconception arising from laboratory studies: that genetic variation was always detrimental and consequently that variation was ultimately bad. Dobzhansky’s answer, one that he would hold for the rest of his career, was that laboratory studies created the misconception by investigating flies only under

464 Dobzhansky 1937, 14.

!190 stable environments.465 Under this environment, only the detrimental side of variation is observed. Under a changing environment, however, variation alone provides the genetic plasticity needed to avoid extinction. Dobzhansky writes, ! Looked at from another angle, the accumulation of germinal changes in the population genotypes is, in the long run, a necessity if the species is to preserve its evolutionary plasticity. The process of adaptation can be understood only as a continuous series of conflicts between the organism and its environment. The environment is in a constant state of flux, and its changes, whether slow or catastrophic, make the genotypes of the past generations no longer fit for survival. [...] A genotypical change means, however, the occurrence of a mutation or of mutations. [...] Mutations are random changes. Hence the necessity for the species to possess at all times a store of concealed, potential, variability. [...] It has already been pointed out (Chapter II) that mutational changes that are unfavorable under a given set of conditions may be desirable in a changed environment. Mutations that decrease viability when taken separately may have the opposite effect when combined.466 ! He connected this misconception even more directly to laboratory studies less than a year later: “All that we really know about mutations encountered in wild populations of

Drosophila is that in the environment in which the flies have been kept in our experiments a large majority of the mutations are deleterious. There is no basis for the assertion that these mutations will be deleterious in every possible environment.”467

Such misconceptions about the evolutionary process were, Dobzhansky believed, caused, at least in part, by laboratory studies that went unchecked by studies in nature.

465 Dobzhansky showed his apprehension when he called this an “attempt” to “indicate one of the possible escapes from the apparent impasse” (1938, 448). 466 Dobzhansky 1937, 127. 467 Dobzhansky 1938, 449. My emphasis.

!191 According to Dobzhansky, in these cases, laboratory studies provided a misrepresentative picture of the nature and consequences of genetic variation. !

THE “NATURAL” EXPERIMENTAL D. PSEUDOOBSCURA

Concepts of the artificial and the natural came to play a much more authoritative and foundational role in Dobzhansky’s thought and practice in the period after he switched to full time studies on D. pseudoobscura and began the “Genetics of Natural Populations” series.

This series set out to describe and analyze the genetic variation in natural populations of D. pseudoobscura in compliance with the program set forth in Genetics and the Origin of

Species. As he positioned his research in relation to that of others, he often exhibited and exploited what he took to be the “naturalness” of D. pseudoobscura.

In 1974, Mayr sent out historical questionnaires to the geneticists of the evolutionary synthesis period, requesting that they summarize their major contributions. When

Dobzhansky replied, the very first thing he drew attention to was his work with fruit flies in nature. Summarizing some of what we have already considered, he writes, ! Drosophila genetics was done only in laboratory bottles until Tshetverikov [Sergei Chetverikov], Timofeeff [Nikolay Timofeev-Ressovsky], [Nikolay] Dubinin, and myself started to do genetics in nature, of course by collecting flies in nature and studying their progenies in laboratory bottles! Surely it is no coincidence that all four got their education in one place [...] when I decided in 1931 to take up studying the “races” A and B of D. pseudoobscura this was met with dead scepticisum [sic]. Should one not stick to D. melanogaster with its abundant chromosome markers,

!192 chromosome maps, and trick chromosomes? Are these “races” so different from “races” which one can build in melanogaster, e.g., by irradiation?”468 ! Dobzhansky here identifies with Russian Drosophila geneticists, which broke with tradition and worked on genetics “in nature.” Dobzhansky often used Dubinin’s work as a useful contrast to his own; Dubinin’s research was similar to Dobzhansky’s in that he studied fruit flies in the wild using similar techniques, but Dubinin continued to work on D. melanogaster, whereas Dobzhansky switched to D. pseudoobscura.

When Dobzhansky first began this work, he wrote to Stern and insisted that although his work is “analogous to Dubinin’s data on melanogaster,” it was perhaps “worth while to have data on a ‘natural’ species like pseudoobscura, and on mountains rather than on garbage cans and [in] store rooms.”469 It is worth mentioning—and this is often overlooked—that

Dubinin’s data, as Dobzhansky was well-aware, were about wild populations of D. melanogaster in Southern Russia. That is, Dubinin’s experiments did not only involve the standardized D. melanogaster of the Morgan laboratory but ones free-living in populations outside of the laboratory. What, then, was so unnatural about them?470

Let us start with the way in which Dobzhansky believed his own studies to be

“natural.” Although Dobzhansky’s collection of D. pseudoobscura took place in the field, these flies underwent extensive laboratory manipulation.471 What mattered to Dobzhansky, however, was that the results found were tracking changes in genetic variation due to natural

468 Theodosius Dobzhansky to Ernst Mayr, 15 December 1970; APS. 469 Theodosius Dobzhansky to Curt Stern, 16 September 1937; APS. Dobzhansky consistently and in a wide variety of contexts referred to D. melanogaster as a “garbage” fly. See the notes 470 Why was data on a more “natural” species worthwhile? 471 See Gannett and Griesemer 2004 for continuities between Dobzhansky’s later experimental practice and the classical genetics done in Morgan’s laboratory.

!193 processes, rather than human-mediated processes. “If a stock was established, even for a single generation,” Dobzhansky worried, “the results might reflect laboratory-mediated events.”472 Of course, such a suggestion is intimately tied to what one considers “natural”: the mountains are more natural, than the store room, for instance.

This stance is made quite explicit in Dobzhansky’s published work of the late

1930s.473 Allow me to paraphrase a long argument made in a 1939 review paper.474 In it

Dobzhansky compares D. melanogaster to D. pseudoobscura (again contrasting his work with Dubinin’s) in an attempt to demonstrate the advantages of the latter species for studies of evolution, thus also securing the important status of his own research. He begins by considering D. melanogaster. He tells us it is a “cosmopolitan” fly, a “scavenger,” never found far from the human habitations, orchards, and gardens on which it depends. This is true in both Russia and America. It’s not even clear to which part of the world it is indigenous.

Thus, he concludes, “All the genetically analysed population samples of D. melanogaster are, then, neither exactly ‘wild’ nor ‘free-living’.” This intentionally refers to Dubinin’s “wild” D. melanogaster, and suggests that, based on geographical origins, D. pseudoobscura is more natural than D. melanogaster (and some other “trashy” flies).

He continues, “To some writers,” and think here of the naturalists in the last section,

“the word ‘domestication’ has become a kind of scarecrow.” It has been used to “cast aspersions on the validity of the resulting data for an understanding of the evolutionary process.” Dobzhansky believed such a general view is untenable, for “domestication is

472 Hampton Carson to William Provine, 19 February 1980; quoted in Provine 1980, 52. 473 See, for instance, Dobzhansky 1939, Dobzhansky and Epling 1944. 474 Dobzhansky 1939, 345-6.

!194 merely a special case of ‘natural’ conditions.” However, “domestication does modify in some ways the balance of forces acting upon the genetic composition of a population” and hence cannot be ignored, even in an animal “so little ‘domesticated’ as D. melanogaster.” In this paper, Dobzhansky is reviewing research on concealed genetic variation in nature, the ultimate aim of which is again to demonstrate that variation is not just created in the laboratory under unnatural conditions, but is extremely widespread in nature as well.475 The drawback, he indicates, of work done on D. melanogaster is that the variation detected might be an artifact (in the most literal sense of the word) of hanging around humans, and will thus not be representative of “natural” variation. Even Dubinin’s “wild” D. melanogaster are not immune to this worry, because when it comes down to it they are not really “natural” at all.476

This argument represents a slight but significant change of opinion. In 1936, when he was writing the Jessup lectures, Dobzhansky could only offer Dubinin’s data as evidence.477

But by 1939, after three years of research on D. pseudoobscura, he had become skeptical of

Dubinin’s conclusions, and this skepticism was grounded in the unnaturalness of D.

475 This we now take for granted largely as a result of Dobzhansky’s studies. See Dobzhansky and Queal, 1938a, 1938b, for early discussions of variability. 476 It is worth noting that similar types of worries extend to the study of D. pseudoobscura with regard to the length of time wild populations should spend in the laboratory. To determine the geographic distribution of different gene arrangements in D. pseudoobscura along the Pacific coast of North America, Dobzhansky and his co-author Alfred Sturtevant analyzed chromosomal inversions in laboratory strains whose wild ancestors were collected from different locations (Dobzhansky and Sturtevant 1938). Each of these laboratory strains were derived from a single wild female. The fact that these strains had spent a long time in the laboratory created a concern, however, about the precision of the results. This was because “the variety of gene arrangements originally present in a given strain may be decreased if this strain is kept in the laboratory for many generations” (Ibid., 30; see also Gannett and Griesemer 2004, 72.). The results of the experiments, in other words, might represent the strain’s time spent in the laboratory, rather than the natural variation created by a natural process. They go on to describe a “different method” which does not suffer from the problem of having flies stored in the laboratory before they are experimentally studied. 477 As he later wrote, “In the first edition of [Genetics and the Origin of Species], I was able to quote only observations of Dubinin, as indication, as proof that natural populations of Drosophila do carry mutant genes which can serve presumably as raw materials for evolutionary change. The amount of this kind of information at that time was quite small.”

!195 melanogaster. Dubinin had concluded that genetic variability fluctuated randomly in populations of D. melanogaster from year to year. Dobzhansky, however, thought D. melanogaster provided a misrepresentative view of genetic variation.478 By 1944 he was habitually advocating the use of D. pseudoobscura over D. melanogaster, because the genetics of the latter were too often “churned up” by man.479

Dobzhansky concluded his 1939 argument by outlining the relative advantages of D. pseudoobscura: it’s found in places where humans rarely reside; it seldom invades “human dwellings” or becomes a scavenger. Its indigenous territory is known and its spread by man has been rare (if completely absent). For these reasons, it is implied, D. pseudoobscura is a more appropriate object of evolutionary study.

This is an important point and worth belaboring. Some have argued that

Dobzhansky’s studies of D. pseudoobscura were necessary because the laboratory populations of D. melanogaster were too standardized. Here we see an additional point—a further limitation of D. melanogaster. He is not just skeptical of standardized D. melanogaster flies that have been maintained in the laboratory, he’s skeptical of D. melanogaster in general, as a species that hangs around humans and has done so throughout its evolutionary history. Wild D. melanogaster populations are unsuitable because they are

478 Dobzhansky had the same argument against Sturtevant’s 1931 study of chromosomal inversion variation in D. melanogaster. See Dobzhansky 1939, 358. As he wrote in the 2nd edition of Genetics and the Origin of Species (1941), “D. melanogaster is not native to Russia, and is probably unable to survive the rigors of winter, except for a small number of individuals which may find hiding places in or near human dwellings. As in many originally tropical species which had become human commensals or parasites, its populations are bound to be occasionally reduced to a few survivors and to be shifted from place to place by human agencies” (Dobzhansky 1941, 181). 479 Dobzhansky and Epling 1944, 3. They write, “As laboratory animals and in the clarity of their chromosomes these species are not much inferior to D. melanogaster, and from the standpoint of analysis of natural populations they are superior in certain important respects. D. melanogaster is at present nearly cosmopolitan and, at least in the temperate zone, is closely associated with man. Its populations are constantly churned up because of unintentional transport with man.”

!196 not really “wild” at all: their evolutionary history involves too much human disturbance.

Even with these free-living flies, he is worried that the variation detected might be an artifact of human interaction. It is for these latter reasons that he claims the value of D. melanogaster is limited. Its evolutionary history is, in a sense, tainted.

Hampton Carson’s claim that Dobzhansky was determined to perform experiments that reflected exactly what natural selection in nature had wrought is corroborated by the above, but needs to be amended (or clarified). This “in nature” is not simply outside of laboratory bottles, but outside of human interference altogether. Dubinin’s analysis of wild D. melanogaster suffers from the same problem Morgan’s domesticated D. melanogaster. D. pseudoobscura, on the other hand, does not suffer from this drawback because it is “shy” and lives away from humans. D. pseudoobscura is a preferable organism for the study of evolution because its genetics carry the stamp of evolutionary changes wrought in nature, not by man.480

As a more general reflection, for Dobzhansky, we might further conclude that

“nature” is not a physical place, but an ideal historical state; “natural” does not refer to an area of origin per se, but to the genetic history of the flies. D. pseudoobscura is “natural” because its evolutionary history, imprinted in its genetics, involves little or no human disturbance. This understanding of “natural” played a role in structuring Dobzhansky’s

480 A related discussion occurs in Dobzhansky’s thoughts about natural selection. For instance, in the Reminiscences, he writes about Genetics of Natural Populations paper XII, “Here you had a chance at long last, that this phenomenon of natural selection, about which people were talking at great length, writing at no less great length, could be experimented with. You actually could reproduce in the laboratory something which went on in nature. And of course, by the same token, it showed that what does happen in nature is an evolutionary change. So, you are observing evolution. It happens before your eyes” (Reminiscences, 497-8). Dobzhansky is implicitly discounting studies of artificial selection, which didn’t count—we can surmise—because the “selection” was done by humans.

!197 collecting practices because it told him where he could find “the natural.” Although “the natural” is not a physical place, this ideal helped him decide what physical places were appropriate. For example, when a friend asked for his advice, he responded, “It is advisable to avoid collecting in orchards or too close to human habitations because in such places the

Drosophila populations consist chiefly of cosmopolitan introduced species (Drosophila melanogaster, D. simulans, D. hydei, D. ananassae) which are not interesting for our purposes. Such ‘garbage’ species are relatively rare in woods or other ‘natural’ habitats.”481

The historical “naturalness” of D. pseudoobscura was important not only for describing concealed genetic variability but also for analyzing it—the other half of

Dobzhansky’s research program set forth in Genetics and the Origin of Species.482

For example, in a 1938 paper, Dobzhansky and Marion Queal analyzed the genetic composition of a number of isolated populations of D. pseudoobscura inhabiting “island- like” mountain forests in the Death Valley region of California and Nevada.483 What they found was that these populations were genetically unique.484 The question, they concluded, was, “What are, then, the driving causes of the historical process which has resulted in the

481 Theodosius Dobzhansky to J. George Harrar, 20 April 1950; APS. Harrar was a plant plant physiologist and president of the Rockefeller foundation (1961-1972). 482 This goal was perhaps best summarized by one of Chetverikov’s students Nikolay Timofeeff-Ressovsky in 1940. The evolutionary, mathematical foundation provided by Haldane, Fisher and Wright, demonstrate “the relative efficacy of various evolutionary factors under different conditions possible within the populations (Wright 1932). It does not, however, tell us anything about the real conditions in nature, or the actual empirical values of the coefficients of mutation, selection, or isolation. It is the task of the immediate future to discover the magnitude of the coefficients in free-living populations of different plants and animals; this should form the aim and content of an empirical population genetics” (Timofeeff-Ressovsky 1940, 104). 483 Dobzhansky and Queal 1938a. This was one of the first papers to work on natural populations of fruit flies— the subject had been “hitherto almost untouched,” they write—and thus they needed to justify their long descriptions of each sample’s geographic location: “The nature and source of the material are so important in the present investigation that we are forced to consider them more carefully than is customary in genetic accounts” (Ibid., 239). 484 Gene arrangements of individuals are determined by inspecting the giant salivary chromosomes.

!198 differentiation of the populations inhabiting separate mountain ranges?”485 They discounted migration and mutation as driving causes, and considered random genetic drift more likely than natural selection. Although they could offer no definite answer, D. pseudoobscura was a good model for asking and answering these types of questions, they argued, because its populations were “as yet practically undisturbed by man’s activities.”486 One did not have to worry that the “driving cause” might be human activity, rather than a natural process associated with evolution.487 Thus making D. pseudoobscura an ideal object for analyses of evolutionary problems.488

Dobzhansky’s intention to capture evolution “in nature” did not make him anti- experimental, or even what we might consider a true naturalist.489 It is surprising how little he knew of the flies’ ecology and natural history, knowledge you might expect from a naturalist who spent time observing them in nature. He was amazed at finding out in the

1940s what the fly ate, and was similarly surprised that it had “a biology of any kind” which lab studies seemed to preclude.490 It wasn’t until 1951 that he discovered that two races of D. pseudoobscura mated at different times of the day, thus providing a reason why they didn’t

485 Ibid., 250. 486 Ibid., 239. 487 Dobzhansky later used further field observations along with experimental studies of fruit flies in “population cages” to corroborate the hypothesis that natural selection caused cyclical changes in these isolated mountain populations (Wright and Dobzhansky 1946). What Dobzhansky thought he could and couldn’t learn by studying fruit flies in population cages in the laboratory adds another dimension to this story, but it will not be dealt with here as it would bring the discussion to far afield of the current point. The artificial studies using population cages were again used to check natural events and would lose all meaning for Dobzhansky in the absence of prior field studies. For a great discussion see Rudge 2000. 488 Dobzhansky and Epling 1944, 3. 489 As one student, Richard Lewontin, recalled, “It was explicitly not part of his plan to give ecological and physiological explanations of this superiority [of heterozygotes] except as an armchair exercise once in a while. No part of his experimental program or, indeed, of what he tried to get people interested in doing, involved real experimental work in the field or real attempts to measure physiological properties that would serve fitness” (Lewontin 2001, 31). 490 Theodosius Dobzhansky to L. C. Dunn, 28 July 1942; Theodosius Dobzhansky to L. C. Dunn, 9 August 1941; Reminiscences, 338; APS

!199 interbreed in nature.491 And when Dobzhansky spent a summer at the Carnegie Institutes field camp in the California Sierra, his true laboratory colors showed through. “Dobzhansky is just emerging out of the milk bottle (T. H. Morgan) stage of Drosophila genetics,” one field biologist remarked. Another agreed, “he is regarded as a bit of a radical among the conventional geneticists [... yet] his outlook is still quite conventional as compared with the

[field biologists here].”492

Furthermore, although Dobzhansky may have been concerned with “the natural” in a way that his immediate predecessors perhaps were not, his experimental practice was fundamentally similar to theirs, and manipulative laboratory work continued to play a dominant and necessary role throughout his career. This manipulation was necessary since, as

I have said, the genetic variation he studied was largely recessive and deleterious for viability, thus this variation was rarely manifested in nature on a phenotypic level. A morphological inspection of wild flies would thus fail to detect the underlying diversity as it had in the past.493 The variation could only be brought out through extensive laboratory manipulation.494

The wild flies were, on their own, experimentally useless. What mattered was their concealed genetic variation, and to observe it required extensive laboratory preparation.

491 Theodosius Dobzhansky to L. C. Dunn, 10 August 1951; APS. 492 Ecologists at the Carnegie Institute. See Kohler 1994, 282. 493 See Dobzhansky 1939, 340. 494 Here is one standard procedure: (1) “Wild males, or single sons of wild females, are crossed to females homozygous for the third-chromosome recessives orange and purple.” (Wild flies, now in lab, crossed with a standardized laboratory strain.) (2) “In the F1 a single male is selected and outcrossed to females carrying orange, Blade, Scute, and purple.” (The wild-laboratory hybrids are further crossed with laboratory strains. (3) “In the next generation (F2) females and males showing Blade and Scute are selected and inbred” (Next generation flies are selected and inbred.) (4) “In the offspring (F3) two classes of flies appear, namely, wild type and Blade Scute.” (These are counted and ratios between mutants determined. From this Dobzhansky estimates the genetic variability—in terms of lethal genes, semi-lethal genes, or genes causing external effects—of the original sample and can compare this to other regions.) See Dobzhansky 1939, 346.

!200 Dobzhansky became familiar with this technical preparation in Morgan’s laboratory working on D. melanogaster, and continued to use fundamentally similar methods with D. pseudoobscura. The laboratory techniques had not in fact changed significantly from the

“artificial” tradition Dobzhansky saw himself as breaking away from. However, with D. pseudoobscura, the variation found would represent the result of processes of which humans played no role.

Dobzhansky’s admiration for the wild D. pseudoobscura went over and above its scientific suitability, and took on a variety of moralized tones. He referred derisively to a colleague’s work as being based on merely “laboratory strains!” He spoke of the flies’

“perfect taste” for natural beauty and wild nature.495 On his 70th birthday a number of his colleagues dedicated papers to him and to one he responded, “It is splendid work, despite being done with that species less noble than Drosophila pseudoobscura, and it is very closely related to what my colleague [...] and myself were trying to do [...] with that nobler species.”496 He even referred to laboratory grown females as not possessing “as high a level of morals” as wild females, because of higher rates of outbreeding in the laboratory.497 These remarks are whimsical, but I think their consistency—like the “friendly” banter between experimentalists and naturalists—expresses a deep-seated preference for a particular kind of study.

495 Dobzhansky wrote to one set of friends that a trip to British Columbia was “most pleasant but scientifically [...] next to sterile”, for D. pseudoobscura could not be found in the mountains. “It is the first time,” he wrote, “that Drosophila pseudoobscura proved to have a less than perfect taste.” 496 Theodosius Dobzhansky to John M. Thoday, 26 February 1970; APS. My emphasis. 497 Theodosius Dobzhansky to L. C. Dunn, 29 July 1951; APS.

!201 A concern to represent what nature had wrought also made Dobzhansky friends among naturalists. For example, when the paleontologist (and evolutionary synthesis architect) George G. Simpson’s research risked losing financial support, he reached out to

Dobzhansky for help. “I had a talk with Dr. Hanson, of the Rockefeller Foundation, in the hope that his organization might support the proposed research on variation and speciation,”

Simpson wrote. “He was very dubious about this, especially on the grounds that the study does not in fact have much bearing on genetics or the problems of experimental biology. I felt, on the contrary, that it has a very crucial bearing since it is one approach to the problem of what actually happens in nature to the raw materials of evolution revealed by genetic and other experimental studies.” And furthermore, “[t]his is certainly necessary to give real meaning to such studies and to coordinate what is known of evolutionary problems from the two approaches.”498 Presumably, Simpson understood that Dobzhansky, despite being a geneticist, would see the necessity of studies which demonstrated “what actually happens in nature” to all that revealed by experimental biologists.

In addition, it only helped Dobzhansky’s case that he adopted (intentionally or not) a

“wild” persona as someone who frequented the mountains and traveled to South American jungles on collection trips.499 Many joked that Dobzhansky worked on D. pseudoobscura simply to get into wild nature, !

498 G. G. Simpson to Theodosius Dobzhansky, 21 January 1941; APS. My emphasis. 499 Although there is no place to pursue this here, it is interesting as well that it takes a kind of “wild” person to collect D. pseudoobscura. He also traveled widely throughout his life and did extensive research in South America, writing travel letters of his wild adventures back to his friends (see the published letter in Bentley Glass’ The Roving Naturalist 1980). His reputation as a wild man probably only increased his credibility among naturalists.

!202 Since my wife and myself were both addicted to excursions, camping trips and so on, there was our chance, and more than one friend and colleague since have suspected that I started to work on Drosophila pseudoobscura and Drosophila persimilis as a thin rationalization of this desire to travel!! The story was going around among geneticists that for rather obscure reasons, Drosophila pseudoobscura occurs in the United States chiefly in natural parks and in Mexico, chiefly near the ancient ruins.500 ! Dobzhansky often referred to himself in this way, or spoke of his research in the “Wild

West,” and he was thought of by many as more vehement than the average geneticist

(including with regard to his temper!) (Figure 6.3).501 !

500 Dobzhansky 1962a, 368. 501 Dobzhansky to Sewall Wright, 12 May 1939, APS. Dobzhansky to L. C. Dunn, 15 June 1941, APS. Dobzhansky to L. C. Dunn, 8 July 1942, APS. On Dobzhansky’s temper see, for example, Brito da Cunha 1998. Kohler 1994, 288.

!203 !

!

! Figure 6.3: A Postcard from Dobzhansky to L. C. Dunn. Dobzhansky’s collecting trips were anything but cosmopolitan. The postcard to L. C. Dunn—and friend, co-author and colleague at Columbia— depicts the view from one such collecting trip. The inscription reads: “Dear Dunn: I hope the view on the other side evokes some pleasant memories. We visited the place yesterday.” (© American Philosophical Society Library, Philadelphia) ! As a parenthetical but related final consideration, Dobzhansky seems to have been aware of how hypersensitive naturalists were to “the artificial.” The synthesis Dobzhansky

!204 helped bring about in biology was nicknamed the synthetic theory of evolution. Dobzhansky however was never satisfied with that name. He preferred to call it the “biological theory of evolution.”502 Why did he prefer “biological” over “synthetic”? He answered, “This theory has also been named ‘synthetic.’ It is synthetic, in the sense that it embodies a synthesis of data from biology as a whole. The word "synthetic" may, however, also mean artificial or factitious, as contrasted with genuine, and this makes the designation "biological" preferable in my opinion.”503 He was worried that people would understand “synthetic” to mean

“artificial.” As we have seen above, this was not exactly what Dobzhansky intended in synthesizing evolution and laboratory genetics through his studies of D. pseudoobscura. He certainly wanted things to remain “natural.” !

THE POLITICAL CONTEXT OF NATURAL AND ARTIFICIAL POPULATIONS ! The trouble with Muller’s [eugenical] program is that, as you correctly say and Huxley denies, it is based on a confusion of a purely biological sort, as well as on a confusion of biology and sociology. He confuses fitness with absence of genetic load, and next confuses biological fitness with human desirability. These two confusions and in fact each one of them separately, would be enough to vitiate his program. I am not surprised that Huxley fails to see this - his own understanding of genetics has always been not as good as he thought it was Dobzhansky to L. C. Dunn, 6 Aug 1961 ! The idea that biologists who spend too much time in the laboratory don’t have enough of an appreciation for “nature” to pronounce on evolutionary matters persisted throughout

502 Dobzhansky’s Reminiscences (398), the Columbia Oral History Project. See also similar remarks in Theodosius Dobzhansky to Everett Mendelsohn, 2 December 1969 and Theodosius Dobzhansky to G. L. Stebbins, 11 September 1973; all at APS. 503 Dobzhansky 1965b, 207.

!205 Dobzhansky’s lifetime. This was, for Dobzhansky and others, both politically useful and unnerving. Dobzhansky, for example, wrote to Ernst Mayr in 1974, “I also agree with you that [geneticist Hermann J. Muller] was ‘rather naive’ in many of his pronouncements on evolution,” but this was to be expected, “from a man of great laboratory achievements but no familiarity with organisms as they live outside.”504 In other words, to say anything informed about the natural process of evolution, one needs some acquaintance with nature.505 (I am reminded, in this exchange, of Wallace’s argument that the Duke of Argyll had little direct experience with wild nature.)506

However, it was not just that Muller’s view of evolution was misrepresentative,

Dobzhansky worried that it was undesirable and politically dangerous. The argument of this section is that we cannot separate Dobzhansky’s interest in natural populations from his interest in the broader social implications of evolution for humans and human societies. The two are interlinked in his thought. His critique of Muller is at the same time political and scientific.

Dobzhansky’s primary interest had always been the philosophical aspects of evolutionary theory and he was outspoken about this later in life. “Although a biologist may

504 Theodosius Dobzhansky to Ernst Mayr, 1974, Mather, Sierra, California; APS. It is also worth flagging that Muller was influenced by Loeb’s conception of biology in which the main aim was to control life processes and to create or engineer new biological possibilities (as we saw from the quote above). This aim was inconsistent with many evolutionary biologists who wanted to understand how nature worked undisturbed by human influence (see Pauly 1987, 177-83). 505 Consider synthesis architect G. Ledyard Stebbins’ recipe for making an evolutionary scientist: “Take a large bump of curiosity and expose it for several years to some spot like Seal Harbor, where from teeming tide pools through green woodlands, lush ponds, and orchid-filled mountain tops, nature presents her infinite variety of living things in tantalizing beauty. Add the knowledge, honesty, and scientific discipline provided by understanding parents, good schools, and the scientists of a great University. Mix carefully, and allow to rise in a atmosphere of tolerance, broad mindedness, and affection. The result is inevitable.” From an unpublished essay written in 1942, “The Objectives and Philosophy of a Modern Evolutionist”; APS. 506 It’s no wonder that Mayr referred to Muller as a typologist as well. See my discussion of Wallace and typology in the last chapter.

!206 do research on mice, Drosophila flies, plants, or bacteria,” he wrote, “the ultimate aim should be to contribute toward the understanding of man and his place in the universe.”507 While this interest can be traced back to his Russian roots, it was really in the late-1930s, and with the work on concealed genetic variability in natural populations, that Dobzhansky realized the broader implications of his own research. He recollected in 1962, ! It was consequently about 1937 or ’38—that was in my life as a researcher crucial—I became convinced that here is a chance of discovering something both of general biological interest, and something which would have bearing and relevance to man and to the human condition. We were playing with flies, but the existence of this concealed genetic variability, or of the genetic load, is a necessary consequence of the occurrence of mutation process, most mutations being harmful and natural selection not being absolutely effective. Natural selection cannot eliminate all these harmful mutations at once. They persist for a greater or lesser number of generations in the populations of flies and man.508 ! As Dobzhansky’s studies progressed, he became convinced of the ubiquity of genetic variation. But it was the evolutionary significance of this variation for the future of both fly and human populations that really interested him. In particular, he was worried about one

(mis)interpretation of this variation—one that we already encountered above—that treated variation as bad, something that a population would be better off without. He gave voice to

507 Dobzhansky 1973, ix, quoted in Paul 1987, 334. He made such remarks on many occasions, for example, “We study evolution of Drosophila because we hope thereby to elucidate the evolution of all life, and evolution of man in particular” (Dobzhansky 1960, 1). And, “The interest of this work, is due particularly to the universality of its application. As is sometimes said, man is not an overgrown Drosophila. Nevertheless there are many things about man and Drosophila which are alike in principle. The mechanisms of heredity are among them. As a matter of fact, the mechanisms of heredity are probably the most universal in biology. If you would study, for example, the physiology of the animal, you would find that the physiology of the fly and the physiology of man are very different. The mechanisms of heredity are surprisingly uniform” (Dobzhansky 1962a, 277). By 1955 Dobzhansky wrote that “the Drosophila work interests me less and less as such, and more and more insofar as it contributes to human problems” (Dobzhansky to L. C. Dunn, 14 Aug 1955, APS). 508 Dobzhansky 1962a, 413-4.

!207 this concern in the first edition of Genetics and the Origin of Species, connecting it directly to the future of human evolution. Dobzhansky writes, ! Judged superficially, a progressive saturation of the germ plasm of a species with mutant genes a majority of which are deleterious in their effects is a destructive process, a sort of deterioration of the genotype which threatens the very existence of the species and can finally lead only to its extinction. The eugenical Jeremiahs keep constantly before our eyes the nightmare of human populations accumulating recessive genes that produce pathological effects when homozygous. These prophets of doom seem to be unaware of the fact that wild species in the state of nature fare in this respect no better than man does with all the artificiality of his surroundings, and yet life has not come to an end on this planet. The eschatological cries proclaiming the failure of natural selection to operate in human populations have more to do with political beliefs than with scientific findings.509 ! Such views went directly against one of Dobzhansky’s most cherished beliefs: that variation was not only ubiquitous but beneficial and necessary, for both flies in the state of nature and humans. He did not deny that most mutations were deleterious to the organisms that expressed them. This, he bemoaned, was an “imperfection in nature.” But this imperfection was a necessary one, because without a large store of variation, as we saw above, a population could not cope with environmental change. “One of the causes of extinction is too narrow an adaptedness to a circumscribed biological opportunity which proves only temporary,” he argued.510

It was thus a mistake to think, as the eugenical Jeremiahs did, of usefulness and harmfulness as “intrinsic properties of a variant gene.” Genes “are useful, neutral, or harmful

509 Dobzhansky 1937, 126. 510 Dobzhansky and Allen 1956, 603.

!208 only in a certain environment,” he argued, “What is good in the Arctic is not necessarily good on the equator; what was good in man in the ice age is not necessarily good now; what is good in a democracy is not necessarily good under a dictatorship.”511 This, he said, might seem too elementary to require repetition “if it were not for the fact that it is so often forgotten in discussions of human evolution, even by some geneticists.”512

The “eschatological cries” were both unfounded and dangerous: if eugenical social policy was built on the erroneous assumption that variation was bad, it might be enacted to create a race of humans with as little variation as possible.513 Ideas “of the genetically new man very easily degenerate into something like the Platonic ideal Man with a capital M— something quite homozygous for the ideal genes and quite free from even a fraction of a lethal equivalent of the genetic load.” It is too easy “to let our imagination strive for something with a body as beautiful as a Greek god, healthy, and resistant to cold and to heat, to alcohol and to infections, with the brain of an Einstein and the ethical sensitivity of a

Schweitzer, the musical talents of an Oistrakh and the poetical talents of a Shakespeare.”514

This, according to Dobzhansky, was unrealistic and furthermore would only lead to a human population unable to cope with change. “Environmental instability,” he wrote a year later,

“presents challenges to the organism—both to an individual or to a population or a species.

To maintain itself in harmony with a changing environment, the organism must be not only adapted but adaptable [...] A species should not only possess genetic variety but also be able

511 “Environment” for Dobzhansky spans the physical, temporal, and social. 512 Dobzhansky 1961a, 461. My emphasis. 513 Dobzhansky was not anti-eugenics. See discussion below. He was against a form of eugenics that was based on what he took to be an erroneous view of the nature of variation and its consequences. 514 Dobzhansky 1961a, 472.

!209 to generate variety. It may then respond to changing environments by genetic changes.”515

Maintaining variation was evolutionarily crucial.

Although it is unclear exactly who (or what eugenics society) Dobzhansky had in mind in 1937, by 1938, as we saw above, he was connecting these mistaken claims with laboratory studies: “In the environment in which the flies have been kept in our experiments a large majority of the mutations are deleterious,” but there is “no basis for the assertion that these mutations will be deleterious in every possible environment.”516 This, unsurprisingly, coincided with the start of his “Genetics of Natural Populations” series (papers I and II were published in 1938). Only by studying natural populations could one see the poverty of this eugenical position—one needed to appreciate the importance of environmental change, but in the lab, this was exactly what was controlled for.

In the period after Genetics and the Origin of Species, Muller became, for

Dobzhansky and others, the embodiment of the position that variation was bad.517 Muller had been an earlier member of T. H. Morgan’s lab, prior to Dobzhansky’s arrival, and had won a

Nobel prize for his work on x-ray induced mutations in D. melanogaster. Muller’s position with regard to the nature and consequences of genetic variation was the exact opposite of

515 Dobzhansky 1962b, 289. As his career went on Dobzhansky began to see variation as not just good for the population but also good for each individual: largely heterozygous individuals were more plastic than largely homozygous individuals because they had two different alleles at each locus, rather than one, and thus had (genetically speaking) more options. In a paper published with the anthropologist M. F. Ashley-Montagu, they write, “Success of the individual in most human societies has depended and continues to depend upon his ability rapidly to evolve behavior patterns which fit him tot he kaleidoscope of the conditions he encounters. He is best off if he submits to some, compromises with some, rebels against other, and escapes from still other situations. Individuals who display a relatively greater fixity of response than their fellows suffer under most forms of human society and tend to fall by the way. Suppleness, plasticity, and, most important of all, ability to profit by experience and education are required” (Dobzhansky and Ashley-Montagu 1947, 588; quoted in Beatty 1994, 210). 516 Dobzhansky 1938, 449. My emphasis. 517 He may have had Muller 1929 paper “The Method of Evolution” in mind. Or his 1935 eugenical text, Out of the Night. John Beatty has called Muller’s position the “foil position” Dobzhansky reacted against.

!210 Dobzhansky’s.518 Muller argued that there was an optimal gene (or genes) for every character trait, nearly irrespective of environment, and that such optimal genes were for the most part already the norm. Mutational changes, at present, were always deleterious and natural selection was needed to weed them out. Dobzhansky summarized this position as follows, ! The classical hypothesis of population structure [Muller’s position] assumes that there exists for every biological species, or may appear to be produced in the future, one normal, or best, optimal genetic endowment. This genetic endowment would give the ideal, the normal, the archtypical man, or ideal Drosophila fly, or the ideal corn plant—the man, the Drosophila, the corn as they ought to be. The way to produce this normal man is to [...] remove, eliminate, purge from the population all the mutant genes. What then would remain would be the ideal constellation of genes.519 ! This view, Dobzhansky wrote, “has the advantages of simplicity, though not necessarily of accuracy.”520 It is a product of the “typological” thinking—a “typological fiction”—that promoted the Platonic ideal of Man.

If anyone was as interested as Dobzhansky in the repercussions of fly genetics for human evolution, it was Muller. A convinced eugenicist by nineteen, Muller promoted eugenical views founded on his research with Drosophila until his death in 1967.521 The problem with civilized human societies, for Muller, was that they had been largely insulated from natural selection, and thus had accumulated a large amount of concealed genetic

518 Dobzhansky strategically named Muller’s position the “classical” position, calling his own the “balance” position (see Beatty 1987b). 519 Dobzhansky 1961b, 290. 520 Ibid., 290. 521 See Paul 1987, Pauly 1987 177-83, Kevles 1985, 176-192, and Carlson 1981, Chapter 28.

!211 variation, what he called our “genetic load.”522 Consistent with his position that variation was harmful, Muller promoted eugenical approaches that would act to remove variation from human populations.523 Muller’s position was thus opposed to both Dobzhansky’s scientific understanding of the nature and consequences of genetic variation as well as his vision of the ideal human society.524 Muller envisioned a homogenous perfection, while Dobzhansky a world rife with variation—both in terms of genetics and forms of life.

Muller’s views, as I said, were based largely on his work with x-ray induced mutations in the laboratory, bolstered by a priori theoretical commitments—he drew the watch analogy, for example. Dobzhansky drew attention to this laboratory work to argue for the poverty of Muller’s position. In Mankind Evolving (1962), Dobzhansky prefaces Muller’s views with the following, ! In working with experimental animals and plants, biologists are often at pains to make the environment of their charges as uniform as possible. Devices are used to keep constant temperature and humidity, the animals are provided with uniform food and plants with uniform soil, etc. Let us suppose that the human species inhabits an environment at least as uniform as the ones used for laboratory animals. It is just conceivable that a genetic endowment might ultimately become selected which would be the best possible one for that particular environment. Any further genetic

522 See Muller 1950. This idea is old. In the last chapter we considered Wallace’s similar views on civilized society. See also Dobzhansky 1960. 523 On the classical view, “to put it in positive rather than negative terms, we might strive to obtain a mankind in which everybody would be a carrier of the normal genetic endowment” (Dobzhansky 1961b, 290). 524 This chapter is not about the “classical-balance” positions and so I will not treat them in anymore detail. See Beatty 1987a, 1987b, 1994. See above note on Dobzhansky’s paper with Ashley-Montagu. In Mankind Evolving, Dobzhansky writes, “Not only do people follow different ways of life, engage in different occupations, have different duties and interests, but human environments change rapidly and most rapidly of all in technologically advanced societies. It cannot be overstressed that the ideal of equality of opportunity, the evolutionary consequences of which were considered in Chapter 9, does not necessarily envisage environmental uniformity; equality of opportunity may also mean freedom to choose among existing environments or to create new ones” (1962, 289).

!212 change could then only be harmful. The reality is otherwise. Not only do people follow different ways of life, engage in different occupations, have different duties and interests, but human environments change rapidly and most rapidly of all in technologically advanced societies.525

In other words, the only place that Muller’s hypothesis holds true is in the very restricted and unrealistic environment of the laboratory. He continues, after summarizing Muller’s “fire- and-brimstone prophecy,” ! Drosophila flies are doing nicely in their natural habitats, despite the fact that they bear enormous genetic loads. The four lethal equivalents which constitute a person’s average load (see above) let most of us enjoy reasonably good health and well-being for something close to three score and ten years. This is so because most of the genetic load remains concealed and unexpressed: it is concealed owing to recessivity or incomplete dominance of the potentially destructive genetic variants. The adaptive norm (p. 126) of the human species consists of persons burdened with genetic loads. Nor is there anything new in this situation—all of human evolution occurred in populations that carried heavy genetic loads. A man or a fly free of genetic loads might perhaps be a superman or a superfly, but as far as anybody knows such a prodigy never walked the earth.526 !

525 Dobzhansky 1962b, 289. Emphasis mine. 526 Dobzhansky 1962b, 295-6. Emphasis mine.

!213 In other words, unless one was familiar with flies in their natural habitats, one could not appreciate that variation was both ubiquitous and beneficial in nature.527 Of course, there was no technical way for Dobzhansky to prove Muller wrong at this time; as John Beatty has shown, both of their positions were underdetermined by the empirical evidence available.528

One tactic Dobzhansky used to vitiate Muller’s position was to draw attention to the fact that it was based on laboratory results rather than a study of nature.529

For our purposes, what is most interesting is that “the natural” played a foundational role in Dobzhansky’s research: grand claims about the structure of human society could be made on the basis of fly research so long as they were backed by data that were valid in nature.530 We cannot thus separate Dobzhansky’s interest in “natural” populations from his

527 Dobzhansky’s student, Richard Lewontin, later recalled his position and its relation to Muller’s: “Dobzhansky certainly had a whole ideological view of the world, which was that human welfare depended on a diversity. Dobzhansky would have said, and I inherited that notion from him, that a population you made essentially homozygous by directional selection would lose the possibility of adaptation to changed circumstances. That’s the general theory of population genetics. Both Muller and Dobzhansky might have wanted to improve the human species, but whereas Dobzhansky would say, ‘maximum diversity is the best thing you could have,’ Muller would say, ‘no, we ought to get the best genotype, and we ought to select for it.’ Dobzhansky’s response, I think, would have been, and my response would be, if you really believe that genetic diversity (and I don’t believe that) is the basis for adaptability, then, although you might get a short-term advantage out of selecting for the best genotypes you can create, you’re really going to screw up the works again because Dobzhansky believe, remember, that homozygotes were narrow specialists. So you could select the homozygote that was better for the present situation, but it would have no ability to protect you against future demands and changes. Since the world is changing, you’ve got to have genetic diversity” (Lewontin 2001, 43). 528 See Lewontin 1981 and Beatty 1987b. 529 Although Dobzhansky was opposed to the eugenical views of both the “eugenical Jeremiahs” and Muller, it is important to note that he was not against eugenics tout court. He was opposed to eugenics that was based on what he saw as an erroneous view of the nature and evolutionary consequences of genetic variation. And importantly an erroneous view was one that did not adequately represent “natural” populations. In reminiscing about eugenics he wrote that “The standpoint which I for one find extremely difficult to understand is that any such thing is contrary to nature. What is contrary to nature and what isn’t, and who is there to tell?” And yet, somewhat paradoxically, he concluded this discussion, however “it is perfectly reasonable to ask, is Muller’s ‘Bravest New World’ project contrary to Nature?” (Dobzhansky 1962a, 449, APS). In other words, the difference between contrary to nature and scientifically unfounded was slight—especially when it worked in his favor. 530 Did he aggrandize “the natural” because it promoted this view of evolution (and eugenics), or did he promote this view of evolution (and eugenics) because it was what he found to exist in nature?

!214 interest in broader social implications. This may go back, as he later recalled, to his Russian roots and to the idea that, to be valid, a “paradigm” must hold in “pristine nature.”

Dobzhansky exploited this as a standard way to argue against Muller both in published text and in correspondence with allies like Mayr. It was effective because it took advantage of a common framework of understanding and communicating that was reinforced in the early-20th century in which “natural” and “artificial” carried descriptive and normative connotations. Laboratory research tended to promote views of evolution that were,

Dobzhansky found, both misrepresentative and undesirable. This, along with what we have seen above, only reinforced his own skepticism of studies that were too artificial, and furthermore caused the artificial-natural distinction to be one that he relied on frequently; he used the distinction to sort through concerns about which objects and methods were valid for biological study. !

CONCLUSION

In this chapter I have argued for two theses. The first was that assumptions about what was artificial and what natural played a role in structuring Dobzhansky’s research practice and his evaluations of others and their work. The second was that Dobzhansky tended to be skeptical of studies that were too artificial, and this was reinforced throughout his career, because he believed they promoted both misrepresentative and undesirable views of the nature and evolutionary consequences of genetic variation. I would argue also that, as the opening quotation by Mayr illustrated, Dobzhansky’s use of the artificial-natural

!215 dichotomy to classify aspects of biology was widespread amongst this generation of biologists.

In conclusion, I’d like to make four points that I think the above case corroborates and that speak to the central themes of this dissertation.

First, we might ask why the artificial-natural distinction is used so frequently during this time period, by Dobzhansky and others? One answer, brought out above, is that the distinction piggybacks on more fundamental and widespread concerns and anxieties about how human actions relate to nature’s normal development—how flies created with no eyes, for instance, relate to speciation and evolution. As I said in Chapter 3, even those cases in which the artificial-natural distinction is used rhetorically feed off the prevalence of these concerns.

Second, “nature” and “the natural,” for Dobzhansky, are ultimately not about place or physical location, as we might prima facie expect. Rather “Nature” is an ideal historical state.

“Natural” refers to the genetic history of the fruit flies, not simply whether they are free- living outside the laboratory. To Dobzhansky, even free-living D. melanogaster were not wild; their evolutionary history was stamped with human presence. Although Dobzhansky could point out the foolishness of the claims of his naturalist forebears—he scoffed at their rash assessment of mutations as “monstrosities”—he nonetheless still found fault with D. melanogaster as an object of study for its close relation to humans in its recent evolutionary past.

Third, as an ideal, “the natural” is endowed with normative content. It is more than just a description of properties. When something is classified as “natural,” it is deemed valid

!216 as an object of evolutionary study and endowed with a host of positive connotations that do work for Dobzhansky. It allows him, as we saw above, to justify his choice of experimental object, to evaluate the work of others, to speak authoritatively and convincingly about evolution, and to promote what he considered a desirable view of human evolution and social arrangement. When Dobzhansky referred to another biologist’s research as being based on merely “laboratory flies!,” for instance, he was both describing the fact that these flies were bred in captivity rather than taken from the field, and exploiting negative connotations that accompanied the term “laboratory,” a place of artifice and artificiality.531 So too when he referred to Muller’s views as “unsurprisingly naive” he suggests that those who have confined themselves to the lab have in some way failed as professional investigators of nature.

Lastly, this chapter has highlighted a number of ways in which humans have been considered disturbing conditions. In Chapter 3, I explained that “disturbing condition” is a phrase common in philosophy of science which, most broadly construed, is used to refer to those factors not included in a representation that, when present in the object represented, impede its accuracy or suitability. Throughout this chapter I’ve presented reasons why

Dobzhansky was skeptical of studies that were too human disturbed. These reasons were diverse and reflected personal, scientific, and political commitments. Dobzhansky worried that artificial studies did not accurately portray the true nature and consequences of genetic variation. Early laboratory studies had mistakingly suggested that mutations were unwanted monstrosities and that variation was ultimately harmful. Dubinin’s studies of D.

531 Dobzhansky to Ernst Mayr, March 29, 1975. Dobzhansky was wrong about the flies. Alfred Sturtevant, the biologist he is referring to, used flies Dobzhansky had sent him directly from the mountains east of Pasadena.

!217 melanogaster were unrepresentative of natural variation because the populations were frequently churned up by man. Only studies of “natural” populations, like those of D. pseudoobscura, undisturbed by human influence, provided a true or accurate picture of the nature and consequences of variation, and thus of evolution.

Humans were disturbing conditions for Dobzhansky partly because he had an ideal of

“pristine nature” in mind when testing hypotheses—nature was, following Dunn, the

“ultimate laboratory of biology.” This “ultimate laboratory” alone provided a true picture of variation and evolution. And following from this, the conclusions drawn from studies of

“natural” populations were not just ones that applied to flies, but to humankind as well.532 We should not forget that this “true” picture of variation and evolution also cohered well with what Dobzhansky wanted to be the case. This is not to say that Dobzhansky had no scientific reasons to count humans as disturbing conditions—laboratory studies suggested conclusions about variation that he, for scientific reasons, saw as false. After all, they did not agree with the data he obtained from natural populations. But it is that wider considerations about what was artificial and what natural, and what he thought this meant for the generality of the results, that played a significant role. For Dobzhansky, the “natural population” was the epistemic foundation for normative claims about science and politics. ! !

532 In 1960 Dobzhansky wrote that many people are skeptical that results drawn from experiments on flies might be legitimately “extrapolated” to humans. He argued that this was the wrong way of asking the question. We should not ask about “extrapolation.” “We do not extrapolate from Drosophila to man, or from man to Drosophila. What we are trying to do is to study the biological laws which are common to both. We study Drosophila rather than man because certain biological laws are easier to study in flies than man” (Dobzhansky 1960, 92). These laws are uncovered in the study of “natural” fly populations.

!218 ! ! CHAPTER 7 ! Communities, Natural Experiments, and “Soft” Science Jared Diamond’s Community Ecology ! ! ! ! ! ! ! ! it must be said that, in biology, the specificity of an object of observation or experiment limits unpredictably any logical generalization. Georges Canguilhem, 1951, Experimentation in Animal Biology !

In all its oxymoronic guise, the idea of the “natural” experiment has been a valuable one throughout the history of biology. It carries the authority of nature with ease, while still possessing the aura of rigor associated with the laboratory. But why exactly is it advantageous that an experiment be “performed” by nature rather than by human experimenters? And why has the artificial-natural distinction been a useful way for biologists to classify types of experimental work? Community ecologists confronted these questions in the 1980s during an internal dispute about what it meant to do “good” science. This chapter will address these questions by focusing on ecologist Jared Diamond’s use of natural experiments and his widely cited reflections on proper experimental methodology. I will argue that, contra any narratives about the collapse of the distinction due to the rise of experimentalism, concepts of the artificial and the natural have been central to experimentation in ecology.

!219 !

THE ARTIFICIAL AND THE NATURAL IN ECOLOGY

Natural experiments, as the preceding chapters attest, are a recurring theme throughout the history of biology. Georges Cuvier and Xavier Bichat classified them as part of a comparative method that overcame the disturbing influence of human manipulation on the “natural” object of study. Charles Darwin used a specific natural perturbation—flowers naturally lacking nectaries—to study fertilization. Early-20th century ecologists Frederic

Clements and Francis Sumner emphasized the experimental, as opposed to natural, side of natural experiments to counteract a widespread trend in favor of experimental laboratory science—a “peace offering to the experimenters,” a fellow field ecologist remarked.533

Despite the nominal uniformity, questions such as what is natural about natural experiments and why this makes them valuable have historically contingent answers.

Bichat’s worries about human manipulation, for instance, were motivated by his commitment to a complex vitalism, a motivation that would hardly apply to physiologists today. It is, however, in contrasts such as these that we may glimpse the importance of natural experiments. Since biologists contrast natural experiments with experiments involving a greater degree of human manipulation—experiments that are more artificial—these contrasts reveal the reasons why human disturbance is thought of as advantageous or disadvantageous.534 In this way, natural experiments can be used as indicators of human

533 See chapters 3 and 5. Darwin to Bentham June 17th [1861]; Albury 1977; Jacyna 1983; Outram 1986; Kohler 2002a; Kohler 2002b, 3. 534 If we were to put experiments on a scale, for instance, from the most natural to the most artificial, natural experiments would fall on the “natural” end; laboratory experiments would fall on the other end. This is not me introducing the artificial-natural terminology. This is how these experiments are often described. See, for instance, Shrader-Frechette and McCoy 1995.

!220 disturbance as some species are used as indicators of environmental disturbance. By looking at particular invocations of natural experiments we learn why less manipulation is often more. In this chapter I will examine a case of contrasting beliefs that arose in 1980s community ecology.

Experiments, it is worth pointing out, were not the only things ecologists discussed in natural-artificial terms; they were but one part of a larger trend. The philosopher Kristin

Shrader-Frechette and ecologist Earl McCoy have demonstrated the ubiquity and significance of this language throughout ecology and the many problems and puzzles that surround its employment.535 In one famous 1986 paper, to which we will return below, the author classifies the following according to this dichotomy: experiments, communities, situations, perturbations, variation, populations, colonizations, disturbances, and causes.536 Setting aside the problems and puzzles posed by such classifications, their recurrence alone is enough to indicate the importance of this dichotomy in ecological discourse. Here I will dive a little deeper and show that this language is fundamental to experimentation in ecology.

Debates in the 1980s over the value of natural experiments were caught up in two wider currents. The first came from outside the discipline. This time witnessed the rise of a pluralistic stance towards science: in biology, philosophy, and history, practitioners claimed that pluralism was a source of strength (or simply a reality), rather than an avoidable weakness. These arguments were put forward in opposition to the unificatory ideals advanced by early 20th century theorists—especially contra the stance that there was a scientific

535 Shrader-Frechette and McCoy 1995. 536 Diamond 1986a.

!221 method best instantiated in “harder” sciences.537 In those areas of biology “furthest” from physics—for example, ecology and evolutionary biology, as opposed to biochemistry—this trend manifested itself in terms of arguments for the autonomy of biology as a science.538 The employment of natural experiments was framed as one way in which biology was different.

The second current came from within ecology. Community ecology was a fairly young discipline at this time without a developed set of accepted epistemic standards, such as what counts as a legitimate explanation, inference or experiment.539 Whether natural experiments counted as “good” experiments was a bone of contention. These two currents will reveal themselves in the case study that follows. I will argue, contra any narratives about the collapse of the distinction due to the rise of experimentalism, that concepts of “the natural” and “the artificial” were central to experimentation in ecology, particularly as a way to clarify and maintain methodological distance between ecology and the so-called “harder” sciences. !

SOFT SCIENCES ARE OFTEN HARDER THAN HARD SCIENCES

In 1987 Jared Diamond, the polymath and at the time influential ecologist, published a popular piece titled, “Soft Sciences are Often Harder than Hard Sciences.” This article had a politics—he expressed strong dissent towards the National Academy of Sciences’ decision not to elect the political scientist Samuel Huntington a member—but his argument aimed to address a more general problem: “Do the so-called soft sciences, like political science and

537 See Platt 1964. 538 See Ernst Mayr 1982, 1985, 1997. 539 Although see Eliot 2011a for an earlier history of community ecology. When I say it is a “young discipline” in the 1980s, I am referring to a particular form of community ecology that was then becoming popular and grew out of the work of Robert MacArthur. The phrase “community ecology” goes back to the beginning of the 20th century.

!222 psychology, really constitute science at all, and do they deserve to stand beside ‘hard science,’ like chemistry and physics?”540 In the article he attacked the labels themselves, suggesting, somewhat facetiously, that soft and hard science be replaced by “hard (i.e., difficult) science and easy science” respectively. “Ecology and psychology and the social sciences are much more difficult and, to some of us, intellectually more challenging than mathematics and chemistry.”541 To Diamond, the rejection of a political scientist to the NAS betrayed deeper issues arising from common prejudices against certain sciences and ingrained ideas about the scientific method.

Diamond’s diatribe was not provoked by the 1987 NAS elections alone. Nor was it simply a result of general criticisms leveled at the “soft” sciences. The issues were closer-to- home than that. Since 1975 Diamond had been centrally involved in a set of controversies within ecology itself over its epistemic and methodological standards. Colloquially these are known as the “null model,” “competition,” or “SLOSS” controversies, and the philosophical issues involved have received considerable attention—particularly with regard to the statistical analysis of null models and hypotheses, and scientific model building and evaluation.542 My narrative will compliment these studies by offering an account of the methodological issues that arose from concerns about different experimental approaches.543

Let us rewind to community ecology, 1975.

540 Diamond 1987a, 34. 541 Ibid., 39. 542 See Cooper 1993, 2003 and Sloep 1993. See also David Quammen’s great discussion in The Song of the Dodo (1996). For recent, brief histories, see Dritschilo 2008, Kingsland 2002a, 2002b. For prehistory see Kingsland 1995. 543 I am working towards a full historical account of these controversies to explain the interrelations between the different facets involved. To save space I will here isolate the experimental issues.

!223 !

ALL THE WORLD’S A LAB: RESERVE DESIGN AND COMMUNITY STRUCTURE CA. 1975544

Collapsing the artificial-natural distinction has been, in one sense, central to the recent history of community ecology. And it was in this sense also central to Diamond’s ecological research. Ecologists have long been committed to what some call a “natural laboratory paradigm,” the idea that the natural world provides us with situations—“natural” experiments—that are analogous to laboratory experiments and can thus be analyzed analogously.545 Archipelagos are a favorite place of natural laboratory theorists—each island is treated as a test tube and by comparing test tubes researchers hope to determine the causal role of particular factors. Islands, as one proponent writes, “being discrete, internally quantifiable, numerous, and varied entities, provide us with a suite of natural laboratories, from which the discerning natural scientist can make a selection that simplifies the complexity of the natural world, enabling theories of general importance to be developed and tested.”546 The idea of nature as laboratory blurs the boundary between the artificial and the natural. We will return to this below.

In 1975 Diamond published two seminal papers, each committed to the natural laboratory paradigm. His laboratory: Papua New Guinea and the surrounding islands, known as the Bismarck archipelago (Figure 7.2). His conclusions in each paper were based on the results of natural experiments conducted for island and mainland bird populations. These

544 The title for this section is taken from the title of a recent co-authored paper by Diamond and shows his long- term commitment to the idea that the world is a laboratory which presents us with natural experiments. See Diamond and Robinson 2010b. 545 Whittaker 1998. 546 Whittaker 1998, 2004.

!224 natural experiments took two basic forms. One consisted of comparing different ecological communities—often on separate isolated islands—that were similar in many respects but that differed with respect to a factor whose influence he wished to understand (for example, the size of an island or the presence of a particular species). He called this form of natural experiment, natural “snap-shot” experiments. The other form was less static. Diamond would consider one ecological community before, during and after a natural disturbance. For instance, he would observe how resilient island communities were to invading species by watching what happened during an invasion. He called these natural “trajectory” experiments. These two forms of natural experiments helped him understand how different factors affected community structure—or in community ecological lingo, they helped him find patterns and determine the processes (competition, predation, etc.) responsible.

In the first paper, Diamond brought MacArthur and Wilson’s recent “theory of island biogeography” to bear on nature reserve construction. He argued that a set of bounded nature reserves surrounded by human-altered habitat resembles a set of islands.547 Supporting his conclusions with natural experiments, he argued for a number of geometric principles pertinent to reserve design (Figure 7.1).548 The most notorious principle he suggested, what became a metonym for the lot, stated that a single large reserve is superior to several small ones adding up to equal area. Large reserves, he argued, support greater biodiversity and lower species’ extinction rates. The controversy which resulted from his paper was aptly

547 See Diamond 1973 for his using natural experiments to directly test the predictions of the theory of island biogeography. 548 Since what is considered a sound conservation strategy—a “better” state of nature—changes over time, it is worth noting that Diamond was aiming to preserve species diversity (the number of species), with a particular emphasis on species that were rare and endemic to the region.

!225 titled SLOSS: single large or several small. Diamond’s paper received an unsurprisingly positive response from the conservation community since it provided scientific arguments they could use to convince government planners of the importance of large reserves.549 The

1980 World Conservation Strategy (a document jointly published by the United Nations

Environmental Programme, the International Union for the Conservation of Nature and

Natural Resources, and the World Wildlife Fund) reproduced Diamond’s diagram in full and endorsed Diamond’s recommendations.550

549 This was before the real beginning of conservation biology as a scientific field, which is typically seen to start with the work of Michael Soulé in the late 1970s. The First International Conference on Conservation Biology took place in 1978, for instance. See Quammen 1996, 529ff. The aims of conservation have changed significantly through time, but at this time biodiversity and species richness were at the top of the agenda and nature reserves were seen as the means. See an interesting recent piece in The Chronicle Review by Paul Voosen (2013). That conservationists had a hard time convincing government planners of the importance of reserves without “scientific” evidence on their side, see Diamond et al. 1976: “biologists have felt intuitively that most existing wildlife refuges are too small to avert extinctions of numerous species. However, because there has been no firm basis for even approximately predicting extinctions in refuges, biologists have had difficulty convincing government planners faced with conflicting land-use pressures of the need for large refuges. Recently several workers have recognized that a predictive understanding of extinction might be obtained from island biogeography” (1976, 1027). 550 IUCN 1980, ; For a discussion see also Quammen 1996, 480.

!226

Figure 7.1: Diamond’s Geometric Principles for Nature Reserve Design. (from Diamond 1975a, 143) (© Biological Conservation) ! In the second paper, Diamond offered an explanation of the origin of differences in ecological community structure.551 Biologists had long observed that communities

(populations of interdependent organisms of different species living together) in close proximity and that were only partially isolated (e.g., islands) tended to differ significantly in species composition. This was so even when those communities were formed of migrators from the same larger species pool (e.g., the mainland) and when migration was recurrent.

What factors, then, might explain the patchy species distribution observed on islands? While

551 Diamond 1975b.

!227 some had argued that community organization was random, or determined by abiotic factors,

Diamond argued that community organization was largely determined through population interactions, in particular, competition.

Once again drawing on the results of natural experiments, Diamond provided his answer in the form of seven community “assembly rules” that described, in very general terms, how species assemble into stable communities on islands. And once again he used the patterns he had found to shed light on the processes involved. As an illustration, one rule

Diamond proposed was, “Some pairs of species never coexist, either by themselves or as part of a larger combination.”552 For example, Diamond found that some species of fruit-eating pigeon never co-occurred: an island may contain one species or the other, but never both

(Figure 7.2). He attributed these differences to competition between species which fought over a similar niche. Together these assembly rules provided an explanation of the origin of differences in community structure. “The working hypothesis,” he summarized, is that

“through diffuse competition, the component species of a community are selected, and coadjusted in their niches and abundances, so as to fit with each other and to resist invaders.”553 This paper was as influential in the ecology world as his reserve design paper was in the conservation world. Not a bad year for Diamond. ! ! !

552 Ibid., 423. His debt to Robert MacArthur’s way of thinking from natural patterns to generalizations was obvious in this paper—MacArthur began his Geographical Ecology, for instance, with the following strong statement, “To do science is to search for repeated patterns, not simply to accumulate facts” (MacArthur 1972, 1). See text below. 553 Diamond 1975b, 343. Diffuse competition is the sum of competitive effects on a species in a community, as a result of resource use of all other species in that community.

!228 Figure 7.2, “Distributions of two Macropygia Cuckoo-Dove Species,” has been removed due to copyright restrictions. It was a map of New Guinea and the surrounding islands of the Bismarck Archipelago showing the “patchy” distributions of two Macropygia (Cuckoo- Dove) Species: M. mackinlayi and M. nigrirostris. This is a case in which Diamond found that some species of fruit-eating pigeon never co-occurred: an island may contain one species, the other, or neither, but never both. This gives rise to what he called a “checkerboard” pattern. Original Source: J. Diamond. 1975b. “Assembly of Species Communities,” in M. Cody and J. Diamond (eds.), Ecology and Evolution of Communities. Harvard University Press: Cambridge: 388. ! One reason why I have juxtaposed these two papers is that they demonstrate the close relationship between Diamond’s simultaneous political, policy-oriented aims and his theoretical ecological research. The two are inter-linked.

Another reason for emphasizing these papers is that they immediately created a controversy. Although lauded by many, Diamond’s papers struck a foul chord with ecologist

Daniel Simberloff and his colleagues at Florida State University, Tallahassee. The debate continues to this day but was especially severe from 1975-86.554 The so-called “Tallahassee mafia” had a number of complaints about Diamond’s work and each of these complaints provoked responses from Diamond and sympathizers, which provoked further rebuttals from

554 David Quammen called the ensuing battle “trench warfare” (1996, 446). In terms of the first paper, Simberloff originally seems to have come to very much the same conclusions as Diamond, but his assessment changed dramatically after 1974 (Simberloff 1974). See Simberloff and Abele 1976, 1982, Connor and Simberloff 1979, Strong et al. 1984. In 2009 Diamond published a paper which he argues finally ends the debate in his favor (Sanderson et al 2009). Unsurprisingly, this provoked a rebuttal from Simberloff et al and a further response by Diamond et al (Collins et al 2011; Sanderson et al. 2011). By 1986, Diamond no longer frequently responds to Simberloff and, furthermore, Simberloff tempers his criticism, see Connor and Simberloff 1986, Soulé and Simberloff 1986.

!229 Tallahassee, ad nauseam.555 The discussions get at times so blunt they are hard to read.556

Both sides pulled out all the stops: both wrote histories which appropriated Darwin; both argued that the other merely had “physics-envy.”557 In 1983 there was an article published in

Science aimed simply at presenting readers with a coherent picture of the many, perplexing stakes involved.558

The Tallahassee ecologists claimed that Diamond’s “natural” experiments were illegitimate means for testing hypotheses. They leveled this criticism at both of Diamond’s

1975 papers and offered what they considered to be more rigorous methodological alternatives. In the next two sections we will look at how experiments got mixed up in debates about the rigor and epistemic standards of community ecology. !

555 For the use of “Tallahassee mafia” see Lewin 1983, 636, and Dritschilo 2008. They claimed that (i) the empirical data that bear on refuge design did not support Diamond’s conclusions; (ii) Diamond’s recommendation was not a consequence of the theory of island biogeography, but was “based on limited and insufficiently validated theory and on field studies of taxa which may be idiosyncratic” (Simberloff and Abele 1976, 286); As Simberloff later put it, “In the absence of clear consensus among workers in the field, a recommendation supposedly dictated by ‘theory’ can be promoted as policy when ‘theory’ could well be written or construed to generate exactly the opposite recommendation” (Simberloff 1983, 630); (iii) the species patterns Diamond used to determine his assembly rules could have arisen simply by chance rather than interspecific competition; (iv) his reasoning was tautological since he used patterns on islands to explain patterns on islands; etc, etc. See, e.g., Diamond and Gilpin 1982; Gilpin and Diamond 1982; Roughgarden 1983; Quinn and Dunham 1983; and Lewin 1983 for an at the time current summary. And references therein for responses from the Diamond side. Also see references in papers in previous footnote. From the Simberloff side see, e.g., Simberloff 1980a, 1980b, 1981, 1983, 1987, Strong 1980, Strong et al. 1984. 556 In one response Diamond and his co-author write, “Diamond considered it unnecessary to belabor this point in every possible quote about assembly rules, because it was not appreciated that anyone would be so silly as to search for effects of competition in all pairwise species combinations of a fauna, until [Simberloff and co- author] did exactly that” (Gilpin and Diamond 1984, 334). 557 For the authority of history, see Diamond 1978, Simberloff 1980a. For each side pulling on physics-envy see, Diamond 1986a, plus sections below, and Simberloff 1980a, 1980c and Connor and Simberloff 1986. 558 Lewin 1983.

!230 HOW EXPERIMENTATION GOT MIXED UP, PT. 1: THE SCIENCE AND POLITICS OF ISLAND

BIOGEOGRAPHY

What Simberloff and colleagues found unsettling was Diamond’s use of community patterns—determined by natural experiments—to infer underlying processes, and following from this, the broad generalizations he drew about design policy and community assembly.

Diamond’s reasoning had been in the form of inferences to the best explanation: the best explanation, he thought, for the patterns he observed was competition between populations of species. Supposedly, at this time Simberloff adorned the door of his office with newspaper clippings of a common theme: one read, “Image On Tortilla Draws Crowds to Hidalgo

Home.”559 The image was of Jesus Christ, and people had flocked from around Mexico to see the pattern on the tortilla. Simberloff compared the patterns Diamond analyzed to the face of

Jesus on the tortilla: he doubted that they reflected anything biologically meaningful. In his less agonistic moods, he seemed simply to claim only that nothing in Diamond’s data entailed that competition must have been the cause, which was true.560

The seeds of hostility that grew around Diamond’s 1975 papers, and that split the ecological community into “competitionists” and “anti-competitionists,” were planted a few years earlier and were always connected to ecology’s epistemic standards. In the 1960s a younger generation of ecologists, led by the prolific and imaginative Robert MacArthur, reacted against what they saw as a largely descriptive discipline. They emphasized mathematics and theory, drew comparisons to physics, and most importantly, and this was as

559 For this story see Quammen 1996, 478-9. 560 Connor and Simberloff 1986, 161.

!231 much an aesthetic choice as any other, they focused on repeated patterns in nature and on generalizations based on these patterns.561 In MacArthur’s own words, ! To do science is to search for repeated patterns, not simply to accumulate facts, and to do the science of geographical ecology is to search for patterns of plant and animal life that can be put on a map. The person best equipped to do this is the naturalist who loves to note changes in bird life up a mountainside, or changes in plant life from mainland to island, or changes in butterflies from temperate to tropics. But not all naturalists want to do science; many take refuge in nature’s complexity as a justification to oppose any search for patterns. This book is addressed to those who do wish to do science.562 ! The concept of pattern or regularity is central to science. Pattern implies some sort of repetition, and in nature it is usually an imperfect repetition. The existence of the repetition means some prediction is possible.563 ! The work for which MacArthur is most famed is his co-authored book, with E. O.

Wilson, The Theory of Island Biogeography (1967). In this book they consciously aimed to move ecology beyond its descriptive phase and offered theories to explain, for example, why fewer species were found on islands than on the mainland. These theories were, at the time, backed by little direct evidence and were largely inferred from natural patterns and theoretical-mathematical assumptions; they made largely qualitative predictions that intentionally ignored quantitative variation.564

561 See Kingsland 1995, Chapter 8. See page 190 for these scientific stances as aesthetic positions. 562 MacArthur 1972, 1. 563 Ibid., 77. 564 MacArthur had been inspired by Richard Levins’ (1966) thoughts on model-building. Levins argued that models may hope to fulfill three desiderata: precision, generality, and realism. In practice these desiderata traded off against one another and gave rise to three strategies. MacArthur’s favorite strategy was to sacrifice precision to gain realism and generality. Such models would only make qualitative predictions. See Kingsland 1995, 188-9, on MacArthur and Levins’ friendship.

!232 Simberloff and Diamond both emerged from the MacArthur-Wilson school of ecology. Simberloff had been a graduate student of Wilson’s at Harvard and for his dissertation had worked with Wilson to test the theory of island biogeography’s predictions on a number of mangrove islands in the Florida Keys. His work showed that the predictions of the theory were more-or-less accurate. After finishing a PhD at Cambridge in biophysics

Diamond took up residence at Harvard as a Junior Fellow, at which time he developed his growing interest in ecology and ornithology. When MacArthur died in 1972, at the young age of 42, Diamond was effectively designated as his successor.565

Diamond’s 1975 papers fit the MacArthur mold. They were mathematical and theoretical. They offered generalizations that were largely qualitative in nature. They suggested that competition was a driving force of community assembly. They used models developed by MacArthur. And most importantly, Diamond used natural experiments and studied large-scale biogeographic patterns. In his Geographical Ecology, a book written shortly before his death, MacArthur drew upon Diamond’s research and defended the rigor of natural experiments: ! since they [islands] have fewer species than the same habitats have on the mainland, they can also be viewed as natural experiments. If we wish to know how the abundance of a species would change if we removed that species’ competitors, we can look for an island which has the species present without its competitors. Purists often claim that these are rather poor experiments. They note that other conditions may also be different on the islands, and that these differences may be responsible for the contrasts in abundance. For instance, the islands may also be missing predators and our species may become commoner more by virtue of release from predation

565 Lewin 1983, 636.

!233 than from competition. So far, so good; we have an explicit and plausible alternative, and we must make the observations needed to discriminate between predator release and competitor release. But some critics go further and say that unspecified unexpected things may be different on the islands, and therefore that we should not be impressed by island-mainland comparisons. One could as validly say that unexpected things may be different between any experiment and its control and that experiments are not to be trusted. But island-mainland comparisons are really dramatic and call for some explanation.566 ! The explanations that MacArthur offered to explain island communities in 1972 were substantially improved upon by Diamond in 1975, but the MacArthur method was still the foundation.

Sometime between 1974 and 1976 Simberloff, on the other hand, had a change of heart.567 In 1976 he wrote a lengthy critical review of his earlier work with Wilson that raised a host of questions about the usefulness and accuracy of the theory of island biogeography— the same theory his earlier research supported.568 Over the next few years a very different understanding of ecology started to emerge from Simberloff’s writing: one that was autecological (close study of individual organisms) rather than synecological (studies of communities), that focused on manipulative field experiments rather than natural experiments, that emphasized hypothesis testing rather than inference to the best explanation, and that was adamantly opposed to generalization. The theory of island biogeography,

566 MacArthur 1972, 111. 567 In 1974 Simberloff wrote a review that demonstrated the positive impact of the theory. He must have changed his opinion soon after. The 1974 paper also included a positive assessment of Diamond’s work, see 1974, 166. 568 Simberloff 1976.

!234 Simberloff said, “has caused a generation of ecologists to waste a monumental amount of time.”569 Ecology of the MacArthur and Diamond type was a “sick science.”570

As mentioned above, there was at this time considerable discussion of the supposed fact that some disciplines—with an emphasis on molecular biology and physics—make rapid progress, while others remain relatively “stagnant.”571 The progressive disciplines were thought to share a particularly effective scientific method: the formation of alternative hypotheses and the attempted falsification of one of these hypotheses through experimentation.572 One proponent of this method recommended that scientists approach

569 Lewin 1983, 636. 570 Simberloff 1980c. 571 Ibid., 50. 572 Simberloff writes, “At least where experiment is possible, it seems to me that application of Popper’s procedure (1963, 1972), clearly state hypotheses and rigorous attempted falsification, is more likely to get closer to an accurate account of nature, and do so efficiently. Platt (1964) provides a nontechnical description of how this procedure, which he terms ‘strong inference,’ has spurred a remarkable series of successes in areas of physics and molecular biology and argues that it is strict formalization that is responsible” (Simberloff 1983, 627). As the quotation indicates, this perspective was influenced by the philosophies of Karl Popper and the physicist John Platt. Popper’s method is familiar to philosophers of science. To test a theory we deduce an observational prediction from it. We set up an experiment to test this prediction. If the prediction is incorrect, we have falsified the hypothesis; if the prediction is correct, then we have not yet falsified the theory and so we continue testing. Accordingly, theories are not confirmed by evidence; they can only be falsified by contrary evidence. Theories that last the test of time are the ones we should, at least for the time being, accept. It is worth noting that this view of scientific testing fits particularly comfortably with a classic view of science as striving for laws of nature: exceptionless generalizations of the form “All A’s are B.” If we test and find an A that is not a B, then we have falsified our hypothesis. John Platt’s 1964 paper “Strong Inference”—which I drew from above—combined a simplified version of Popper’s philosophy with a reverence for the methods pursued by molecular biology and physics. He argued that the rapid progress of the latter scientific fields was due to their underlying methodology: “strong inference.” If soft sciences like ecology were to advance in the same manner as hard sciences, he strongly recommended they systematically adopt his four step method (which reads very much like Popper): (1) Devise alternative hypotheses; (2) Devise a crucial experiment, with alternative possible outcomes, each of which will exclude one or more of the hypotheses; (3) Carry out the experiment so as to get a clean result; (1’) Recycle the procedure. Platt saw his theory of strong inference as a “yardstick of effectiveness.” “Surveys, taxonomy, design of equipment, systematic measurements and tables, theoretical computations—all have their proper and honored place,” he wrote; unfortunately, “all too often they become ends in themselves, mere time-serving from the point of view of real scientific advance, a hypertrophied methodology that justifies itself as a lore of respectability” (Platt 1964, 351). He believed the triumphs of molecular biology using cleverly designed simple model systems have “not fallen to the kind of men who justify themselves by saying, ‘No two cells are alike’” (Ibid., 349). In other words, the complexity of a science is irrelevant to what methods should be used; what are needed are clever people to devise simple tests.

!235 each other’s research by asking, “But sir, [sic] what hypothesis does your experiment disprove?”573 No answer was the mark of a “soft” science.

Simberloff and the Tallahassee ecologists took these discussions to heart and advocated this account of scientific method throughout the 1980s. “Sciences that have progressed rapidly (physics, chemistry, molecular biology),” they wrote in a coedited volume, “have made great use of sorts of evidence that ecology has not, of vigorous hypothesis testing and experimentation.”574 Simberloff wrote that “the crowning achievements of molecular biology were achieved in spite of complicated inputs, by clever experimental procedures that could unambiguously test well-chosen hypotheses”575

Analogous procedures would provide ecologists with the same payoff.

This stance on proper scientific methodology caused the Tallahassee ecologists to be skeptical of Diamond’s natural experiments. What Diamond referred to as “natural experiments,” Simberloff and colleagues called “nonexperimental observations.”576 While the Tallahassee ecologists understood that experiments in ecology couldn’t be as simple as those in molecular biology—given the differences between their objects of study—they did often recommend the use of experiments that were as simple as possible, involved greater experimental manipulation, and that aimed exclusively at falsification. In 1980, Simberloff wrote,

573 Platt 1964, 352. 574 Strong et al. 1984, viii 575 Simberloff 1983, 631. 576 Following the lead of molecular biology meant that the best way to test a proposed hypothesis was through controlled and replicated experiments using simplified experimental systems. Given the nature of the objects of study in molecular biology, this method was powerful and could provide in many cases an unequivocal result, falsifying a proposed hypothesis. Molecular biologists, as Diamond conceded, could unravel a nearly universal genetic code through short-term laboratory experiments by employing this method. Diamond and Case 1986, x.

!236 ! Whatever the reason, the predominant absence of critical manipulative work makes it difficult or impossible for ecology to progress through the method of “strong inference” proposed as a model by Platt (1964), that has worked so well in molecular biology: the posing at every step of a series of alternative hypotheses whose relative merits are swiftly and cleanly elucidated by a concise, economical experiment. The dearth of experiment may be because ecological experiment is more difficult than that in molecular biology and the physical sciences. But if that is the excuse, ecologists must retract inflated claims and admit their inability to form universal laws or theorems in the tradition of other sciences, and aim instead at idiosyncratic description and mechanistic explanation of specific systems.577 ! Ecologists should “insist that mathematical or verbal theory without direct, rigorous field testing no longer be recognized as a significant contribution.”578 So much the worse for

MacArthur. The “key to a clear result,” Simberloff wrote, was “a system sufficiently simplified, by whatever means, to allow an unambiguous test of the hypothesis.”579

If the goal was to unequivocally falsify hypotheses about ecological reserve design or community structure, there were much simpler alternatives to Diamond’s natural experiments

—which involved a decade of research—and ones which better fit the task of falsification.

For example, Simberloff and Lawrence Abele contended that in his 1975 paper on reserve design, Diamond didn’t aim to disprove any existing and clearly articulated hypothesis at all, and instead drew very general conclusions from patterns found within a large collection of observational data.580 They furnished, as well, a simple field experiment which disproved

Diamond’s primary reserve design principle: that a single large reserve can support more

577 Simberloff 1980c, 51. My emphasis. 578 Ibid., 52. 579 Ibid., 631-2; my emphasis. 580 Simberloff and Abele 1976.

!237 species than several small. The field experiment was done on arboreal arthropod species living on small mangrove islands. After taking a species census of a number of islands, they experimentally subdivided these islands into smaller ones. Following a recolonization period of three years, they found that the number of species on the new subdivided islands was greater than on the initial larger islands, thus providing evidence against, even purportedly falsifying, Diamond’s principle.

Diamond and his colleagues retorted that Simberloff and Abele’s simple arthropod experiment was biologically unrealistic: the species studied had little in common with those we design reserves to protect and the protection of particularly sensitive species was the goal, not simply overall species quantity.581 Even worse, they argued, Simberloff and Abele failed to appreciate that this research was politically non-innocent: developers might exploit their research to ground arguments against the need for nature reserves.582 Critical reviews of

Diamond’s work, in other words, could be politically dangerous.

With regard to Diamond’s second paper on assembly rules, Simberloff and Edward

Connor offered an ultimatum. Either (i) Diamond had to prove that competition was the primary determinant of community structure through “observed active replacement of one species by another [...], [field] experiment, or very detailed autecological study,” or (ii)

Diamond had to test his patterns against a “null model” to show that they differed significantly from what would be expected if the patterns were created by chance alone.583

581 Diamond et al. 1976. 582 See Kingsland 2002a. See also Diamond et al. 1976. 583 Connor and Simberloff 1979, 1986. For the quote see 1979, 1138.

!238 Again, Diamond and colleagues questioned both horns of the dilemma. With regard to the first horn, Simberloff and Connor, they rebutted, were claiming that evidence from natural experiments—in this context, observing the result of naturally occurring species invasions—was “intrinsically inferior to evidence from” field experiments in which the

“invasions” were human-mediated.584 They agreed with Simberloff “that the failure of several experimental [i.e., human] introductions of each species [of lizard] onto islands occupied by the other is supportive evidence for a role of competition.” But “Why is it less strong evidence,” they asked, “when the successful self-introduction [i.e., not by humans] of the pigeons Ptilinopus superbus and P. insolitus to Umboi between 1913 and 1933 was followed by the near-extinction of the formerly abundant P. solomonensis?”585 The only difference between a natural experiment and a field experiment in this case appears to be the cause of the experimental perturbation, in one case humans, in the other, naturally occurring.

Why they asked should human manipulation be, in this case, considered evidentially relevant?

A full explanation of the second horn would take us too far a field. In summary,

Simberloff and Connor argued that Diamond’s patterns did not significantly differ from what could be expected by chance, and Diamond and Gilpin (among others) continued to argue that the “null models” offered by Simberloff and Connor were problematic, particularly in that they tended to “swamp” or significantly underestimate the effects of competition, even

584 Gilpin and Diamond 1982, 83. 585 Ibid.

!239 when those effects were known in advance.586 If nothing else, the null model controversy created a climate of suspicion around the the scientific credibility of natural experiments and the usefulness of natural patterns.

Paradoxically, both sides of this debate employed “physics-envy” as a rhetorical strategy.587 The paradox is only illusory however since each side drew on a different notion of physics. Simberloff chided Diamond for his use of theorizing; lofty theorizing, he thought, was an indication of physics-envy.588 Diamond, on the other hand, as I pointed out above, often drew attention to Simberloff’s reverence for the inferential, evidential, and especially, experimental practices used in molecular biology and physics. Ecologist Robert May drew attention to the irony in Simberloff’s position when he wrote, “it is paradoxical that some of those who are most sensitively aware of the need to keep sight of alternative explanations for observed patterns in community structure seem, at the same time, occasionally to accept that there is only one True Way to do science.”589

A useful way to characterize the methodological claims made by Simberloff and his

Tallahassee colleagues is to say that they advocated a form, albeit weak, of what David

Rudge has called “methodological reductionism”: the claim that methodologies are legitimate in biology only in so far as they are based upon “proven” methodologies in the physical sciences.590 They often compared the inferential, evidential, experimental, and explanatory

586 Two ecologists at the University of Berkeley, for example, showed that “when a model archipelago whose species distribution is known to be affected by competition is established by computer simulation, the Connor- Simberloff null hypothesis fails to detect those effects.” Quote from Lewin 1983. 587 See Milam 2010a, for an interestingly similar fundamental rejection of methods from the physical sciences by evolutionary biologists in the 1960s. 588 And Simberloff rightly pointed out that MacArthur often contrasted ecology and physics to emphasize the impoverished theorizing of the former. See Simberloff 1980a, 23. See also Kingsland 1995, 198. 589 Quoted in Lewin 1983, 636. 590 Rudge 1996, 250.

!240 practices used in ecology to those used in laboratory physics and molecular biology and used these comparisons to support normative prescriptions about what methods ecology should and should not use. The methods Diamond used, they claimed, offered “but ‘soft corroboration,’” and provided “flimsy” evidence that had “little potential to falsify theory.”591 !

HOW EXPERIMENTATION GOT MIXED UP, PT. 2: MAKER’S KNOWLEDGE AND THE ACID

TEST OF UNDERSTANDING

While the Tallahassee criticisms were perhaps foremost on Diamond’s mind as they challenged his work directly, there were other expressions of methodological reductionism in the air—even among those who sympathized with Diamond, for instance his collaborator,

Michael Gilpin.592 Michael Gilpin and his colleagues described their approach to community ecology as “community reconstitution”: the study of synthetic Drosophila communities in the laboratory. Our approach, they write, is “the construction of communities from individual parts. We believe that the acid test of one’s understanding of any complex system—whether it be a clock, a virus, or an ecological community—comes in trying to reassemble the system

591 Strong et al. 1984, ix. Diamond’s methods were unconventional—which I should emphasize doesn’t mean unscientific—in others ways which could not have helped his case. For instance, in order to determine accurately the bird distributions on New Guinea, Diamond utilized the indigenous peoples. Learning to speak their native languages, he could treat them as “walking encyclopedias of bird lore.” He wrote, “they could distinguish at a distance obscure sibling species in such taxonomically difficult genera as Sericornis; and could accurately describe other species known to them only from single individuals observed up to 10 years previously. When such ‘walking encyclopedias’ of bird lore confirm the permanent local absence of a species that is regularly encountered in other areas, one can have confidence that the species is actually absent and not merely overlooked” (Diamond 1973, 762). 592 See Mertz and McCauley 1980 about the relationship between laboratory ecology and field ecology at this time. As they argue, “As a result of their different domains [...] field and laboratory ecology have begun to develop different vocabularies, differing ways of looking at nature, different intuitions about natural processes, and, consequently, a partial communications breakdown. There is sometimes a mistrust by field workers for laboratory results, and there is a growing possibility that this mistrust may become reciprocal” (1980, 95).

!241 out of pieces. Reconstruction is the procedure that accounts for many of molecular biology’s successes.”593 Success that community ecology might enjoy as well, it is implied.

Gilpin also further explored the “synthetic approach” to ecology by treating restoration projects as experiments in community assembly. He traced this approach back to the 17th century Italian philosopher Giovanni Battista Vico who wrote, “the condition of being able to know something truly, to understand it as opposed to merely perceiving it, is that the knower himself should have made it.” Molecular biologists, Gilpin and colleagues argued, “demonstrate their understanding of biological subsystems such as membranes and cellular organelles, and also processes such as photosynthesis [...] by taking them [...] out of cells and then putting them back together and making them run in vitro.” We “understand thoroughly only as far as we are able to reassemble, adjust and control.”594

The question is, how well does this method work in community ecology? Through the volume they attempt to use restoration projects as synthetic experiments to take advantage of a method that has been very productive in other sciences. It might be somewhat telling that

Michael Gilpin began his career as a physicist—although, then again, Diamond began as a biophysicist.595

593 Gilpin et al., 1986, 23; emphasis in original. That this is the acid test of one’s knowledge is stated by others as well. See Bradshaw 1987; add refs from experimental ecology volume from 1998. They write, “This approach tests whether we really have identified the significant components of a higher system, just as does the approach of a biochemist attempting to reconstitute the mitochondrial electron-transfer system from its components” (Ibid., 40). 594 Jordan et al., 1987, 11. 595 From Diamond’s chapter in this volume, we can see that he agrees with their use of restoration as an “acid test” of our understanding. So the point is not that Diamond was against synthetic type projects. The point is that because these types of projects are—restoration notwithstanding—not the types of projects ecologists can usually pursue—as Diamond says, they offer a “unique insight”—the rest of ecological research should not be viewed as less significant. Again, such a point of view comes from thinking about science in terms of a scientific method, that pursued in molecular ecology.

!242 There were thus a host of threads running through community ecology at this time which shared in common a reverence for the methods used by the “harder” sciences.

Laboratory physics and molecular biology, however, had little use for the types of experiments Diamond had utilized to gain an understanding of ecological communities. On what grounds could the employment of his methods be justified, if their utility had not been formerly proven by the “harder” sciences? !

DIAMOND ON EXPERIMENTAL PLURALISM, TRADEOFFS, AND POPPERPHILIA

Amidst the quarreling about assembly rules and principles for reserve design,

Diamond published two papers on proper experimental design in ecology.596 These have become widely cited classics in the literature. The relevance of these papers to the controversies of which Diamond was involved should not be overlooked.

Diamond began by pointing out that ecologists employ three types of experiments which grade into one another, but that have coexisted uneasily: laboratory, field and natural experiments. These experiments fall, following philosopher Robert Brandon, along a continuum of manipulation: “If our response variable—that which we measure and record in an experiment—is thought of as being a function of some number of independent variables, then changing and/or controlling the values of some or all of the independent variables is what constitutes a manipulation.”597 Laboratory experiments involve the most extensive manipulation, field experiments are intermediate, and natural experiments the least (since the independent variables are not intentionally and directly controlled by the experimenter). In

596 Diamond 1983, 1986. 597 Brandon 1994, 61-2.

!243 field and natural experiments, the experimental controls, and sometimes even the particular experimental perturbation (e.g., a fire), are provided by the way in which natural sites are arranged, rather than by human experimenters (see Appendix 2).

These experiments, Diamond argued, involve tradeoffs. Diamond explained that tradeoffs occur along eight axes: regulation of independent variables, site matching, ability to follow response trajectories resulting from perturbations, temporal scale, spatial scale, scope

(range of species and manipulations that can be studied), realism (whether results apply to natural communities), and generality (number of natural communities to which results apply)

(see Appendix 3).598 These axes constitute a multidimensional space of experimental types.

Each axis of this space represents a different feature of an experiment, for example, the first axis is the degree of experimental control over independent variables. Each point in this space represents a combination of such features. The argument that there are tradeoffs between experimental types is equivalent to saying that not all points in this space can be realized.

Laboratory experiments, for example, have the virtue of allowing one to control all independent variables (1st axis), and are thus the method of choice for valid causal designation and for unequivocal hypothesis testing. They, however, suffer from being

“uselessly unrealistic” (7th axis): laboratory experiments involving competition between species of paramecium “told nothing about whether competition is important in natural populations,” Diamond writes.599 Laboratory experiments focused on species that could be

598 Diamond 1986a, 4-5. 599 Diamond 1983, 586. Diamond is here referring to the experiments of ecologist G. F. Gause from the 1930s. See MachArthur 1972, for an overview that Diamond would likely have agreed with.

!244 used in a laboratory setting, such as cultures of paramecium, yeast or beetles; often the species studied in artificial laboratory communities never even came into contact outside of those circumstances. Natural experiments, on the other hand, sacrifice all human experimental control, but are realistic and permit one to examine conditions that would be impossible (too large), impractical (too much money involved) or unethical (violate basic animal rights) to study otherwise. Field experiments are intermediate along all axes and are advantageous for this reason.

Each of Diamond’s axes represents a particular experimental virtue or advantage. We should understand these experimental virtues as ways of evaluating the adequacy of experimental methods.600 Tradeoffs among virtues is a common theme in the philosophy of science. We often speak of theoretical virtues, such as simplicity or accuracy, as trading off against one another. One community of scientists may, for instance, accept a simpler theory even when it is less accurate, because they value simplicity over accuracy.601 Similar remarks have recently been made about scientific models.602 Diamond’s experimental virtues, however, apply to neither theories nor models; they are experimental. They serve to evaluate experimental types based on a widely acknowledged set of criteria.

Diamond felt that the Tallahassee ecologists failed to appreciate the non-overlapping nature of these virtues. Simplicity of the experimental system or degree of experimental manipulation are certainly virtues, but experiments with these virtues may be lacking in other important respects. Proper experimental methodology meant understanding and respecting

600 See Stegenga 2013. 601 See Douglas 2009. 602 Weisberg 2006. Scientific modelers are often forced to heed tradeoffs between different modeling virtues, such as precision and generality.

!245 these experimental tradeoffs. As I said above, in reference to Simberloff and Abele’s simple arthropod experiment, Diamond and colleagues retorted that the experiment was unrealistic.603

Diamond went one step further by arguing that these tradeoffs give rise to experimental pluralism. He begins by recognizing that each type of experiment has inherent advantages and weaknesses. Tradeoffs, say between realism and the regulation of independent variables, make it such that in community ecology, one has to use each of the different types of experiments—laboratory, field and natural—to realize different aims.

Tradeoffs make pluralism unavoidable and necessary. “[E]ach methodology,” he writes,

“yields some information inaccessible to the others.”604 Proper experimental design involves respecting the tradeoffs between experimental types and utilizing that method most appropriate to the question asked and species studied. Obviously this argument depends on our scientific goals: we are forced to accept pluralism because we want to maximize all of these virtues but find that they trade off against one another in practice. Furthermore, because the major tradeoff is between regulation of independent variables and realism, we are forced to accept a particular type of pluralism, that between laboratory, field, and natural experiments.

Diamond’s argument has a clear objective, to justify the use of a particular set of experimental approaches in ecology, creating space for natural experiments. His argument is aimed at justifying experimental pluralism in contrast to the monistic ideals advocated by the

Tallahassee ecologists.

603 Diamond et al. 1976. 604 Diamond 1986a, 22.

!246 Diamond is not so naive as to think that tradeoffs between experimental virtues hold equally across different sciences. From a philosophical point of view, this is perhaps his most interesting point. He explains that in comparison to community ecology, laboratory experiments “are often much more realistic in physiology” or in molecular biology or physics.605 In other words, the regulation of independent variables (1st axis) and realism (7th axis) do not trade off against one another to the same degree in these latter sciences. Such sciences need not heed the experimental tradeoffs that community ecology must manage.

All of this Diamond brought to bear on his quarrels with the Tallahassee ecologists.

“[T]here has been concern that the choice of experimental method might be critical to the conclusions reached,” he writes, “especially, that conclusions about the role of competition

[for example] might be an artefact of natural experiments and might not be sustained by field experiments.”606 He furnished independent and corroborating field experiments against this, but the issue was not simply whether Diamond’s experiments would be confirmed by independent methods, but whether natural experiments were a legitimate method at all.607

Two related arguments, he said, were used to buttress claims for the the superiority of one method over all others. These arguments were unmistakably those promoted by the

Tallahassee ecologists. They were “Popperphilia,” an unhealthy obsession with philosopher

Karl Popper’s falsificationism, and the belief that “ecology would progress faster by imitating the rigorous approaches of hard sciences like physics and molecular biology.”608

605 Diamond 1986a, 5. 606 Diamond 1983, 587. 607 He continued, “some proponents of each method have claimed that their method is inherently superior and is the method of choice” (Ibid.). 608 Diamond 1986a, 20-1.

!247 These arguments shared a methodological reductionism and promoted methodological unity among the sciences. In doing so they cut counter to Diamond’s third and forth points: different sciences face tradeoffs in experimental virtues to different degrees.

The more general problem, however, was that “Even scientists who work in pluralistic fields tend to view how science ‘should’ be pursued in ways that are mismatched to their field’s special needs.”609 “Many ecologists,” falling victim to this tendency, “feel defensive about whether ecology is a rigorous science and are determined to avoid anything smacking of nonrigor.”610 For Diamond, proper experimental practice meant respecting the differences between the sciences. Instead of holding up the so-called hard sciences as an ideal, ecology must cope with “its own distinctive problems.”611 To model ecology off other sciences, especially those which rely predominantly on short-term laboratory experiments, is wrongheaded and, given the applied nature of ecological knowledge, potentially ethically and politically questionable.

In summary, what prompted Diamond to reflect on proper experimental practice were a set of methodological prescriptions, justified by their use in the “harder” sciences but which piggybacked on widespread anxieties about ecology’s rigor. Natural experiments, and other less manipulative methods, fit awkwardly with these prescriptions which often equated rigor with simplicity, ability to falsify hypotheses, and experimental manipulation. Diamond worried that rather than see these prescriptions as mismatched, ecologists would see such methods as inherently bad. Central to his arguments against this assessment were his

609 Diamond and Case 1986, x. 610 Diamond 1986a, 12. 611 Ibid.

!248 experimental tradeoffs and the particular methodological pluralism they warranted. Caught in the fray of wider intellectual currents favoring pluralism over monism, Diamond’s reflections on experimental practice, and especially his tradeoffs, have been frequently repeated.612 !

WHEN LESS MANIPULATION IS MORE

We can interpret Diamond’s methodological papers from this period as a defense of natural experiments. Diamond’s argument (reconstructed by me) is that, given the nature of the phenomena studied in ecology, generality can be purchased only by sacrificing experimental control. The benefits outweigh the costs, however, when the goal is to apply ecological knowledge to problems of conservation. Such considerations, as we will see, force upon ecologists a keen awareness of the artificial-natural distinction.

According to Diamond’s schema, a tradeoff exists between the “regulation of independent variables” (axis 1) and “realism and generality” (axis 7 and 8) (Table 7.1). The former is a property of an experimental system; the latter are properties of the relationship between the experimental system and other “target” systems of interest. The first axis is fairly straightforward but it is worth noting that Diamond is here referring to the human, active,

612 At the same time that Diamond was arguing for plurality in science and the uniqueness of ecology as a science, the same trend was operating in philosophy. This is perhaps due to the fact that Ernst Mayr, whom we encountered in the last chapter, had a significant influence on both fields. (He wrote seminal works in philosophy and history of science, as well as being an important biologist and primary synthesis architect. His influence on early philosophers of science is unmistakable. His influence on some ecologists, e.g. Diamond, is clear from the fact that Diamond cites him authoritatively on plurality and the uniqueness of biology and also that they published together.) Regardless, philosophers and historians of science who came of age in the 1980s were particularly concerned to show that biology was different from other sciences, primarily physics, and that history and philosophy of science should reflect this (see for instance, Beatty 1980). One of the major points of contention was whether biology has laws in the same sense that physics or chemistry has laws—i.e., exceptionless generalizations. One of the leading philosophers to pursue this problem, Robert Brandon, also built on Diamond’s analysis of experiments—maybe unsurprisingly in light of Diamond’s views presented so far (Brandon 1994, 1996). In this section we shall consider his interpretation and elaboration of Diamond’s points.

!249 intentional regulation of independent variables, such as temperature, amount of light, etc.

This is why he lists natural experiments as having no regulation of independent variables whatsoever. Although both natural and field experiments technically involve controlling many independent variables (as far as is possible anyhow), this is often done through site- matching (axis 2) rather than through active intervention by human experimenters. Natural experiments, for example, often involve comparing nearby islands to ensure that the abiotic and biotic factors comprising them are similar—just as we try to ensure that laboratory bottles are identical. In what follows I will be concerned with the first axis, which I will refer to as “experimental control.”

The advantages of experimental control are well known and were briefly mentioned above. It is associated with the statistical design category internal validity: “the result of an experiment E is internally valid if the experimenter attributes the production of an effect Y to a factor (or set of factors) X, and X really is a cause of Y in E.”613 The more control we have over independent variables, the more confident we can be of an experiment’s internal validity.

The other two axes, realism and generality, also warrant further treatment. Diamond says that an experiment possesses “realism” if “there are any natural community and any natural perturbation, even a single one, to which the results of the experiment apply or can be readily extrapolated.” Realism is thus a binary axis; an experiment possesses it or it does not.

An experiment possesses “generality” on the other hand if “it is immediately known to apply to many communities.”614 Diamond’s definition of generality makes it sound as though it too

613 See, Guala 2005, 142; Shadish et al 2002 614 Diamond 1986a, 5.

!250 is binary, but his examples show that he intends this concept to do work by indicating some measure of degree: generality is a function of the number of natural communities to which an experiment’s results can be applied. It is also used comparatively: an experiment is more general than another when its results apply to more natural communities.

We can subsume “realism” under “generality” and apply it to the three types of experiment outlined by Diamond above. Laboratory experiments lack generality; typically the results do not apply to any “natural” community and thus they lack realism. Field experiments are intermediate; typically the results apply to one or a few natural communities and thus they often possess realism as well. Natural experiments possess the most generality; typically the results apply to many natural communities and thus they also possess realism.

The tradeoff I’m interested in is therefore between “experimental control” and

“generality.”615

Table 7.1: Abridged table from Diamond 1986a comparing the advantages and disadvantages of different types of experiments in ecology. (Full table is given in Appendix 3.)

Type of experiment Axis LE FE NE 1. Regulation of independent variables Highest Medium/Low None 7. Realism None High Highest 8. Generality None Low High !

615 For Diamond, then, technically one cannot have generality without realism. There is also something else strange about Diamond’s schema. Generality and realism only refer to an experiment’s results applying to “natural” communities. Thus a laboratory result may apply to many other laboratory-synthesized communities, but would still lack realism and generality.

!251 The easiest way to think about Diamond’s axes, as I said, are as constituting a multidimensional space of experimental types. In this case we are considering a two- dimensional space: “experimental control” on one axis and “generality” on the other. When I claim there is a tradeoff between these axes, I mean that some points in this space cannot be realized. In particular, points in which the two axes are simultaneously maximized. An experimenter can maximize control, or maximize generality, but not both simultaneously.

Diamond intends this to be a general, “intrinsic” feature of experimentation in community ecology (i.e., not a “common but curable” deficiency in experimental design). This tradeoff, as noted above, may not occur in other sciences. Chemists, he finds, “are often initially puzzled as to why ecologists ever resort to any other approach,” as their laboratory experiments possess both experimental control and generality.616

Diamond’s notion of generality calls for one further comment. If divorced from the particular context in which he is writing, it might appear to be uselessly vague or even meaningless.617 He is interested in generality of a particular type, the kind that warrants application to “natural communities.” But one should rightly ask, which natural communities and why? Since this depends on what counts as “natural,” a particular concept of the natural thus plays a foundational role in his assessment of the advantages and disadvantages of experiments. His drawing of the line between artificial and natural is closely intwined with

616 Diamond 1986a, 5. He repeats this example throughout his life, see Diamond and Robinson 2010b. 617 For example he never addresses the following problem. Diamond’s reflections on natural experiments highlight a conceptual ambiguity that I think often accompanies discussions of natural experiments but that I did not delve into above. In Diamond’s case, we might ask, are natural experiments better (in terms of generality) because they allow us to “experiment” on the organisms we’re ultimately interested in building a reserve for and thus provide results valid for those organisms (Diamond’s New Guinea birds are a good example), and/or, are natural experiments better because they can be “performed” on a larger set of different types of organisms or species, supplying results that are invariant across many species (possibly even universally invariant)? Diamond, I say, tends to waver between the alternatives, although they highlight very different types of experimental advantage.

!252 his conservation efforts and associated with ideas about communities that we have not, could not or should not manipulate through experiment or development. From the early 1970s through the 1980s he was centrally involved in designing, implementing, and monitoring nature reserves for terrestrial animals in Indonesian New Guinea (the western half of New

Guinea).618 In many cases the areas proposed for the reserves had never been scientifically assessed—in one case, for instance he “could not name a single species of animal that was definitely known to occur.”619 His natural experiments in Papua New Guinea (the eastern half of New Guinea) provided the only indications of how community structure in the surrounding area would be affected by the imposing development plans. Field and laboratory experiments, while internally valid, would be in this case inapplicable. When Diamond stresses the importance of generality, it is this kind of generality he has in mind. It is his commitment to ecology as an applied science that necessitates that generality enter a discussion on proper experimental practice: he is asking ecologists, especially the Tallahassee ecologists, to widen their notion of scientific rigor to take account of how their research will be applied.620

618 Diamond 1984, 1986b. 619 Diamond 1986b, 501. 620 As Diamond writes, clearly referring to Simberloff, “Scholars working in academic disciplines are driven by the quest for knowledge, independent of external deadlines. Hence one of the cardinal sins of pure science is to leap to a conclusion prematurely, before adequate supporting evidence has been assembled. Thus when pure scientists without practical experience of conservation programs write about conservation, one often hears the caveat that decisions must be based on detailed autecological studies of individual species. However, conservation is an applied science and is subject to different rules from those governing pure science” (Diamond 1986b, 501).

!253 Diamond is not alone in acknowledging that this tradeoff is an important feature structuring experimentation in ecology, or even in biology at large.621 And it is often, as well, expressed in artificial-natural terms. For example, Robert Brandon writes, “The most manipulative population biology studies are those that most tightly control the relevant independent variables. Typically this is only possible in laboratory studies. But it is all too easy to create in a laboratory setting conditions that are nowhere found in nature and that have only dubious relevance to what is going on in nature.”622 Brandon’s quotation highlights a further feature of Diamond’s tradeoff. Diamond, I believe, intends these axes to have more than just a correlational relationship. Field and natural experiments in community ecology possess on average greater generality because they involve less experimental control. To pursue this further let me delve into Diamond’s views on what gives rise to this particular tradeoff in community ecology.

Part of the reason he thinks this tradeoff arises is that communities are large, sometimes endangered, and always complex. Their size and endangered status forces us to seek more tractable, but less realistic, experimental “models,” such as yeast or Drosophila,

621 Ecologist Peter Morin, for example, writes that “Laboratory experiments can be much more precise than field experiments because extraneous factors are readily eliminated or controlled and high levels of replication are possible. That precision comes at the expense of ecological realism” (Morin 1998, 50.). In other words, precision in the laboratory trades off for realism in the field. Ecologist Nelson Hairston similarly draws attention to an experimental choice ecologists must make “between confidence in our results and their generalizability” (Hairston 1989, 121). Philosophers too acknowledge Diamond’s tradeoff. David Rudge writes “that the presence of a trade-off between how artificial an experiment can be and how informative the results are for studies of natural populations is an important difference between experiments in biology and most other sciences” (Rudge 2000, 183). See also Peters 1991, 137-41. 622 Brandon 1996, S452. Brandon goes on to draw a comparison to physics. In physics, the phenomena created in the lab are “highly artificial” and “have existed either rarely or nowhere else in the universe.” But, he says, and again in contrast to community ecology, physics is a law-driven science and thus “no phenomena are too ‘artificial’ to be of relevance to” these universal laws.

!254 and their complexity makes the active regulation of independent variables an uncertain affair.

Diamond tends to refer to such issues as problems of “complexity.”

But complexity is only part of the reason why this tradeoff arises. It is also the case that the processes and patterns under study in community ecology tend to be highly context sensitive; they tend to be connected to particular times, places, situations.623 Just because whales don’t fit into test tubes does not by itself explain why learning about a community of yeast in the laboratory does not tell one about communities of whales, or even communities of “wild” yeast. This has to do more with the variability and uniqueness of biological phenomena, than their complexity. Think of a general question asked in community ecology, for example: What factors limit the species composition of an ecological community? What factors determine species distributions and abundances? Given the nature of the phenomena studied in community ecology, Diamond argues, answers to these general questions will be highly context-sensitive. Answers will not come in the form of universal laws, like those found in physics, but will instead be of the form, “For a community of species with

623 See Brandon 1996 and Waters 2007, 2008.

!255 properties A1 and A2 in habitat B at latitude C, limiting factors X2 and X5 are likely to predominate.”624 Diamond refers to this feature of ecology as problems of “conditionality.”625

For Diamond it is a result of both the complex and conditional nature of the phenomena studied in community ecology that gives rise to the tradeoff between experimental control and generality.626 We should think of Diamond’s concepts of

“complexity” and “conditionality” as providing, albeit only in vague outline, the conditions under which less manipulation is more. While experimental control will continue to increase our confidence in an experiment’s internal validity, when these conditions are met, the results found will not help us understand “natural” communities. Yet this, Diamond tells us, is the ultimate goal.

As a contrast case, consider a quintessential laboratory science like experimental physics—that is, a science in which Diamond believes this tradeoff does not arise. According

624 Diamond and Case 1986, x. This might be contrasted with the Morgan laboratory’s studies of heredity done in the Morgan laboratory that we encountered in the last chapter. Morgan and the fly group were able to discover principles of heredity that held across great changes in context: that are the same, or similar enough, between humans and fruit flies. The case could be made even more easily in physics, where the phenomena discovered and studied often hold throughout the entire universe. Brandon characterized the phenomena under study in ecology as less “projectible” than the phenomena under study in molecular biology or laboratory physics: they do not travel as well from one context to another. Brandon 1996. The term “projectible” Brandon takes from philosopher Nelson Goodman. A different way of thinking about projectibility is in terms of what Ken Waters calls “causal invariance space” and what Woodward calls “causal stability” (see Waters 2008 and Woodward 2010). These two notions are tied more closely to the causation literature. Because of this these notions, would have taken much more detail to explain, and commit one to a particular causal reading of experiments. 625 “Conditionality” of results is something which many biologists, philosophers, and historians have discussed, but in contexts other than experimentation (but see Brandon 1996 and Rudge 1996). These discussions usually trace back to Ernst Mayr’s discussions of what makes biological science unique. One thing that Mayr draws attention to is the universal uniqueness of the entities in biology and the variability this presents biologists with. This variability makes generalizations and even predictions in biology much harder to acquire. Something like this argument is behind Brandon’s notion of “projectibility” and was likely also influential on Diamond’s thinking. See Mayr’s discussions in, Mayr 1988. 626 Diamond and Case present a hypothetical case to bring out the moral: “Suppose for comparison that the genetic code, instead of being determined solely by DNA, was codetermined by seven classes of macromolecules, whose relative role in a given species varied with age, season, weather conditions, and time since the last glaciation and also tended to differ between large and small species, ectotherms and endotherms, and herbivores and carnivores. If this were true, we would surely not have our present complete understanding of the code. Yet this is exactly the problem that ecologists face” (Diamond and Case 1986, x).

!256 to philosopher Nancy Cartwright, it is precisely experimental control which permits physicists to generalize their results beyond the laboratory walls.627 In the laboratory, physicists create ceteris paribus conditions, that is, conditions in which all of the disturbing factors are either removed or controlled for. Under these very special conditions, the phenomenon under study exhibits itself in a very systematic way, a way that allows us to generalize beyond the laboratory to all such situations in which the phenomenon plays a role.

“Situations that lend themselves to generalisations are special, and it is these special kinds of situations that we aim to create [...] in our experiments.”628 According to Cartwright what is revealed in such situations is the phenomenon’s nature: how it acts in isolation or without impediment (she consciously intends this to sound Aristotelian). She writes, for example, of the “Gravity-Probe” experimental team, ! by designing the experiment to ensure that the nature of relativistic precession can manifest itself in some clear sign, by blocking any interference and by opening a clear route for the relativistic coupling to operate unimpeded - to operate according to its nature, by doing just this, the Gravity-Probe team will create an experiment from which it is possible to infer a general law.629 ! How different this is from Diamond’s methodological prescriptions for community ecology. In Cartwright’s presentation of physics there no tradeoff, it is precisely the experimental control which gives rise to the generality of the experimental results by removing the disturbing conditions and allowing the phenomena to exhibit its typical

627 We also considered Cartwright’s views in Chapter 2. 628 Cartwright 1999, 86. 629 Cartwright 1999, 87.

!257 behavior unimpeded. This typical behavior is then carried “from one circumstance to another.”630 Physics, according to Cartwright, partakes of the analytic method: “to understand what happens in the world, we take things apart into their fundamental pieces.” We can then

“carry the pieces from place to place, assembling them together in new ways and in new contexts. But you always assume that they will try to behave in new arrangements as they have tried to behave in others.”631 Underlying her argument is the assumption that physical phenomena are not very context-sensitive, that there exist stable natures to be discovered and to be built into universal generalizations.

In contrast, Diamond’s tradeoff suggests that in community ecology experimental control introduces disturbance rather than removing it. Recall my previous discussions of disturbing conditions; these are factors not included in a representation that, when present in the object represented, impede its accuracy or suitability. For Cartwright, the disturbing conditions must be accounted for and removed by extensive experimental manipulation. In the case at hand, however, experimental control impedes the accuracy of the experimental system to represent a natural system of interest. In community ecology, then, rather than removing disturbing conditions, experimental control is thought to be a disturbing condition.

“Outside the supervision of a laboratory,” Cartwright argues, “what happens in one instance

630 Ibid., 83. See also Carwright’s discussion of what makes the “ideal” experiment in physics, and why such ideal circumstances are also the most artificial (Cartwright 1999, 84ff). It may be better to say that the phenomena under study in physics are more projectible, rather than to follow Brandon. See also Daniel Steel’s discussion of Cartwright in Steel 2008. 631 Ibid., 83. Cartwright’s physics also partakes of the engineering or maker’s knowledge ideal. 632 Cartwright 1999, 86.

!258 is rarely a guide to what will happen in others.”632 According to Diamond, however, what happens in the laboratory is an even worse guide, in community ecology.633

This is again not to say that Diamond sees manipulative experimental work as uninformative. In his own research he uses all three types of experiment: he uses laboratory experiments to study food processing by avian intestine, field experiments to study bower decoration by bowerbirds, and natural experiments to study bird communities in the New

Guinea area. He recognizes the virtues associated with each of these types, but he pushes us to consider the tradeoffs encountered in this science that are not encountered in other sciences. And most importantly, this highlights the poverty of methodological reductionism.

In community ecology, according to Diamond, laboratory experiments inform us about “the possible” rather than “the actual.”634 “Like mathematical theory,” he writes, “LEs [laboratory experiments] take specified simple starting conditions and reveal a range of possible

633 Diamond is even skeptical of the generality of field experiments because of their increased experimental control. “There is an acute question,” he writes, “concerning the extent to which the result of an FE [field experiment] possesses generality and applies beyond that single site in that year” (Diamond 1986a, 5). This is both because of the complexity of ecological phenomena—fences for instance can affect those being fenced-in in at least nine different and unpredictable ways—and their conditionality—field experiments minimize natural variation and thus only apply to the community directly studied and often only in that year. This contrast with laboratory physics warrants one further comment. According to my definition, disturbing conditions only make sense relative to representations—they interfere with the accuracy or suitability of representations. Why experimental control is considered a disturbing condition in community ecology, and yet helps to remove disturbing conditions in laboratory physics might be attributed to a difference in representation, that itself can be attributed to differences of the aims of these sciences. In community ecology, the experimental system is a representation of a natural system, maybe many natural systems, of interest (this is at least what Diamond has in mind). In this case, experimental control introduces disturbance because it tends to make the experimental system less like any, actual natural system. In laboratory physics, however, the experimental system is a representation of no actual system—that according to Cartwright (and Galileo, whom inspires her) would be completely beside the point. Physics is a law-driven science. The experiment is set-up to test a general law, or to determine a generally applicable constant, and thus creates ideal conditions under which the law can be tested. It is thus a representation of these ideal conditions. In such a case, experimental control is precisely what gives rise to a situation in which this law manifests itself without hindrance or impediment. Thus whether experimental control counts as a disturbance or not depends on what the experiment aims to represent. 634 For a much more nuanced discussion of the important roles of simple mathematical models and laboratory experiments in ecology, see Odenbaugh 2005b and 2006. For discussions about biologists being more interested in the “actual” than the “possible,” see Beatty 1986b, Brandon 1994, 1996, Waters 2007.

!259 outcomes, which field biologists can then evaluate for the LEs’ relevance to actual communities.”635 They are thus a starting point, but cannot be the end of inquiry.636

These considerations compel community ecologists to have a keen interest in “the natural” and “the artificial” when designing and performing experiments—or should compel them, if following Diamond.637 Given the complex and conditional nature of the phenomena under study in community ecology, the artificiality of the experimental circumstances matter for the knowledge produced. If one wants to learn about communities of whales, or communities of birds in New Guinea, one is forced to utilize natural experiments since more artificial, and experimentally tractable, substitutes will be misrepresentative.638 It is for these reasons that the artificial-natural distinction has been a useful way to classify the types of experiments found in community ecology.639 Community ecologists, and many biologists in general, are compelled to be keenly aware of the artificial-natural distinction because the distinction has an epistemic bearing that it does not have in some other sciences, in particular,

635 Diamond 1986a, 6. 636 That biologists in general are most interested in “the actual” rather than “the possible” has been called attention to in a number of philosophical discussions: see Beatty 1986b; Brandon 1994, 1996; Waters 2007. We should remember however that all experiments require controls, natural experiments being no different in this regard. Natural experiments employ the strategy of site-matching to this end. However, site- matching does not introduce disturbance according to Diamond, since setting up and performing the experiment does not alter the natural sites in any way. Natural experiments are experiments without active experimental control and it is only this kind of control which introduces disturbance. 637 Brandon says the same thing but chooses “the actual” rather than “the natural.” He also discusses evolutionary biology predominantly. 638 This may also explain why “model” organisms are extremely useful in some areas of biology and yet less so in others. And of course, what Diamond argues is a matter of degree and applies to different sciences to different degrees: the artificiality of the experimental circumstances matters less moving from community ecology, to behavioral ecology, to physiology, to molecular biology, to biochemistry, to laboratory physics. 639 For community ecologists, how artificial an experiment is matters more fundamentally than in some other sciences. This also explains why the “harder” sciences, as Diamond portrays them, do not classify types of experiment according to the artificial-natural distinction. As Brandon says, the phenomena created in physics are “highly artificial, they would have existed either rarely or nowhere else in the universe. But physics, at least certain areas of it like high energy physics, is a science driven by the search for fundamental laws. No phenomena are too ‘artificial’ to be of relevance to it” (Brandon 1996, S452). There is no need to draw the artificial-natural circumstances because what happens under artificial conditions just is what happens in nature.

!260 according to Diamond, the harder sciences. The point of Diamond’s discussion is, in effect, to demonstrate that it makes a difference for the knowledge produced whether the experimental conditions were “artificial” or “natural.” As Brandon said, in ecology, “it is all too easy to create in a laboratory setting conditions that are nowhere found in nature and that have only dubious relevance to what is going on in nature.”

This further suggests why the artificial-natural distinction itself matters more to some sciences than to others, and thus arises in the discourse of some sciences more than in others.640 The reasons for this are tied to the nature of the phenomena under study in such sciences as well as the goals of their inquiry (which are, obviously, conditioned by the phenomena studied).641 We should, in fact, see Diamond’s “complexity” and “conditionality” as also providing, again in vague outline, the conditions under which the artificial-natural distinction is going to matter in a science. Sciences which work on small, simple, isolatable, and less context-dependent phenomena and which strive for universal laws will not require or express the distinction in the context of experimental practice because human manipulation is less problematic (and, according to Cartwright, necessary). In sciences like community ecology, which work on complex and conditional phenomena and whose generalizations typically hold over only a small subset of the natural world, human manipulation will tend to

640 In Chapter 3 I argued that the distinction plays a much more dominant role in the discourse of those areas of biology “furthest” from physics. In this chapter we have seen both examples of this and reasons for it. The example is that the distinction is not used in these other sciences to classify types of experiments, nor is it considered to be a particularly important factor in deciding what experiment to perform or in structuring experimental practice. In fact, if the distinction matters at all in these sciences, it is probably for the exact opposite reason: as Cartwright says, for laboratory physics “natural” phenomena are unruly and uninformative, and thus the more artificial the better. This also points to a further similarity between the artificial-natural distinction and the normal-pathological distinction. In physics, there is no normal-pathological distinction because there are no “pathological” states because there are no “normal” states. A similar thing might be said about artificial and natural: there are no “artificial” states in physics because its all just a “natural” state. 641 And there is certainly a feedback between the two.

!261 obscure what researchers wish to know. The artificial-natural distinction gives them a way of expressing this problem and suggests a language they can adopt to navigate solutions. All this is to say that, in such sciences, less manipulation can be more. !

CONCLUSION: ARTIFICIAL AND NATURAL IN HARD AND SOFT SCIENCE

In summary, behind Diamond’s frustrations with the National Academy of Sciences elections of 1987 and the unfortunately ingrained labels “hard science” and “soft science,” were concerns about the undecided future of community ecology and about a method he had found particularly revealing: the “natural” experiment. When this method came under direct attack in 1975, Diamond defended it by drawing attention to intrinsic experimental differences among experimentation in the sciences. His interlocutors had mistakenly assumed that experimental methods were valid only insofar as they were based upon “proven” methodologies in the physical sciences. Experimentation in community ecology is difficult, and thus intellectually stimulating, he argued, precisely because it faces tradeoffs not encountered in the “harder” sciences. Specifically, it faces a tradeoff between experimental control and generality. What natural experiments lack in experimental control, they gain in generality, and are thus the method of choice for supplying results applicable to endangered ecological communities, such as bird populations of New Guinea—“natural” communities in

Diamond’s parlance. Given Diamond’s environmentalism and the applied nature of his work, it is easy to see why this was important to him. Could laboratory experiments or even a host of field experiments provide adequate grounds for environmental decision-making when they

!262 tended to lack generality? When they studied phenomena isolated from their “natural” times, places, and environments? Diamond’s answer was a controversial, “no.”

Throughout this story concepts of the artificial and the natural have entered at multiple levels and played various, important roles. Most conspicuously, they entered as a way to classify the types of experiments done in ecology and to constrain the application of types of experimental knowledge. Natural experiments were those which provided results most applicable to “natural” communities. This object of study itself (obviously) depends on an assessment of what is to count as natural. As we observed above, this concept is an ideal.

For Diamond, this concept at times means those communities not experimentally studied and that exist in the absence of human disturbance, but more often than not, it refers to only a subset of those communities, to the type of “wild nature” some believe we ought to preserve and protect. This ambiguous notion of nature has obvious rhetorical value: few community ecologists in the 1980s would dare be caught studying systems with no connection to nature whatsoever, even though what is to count as a “natural community” is rarely specified.

Diamond’s tradeoffs thus tapped into this rhetorical value. Simberloff and colleagues’ continuous references to Diamond’s “nonexperimental observations,” as opposed to “natural experiments,” was an attempt to remove any rhetorical power associated with the words

“experiment” or “natural.” Finally, in the background of this entire research looms the natural laboratory paradigm, the idea that nature is a laboratory. While this might seem to collapse the distinction, it reappears at a more basic level. For Diamond, when it came to solving the problems of conservation biology, nature’s laboratory was distinct from and advantageous to human laboratories.

!263 While there is something correct about Diamond’s analysis, there is something awry or problematic about its rhetorical character. It is certainly true that when making risky environmental decisions under significant temporal, economic, and political constraints, it is better to rely on organisms and environments that are most closely related to those of interest, even when their study may be considered less scientifically rigorous. It is better to rely on less manipulative experiments with birds on eastern New Guinea, for instance, than laboratory or field experiments with yeast, when designing a nature reserve for birds on western New Guinea. However, it is locating these scientific issues about context- dependency and generality within the abstract notions of “artificial” and “natural” which, while commonplace, adds the rhetorical but problematic flavor. It suggests that it is because of their artificiality alone that knowledge of “artificial” communities is invalid when applied to “natural” communities, when in reality this has to do with relevant similarities between the experimental and “target” communities. The particularities of each case are in effect masked by these abstract concepts.

In this chapter humans are treated as disturbing conditions in a way both similar to and different from their treatment in previous chapters. Let me start with the similarity.

Humans are disturbing conditions in this case because human experimental control is the factor which impedes (rather than facilitates) the representativeness of the experimental system. Natural experiments are a way of trying to retain the power of experimental work while at the same time allowing as little human interference as possible. Again, humans are,

!264 in this sense, treated as the “outside” factors which, when present, are thought to be grounds for calling into question the validity of the results.642

The difference might be put like this: in previous cases, humans were considered disturbing conditions because they were seen as “outside” of nature (or natural nature), and thus outside of the natural processes under study. This was so during the 19th century when humans were not considered part of the “state of nature.” It was also so for Dobzhansky, although maybe to less of an extent, what he worried that even “wild” or free-living

Drosophila melanogaster were artifacts—in this sense, our artifacts spread far abroad, metaphorically speaking, into what we would otherwise consider the natural world. In

Diamond’s case, however, human involvement, in and of itself, does not impinge on the validity of experimental results. Natural experiments can make use of human-mediated events, such as when the peacock bass was accidentally released by a business man into a tributary river and proceeded to spread throughout Panama’s Gatun Lake.643 Humans become disturbing conditions, for Diamond, in the context of experimental control. It has, in this case, less to do with human actions and more to do with features of the ecological phenomena studied themselves.

This point is perhaps best made by returning to something that was covered in

Chapter 2. The collapse of the distinction on ontological grounds only opened the door for the distinction to be drawn on epistemic grounds. For Diamond, artificial states are only one set of possible states of nature, but they are not the ones we’re interested in, nor are they

642 Here I have in mind external validity, not internal validity. This distinction is however rarely drawn by ecologists, probably because results without external validity are simply considered invalid tout court. 643 See Diamond 1986a, 13. Also Zaret and Paine 1973. The lake itself was artificially created in 1907 with the building of the Gatun dam.

!265 often representative of “natural” states. Human actions, including experimental control, represent only a small and idiosyncratic part of a context-dependent ecological world. To learn about communities of New Guinea birds, we have to use those communities—or ones as closely related as possible—because inferences from other, more remote, experimentally isolated, ecological systems will be unreliable. Humans are disturbing conditions, not because ecological nature is too “unruly,” as Cartwright says of physical nature, but because it is too context-dependent and too complex; there are no stable natures to be discovered.

And so we see, even in experimentation, the knowledge-making practice that supposedly accompanied the collapse of nature and artifice so long ago rendering possible the rise of modern science, the categories of “natural” and “artificial” remain in opposition, continuing to be essential for the actors themselves.

!266 ! ! CHAPTER 8 ! Conclusion ! ! ! ! ! ! ! ! The impulse behind different manifestations of interest in Nature and Art appears to be a fundamental gesture of the human spirit that has, at various times and places, found diverse channels for its expression. Edward Tayler, 1966, Nature and Art in Renaissance Literature !

WE (AT LEAST MANY OF US BIOLOGISTS) HAVE NEVER BEEN ARTIFICIAL

According to historian Philip Pauly, Jacques Loeb was one of the first biologists to decide that nature was fading away.644 That it was becoming trivialized. This, Pauly speculated, foreshadowed the deterioration of its authority as a concept central to scientific thought and practice in the 20th century. Loeb advocated a blurring of the artificial-natural distinction. “Natural” states of affairs have no privileged place, he said, among what we should study or what we should aim to create. It appears that Loeb’s ideas were taken up by many in biochemistry, molecular biology, genetic engineering and synthetic biology. Some envision a future in which “Building a house would entail no more work than planting a seed in the ground.”645 How far we are from Aristotle’s oak seed—the quintessential example of

644 Pauly 1987, 199. 645 For example, see the popular book by G. Church and E. Regis, Regenesis: How Synthetic Biology Will Reinvent Nature and Ourselves (2012).

!267 natures as non-human ends. This idea, that the nature, in Aristotle’s sense, of an engineered seed would be to develop into a house, presents a striking challenge not only to Aristotle’s understanding of artificial and natural but also our own.

Yet, in other areas of biology, in particular evolutionary biology and ecology, classifications according to the artificial-natural distinction are still commonplace, have deep historical roots, and structure the way biologists do their research and decide what counts as valid for biological study. The argument of this dissertation has been that in these areas of biology decisions concerning the choice and evaluation of objects, and evidential and experimental practices, as well as the assessment of styles of scientific persona, reflect the artificial-natural distinction. To the extent that we understand science to be knowledge- making through the investigation of phenomena according to specific methods and practices done by individuals, these decisions, and thus the distinction framing them, shape biological science. This argument undermines a commonplace view of contemporary science according to which the collapse of the artificial-natural distinction was a prerequisite, and according to which the distinction lacks significance or relevance. I have argued, instead, that this distinction is not an outdated, pernicious relic left-over from antiquated anxieties, but has continued to exert a significant influence on scientific practice, thus meriting philosophical and historical treatment. We cannot simply dismiss it with a tinge of nostalgic regret, a biologist’s thought and activity are incomprehensible without these concepts.

To say that because the distinction is being undermined, it is therefore fading away is to forget that the distinction was problematic to begin with. It has always been contested

!268 terrain, and our idea of nature has always been “impregnated with artifacts.”646 As Robert

Kohler pointed out, just because we understand the sharpness of the distinction to be somewhat dulled, we need not conclude that it is unimportant, especially for biologists in the field. Instead we should ask why the distinction remains important to these biologists, given that we see many problems with its coherence. What’s real is real in its consequences, and in evolutionary biology and ecology, the distinction has notable consequences.

The goal of this dissertation has not been to address a single question nor to offer a single culminating narrative. The argument was made ostensively, by providing examples which show these points to be true of at least some biologists and their science. The result has been a problematization of these concepts: showing that they have an influence on practice that is hidden by their familiarity and deep historical roots in sciences like evolutionary biology and ecology unlikely to be unsettled. If it is true that the artificial- natural distinction is being challenged on more fundamental grounds now than it has in the past, then the future of disciplines like evolutionary biology and ecology, which treat “the natural” as epistemically privileged, will be an interesting one.

In the introduction I drew attention to seven conclusions which were supported by the analyses in this dissertation. I would like to end with these as well: (1) Invoking the artificial- natural distinction is routine in biology and has been central to recalcitrant debates.

Biologists, of the stripe considered here, have a habit of approaching their discipline and its puzzles with these two alternatives in mind. (2) There is a standard historical narrative that relates the dissolution of the distinction to the rise of modern science, and that serves as a

646 Quote from Merleau Ponty in 1956; quoted in Bensaude-Vincent 2007, 293.

!269 touchstone for discussions of the artificial-natural distinction in science. This narrative is highly problematic and cannot accommodate the life sciences. In the life sciences this distinction has been central to its modern history. (3) When this distinction is drawn, humans are treated as, what philosophers call, “disturbing conditions.” (4) The concepts “nature” and

“natural,” in the cases considered here, do not refer to a physical place, but to an ideal state involving no human disturbance. (5) Artificial things are often treated as lacking the reality of natural things, and this is because the former are in some way human disturbed. “The natural” is often, for this reason, given a privileged epistemic position in biological studies: it is a crucial feature of the proper object of knowledge. (6) Underlying the artificial-natural dichotomy are normative concerns about which objects and methods provide valid biological knowledge and which do not. This entails that classifications according to the artificial- natural dichotomy are not innocuous; they are consequential. (7) Finally, these concepts, as well as their relation to one another, have been in the process of continual reconceptualization. As a result phenomena, practices, etc., once deemed artificial may no longer be so called. This changes how biologists act towards them. !

LIMITS OF THE DISSERTATION

It should go without saying that the full range of uses of “the natural” and “the artificial” cannot be dealt with in a single dissertation. I would like, furthermore, to point out four major shortcomings of this dissertation research. First, the concept “natural” is the antithesis of many concepts besides “artificial.” These include most importantly, “unnatural,”

“cultural,” “nurture,” and “synthetic.” Each of these distinctions has a complicated history

!270 and a literature devoted to its study.647 These antitheses of nature differ, sometimes importantly, from one another. In this dissertation I deal primarily with the artificial-natural distinction, along with those cases in which concepts like “synthetic” are used as synonyms for “artificial.” Future work is needed to draw any substantial conclusions about the relations between the antonyms I have considered here and nature’s other antitheses.

In the later chapters I opted for detailed analyses of particular actors at the expense of general analyses of the fields in which they work. Both of these scholarly orientations obviously have advantages and disadvantages. The disadvantage of my analysis is its substantial reliance on generalization from individuals to wider trends. I use Dobzhansky,

Darwin, and Diamond as models and draw general conclusions from my analyses of them. As with all models, there always remains a possibility that the results are idiosyncratic and unrepresentative of larger patterns. I have tried to situate these models within their larger contexts to indicate that such a possibility is unlikely.

As mentioned in the introduction, the cases I consider are biased towards those in which, when the artificial-natural distinction is drawn, “natural” corresponds to good, valid, or advantageous and “artificial” to bad, invalid, and disadvantageous. As I’ve said, I made this decision for three reasons: (i) my primary interests are in the research practices of ecology and evolutionary biology, and in these disciplines the distinction is typically drawn in that way; (ii) I want to complicate a trend in historical-philosophical research that focuses on the rise of laboratory science in the 20th century and the unnaturalness of modern science;

(iii) I simply have insufficient space, time, and mental capacity to do justice to all subfields

647 See Daston 1998.

!271 of biology. I fully admit that this bias has, if anything, only taken away from the full quality of a much larger, and still interesting, project.

I have in general steered clear of dealing with other areas of biology, such as synthetic biology or molecular biology, other natural sciences, and the design sciences (as Herbert

Simon characterized them, architecture, engineering, artificial intelligence, etc.). Each of these areas would accent my accounts of the relationship between the artificial-natural distinction. I have also been forced to leave out other areas of evolutionary biology and ecology, such as invasive species research, in which the artificial-natural distinction plays an important role. !

THE FUTURE

This project points to much future work, not least because it is very much an unfinished story. I will here detail just two future projects. The first could be considered within the scope of the dissertation, the second is more an outgrowth.

(1) In Appendix 1 I briefly consider evolutionary biologist Ernst Mayr’s classification of biologists according to the artificial-natural distinction—“naturalists” and

“experimentalists”—and how these classifications relate to the classification of research organisms, like fruit flies, as artificial and natural objects. I plan to extend this research further into the context of the increasingly hostile relations between molecular biologists and organismal biologists throughout the 1960s, especially as it occurred within the biology department at Harvard, ending in the splitting of that department along precisely those lines.

This will help extend the project into other areas of biology, as well as to other areas of

!272 analysis, since the classification of humans is importantly more complicated than that of objects. The guiding questions of this research will be: (i) what did it mean to be a

“naturalist” in the 20th century, and (ii) how were the categories “naturalist” and

“experimentalist” used to express, dictate, and partition the legitimacy of particular groups of scientists for gaining particular types of knowledge? I plan to use philosopher Ian Hacking’s idea of “interactive kinds” and “looping effects” to aid this analysis.

(2) The second project is more straight-forwardly philosophy of science.

Experimental practice in sciences like ecology and evolutionary biology often involve heeding tradeoffs between different experimental types, as we saw in Chapter 7. The nature of these tradeoffs has, however, received little philosophical attention. This is surprising given that, firstly, tradeoffs in model- and theory-evaluation are mainstays of current philosophy of science, secondly, that, historically-speaking, the analysis of tradeoffs in experimental practice was a direct response to proposed tradeoffs in model-building, and, finally, that these experimental tradeoffs are commonly drawn on to explain methodological pluralism and to underwrite arguments for differences among the sciences. In one set of papers, inspired by Chapter 7, I focus on the types of criteria that give rise to these experimental tradeoffs in order to determine why they are felt more strongly in some sciences than in others. I argue that they depend on (i) our knowledge of the natural system we are investigating, (ii) practical limitations of our laboratory apparatus/environment, (iii) the goals of our research, and (iv) the nature of the natural system itself. In another set of papers I am again concerned with the nature of experimental practice and evidence, this time through a single specific research question: is the “founder effect” evolutionarily significant? The

!273 founder effect is a famous and controversial phenomenon in evolutionary biology that is used to explain speciation. Few other phenomena in biology have been tested in such a variety of ways, put to such scrutinizing examination, reformulated so many times, and yet still remain almost as controversial as when initially proposed. I plan to use the history of experimental tests of the founder effect—many of which were performed by actors in this dissertation—to help shed light on philosophical questions about experimentation, such as how biologists navigate the evidential and methodological diversity which characterizes their science and how they deal with discordant evidence.

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!304 APPENDIX 1 A Note on the Artificial-Natural Classifications of Persons ! Although the case study presented in Chapter 6 was primarily about the classification of fruit flies and biological processes, there was another very important type of classification operating as well. This is the classification of types of persons, more specifically, types of scientists: naturalists and experimentalists. In Chapter 3 I explained this classification as also drawn in artificial-natural terms. Experimentalists study nature under artificial circumstances, whereas naturalists study nature under natural circumstances; or as Dobzhansky simply explained, “Naturalists study nature.”648 As I said, although “naturalist” and

“experimentalist” are both actor’s and historian’s categories, the latter are going to be my focus here.649 In what way is such a classification of persons consequential? In what ways does it also involve normative as well as descriptive dimensions? Let me just give one pertinent example.

The historical study of the evolutionary synthesis period was stimulated by a set of workshops in 1974 and organized by Ernst Mayr and the historian William Provine. One of the central agendas of this conference was a classification of historical actors according to the naturalist-experimentalist dichotomy because the collapse of this dichotomy was thought to be central to the synthesis. It was important to the conference attendees to try to figure out

648 Dobzhansky 1966. 649 The former involves the more complicated problem of, what Ian Hacking calls, looping effects, “classifications that, when known by people or those around them, and put to work in institutions, change the ways in which individuals experience themselves—and may even lead people to evolve their feelings and behavior in part because they are so classified” (Hacking 1999, 104; Hacking 2001, Chapter 6, “Making Up People”).

!305 which actors fit into which category, and principally which actors actively acted to collapse the dichotomy. Dobzhansky, as I have said, was seen as centrally involved in this collapse, and he himself endorsed this. But who were his precursors? What about his earlier Russian training? Did it put him in a position to see the importance of an evolutionary perspective on genetics and to work on natural populations, when such things were considered “old- fashioned”?

One of Dobzhansky’s precursors and mentors, who has been mentioned above, was

Sergei Chetverikov. Chetverikov’s story is a tragic one. It is (and was during the 1974 conference) often claimed that a paper he wrote in 1926 should have been one of the founding papers of the modern synthesis. His paper, more decisively than those that came before him, showed that mutations were not artifacts of the laboratory environment, but existed in nature in the form of recessive alleles—what Dobzhansky called concealed genetic variation. Such historical counterfactuals are hard to assess. What actually did happen was that Chetverikov had an influence on some Russians and Europeans when he gave a talk at the International Congress of Genetics in 1927, including Dobzhansky who cited him repeatedly, but his papers were not translated into English until 1961, he was arrested under mysterious circumstances in June 1929 and never again undertook serious research, and as a result he remained largely unknown in the relevant literature till his death in 1959.650 His name was a common occurrence at the 1974 evolutionary synthesis conference and many of

650 Adams 1980, 268-9. Stern and many other Europeans would have seen him speak in 1927 at the Fifth International Congress of Genetics in Berlin. See Chetverikov 1961 for a translation of his paper and Michael Lerner’s instructive introduction and intellectual biography. See Adams 1980 for an early history of Chetverikov in the Russian context.

!306 the attendees saw him as one of the first to cross the naturalist-experimentalist divide (one of the first to instigate a synthesis).

This assessment was phrased in a particular way, however. It was Chetverikov’s initial training as a naturalist that allowed him to overcome the problems associated with the experimentalist tradition which he later joined—namely that experimentalists misunderstood natural variation because they didn’t have an appreciation for working outdoors with unstandardized nature. To some, it was his being a naturalist that made him, in this sense, a better biologist. The historian William Coleman was observant of this trend. Consider the following abridged conversation about Chetverikov from 1974 between Coleman and

Dobzhansky: ! Coleman. [...] let me turn briefly to one of the dichotomies which have been discussed here at great length. [...] I’d like to speak to the theme of naturalists and experimentalists and suggest that this distinction was perhaps not as clear in the writers of the teens or twenties as it seems to us as historians. [...] I cite two passages from Chetverikov [...] [One] declares: “I repeat that by means of laboratory treatments we cannot produce mutations and consequently the reverse is not dependent on artificial conditions of investigation.” One way you could look at this is, of course, in terms of the artificial production of mutation; another, however, is in terms of Chetverikov’s own values, namely, that if you can’t produce mutation artificially (which is what you really want to do if you want to understand it) then you must go out and look in nature. Thus, you could read Chetverikov’s approach as being a turning to natural populations as the last resort, once the experimental approach had failed. Now that’s forcing the issue; I can see it by the shaking heads. [... However, these] passages suggest to me that in Chetverikovs mind you don’t have a sharp division between what the naturalist does and what the experimentalist does. [...] the distinction between one and the other is really a judgement which we have imposed with hindsight.

!307 Dobzhansky. I believe in Chetverikov’s mind, artificial production of mutation was decidedly interesting and important, but for his particular theoretical construct finding artificial mutations would be totally inadequate. [...] So for Chetverikov and his group finding mutations in nature was something else. It was finding the existence of mutants in natural populations which was regarded as a test of their theory. Ten times as many mutations induced by x-rays would not do. Coleman. The question is not the production of a mutation, but one of its detection. This is what Chetverikov is talking about. And he points out time and again that you don’t detect there simply by observing organisms in nature; you have to bring them back and work with them in the laboratory. [...] that the activity of the experimentalist and of the naturalist are not really as sharply defined as they might seem to some of us is because the detection process itself is a laboratory operation, not one carried out among natural populations. Dobzhansky. No doubt-detection has to be made in the laboratory [...] But it was detecting mutants in the progeny of population samples brought from nature quite emphatically, deliberately, and consciously which those people regarded as a test of their theory. The fact that they were detected in the laboratory - that meant nothing. Where else could they be detected? [...] Coleman. [...] why are we reluctant to call the latter operation an experimental operation, labelling Chetverikov, in a value judgement, calling him a naturalist651 ! There are two points to be made about this transcript. The first is that Dobzhansky’s assessment of what Coleman says about Chetverikov fits well with our above analysis of

Dobzhansky himself. To Dobzhansky, what makes Chetverikov a naturalist, rather than an experimentalist, is that he focuses on what nature had wrought. It doesn’t matter that he uses the laboratory, a “controlled” environment, as a way to reveal the underlying natural mutations. The significant thing about Chetverikov’s work, what makes him fundamentally a naturalist, is that the focus is to reveal phenomena created in nature; only those phenomena

651 From 1974 Evolutionary Synthesis Conference Transcripts, APS.

!308 can count as a “true test” of his theory.652 That Chetverikov had a great influence on

Dobzhansky’s own thinking in this regard is significant.

The second point relates to the classification of persons. Coleman recognizes that calling Chetverikov a “naturalist” is a value judgment, not just a description of his research.

It’s not even clear to Coleman what Chetverikov truly thought or how he should be classified based on his practice; as he suggests, passages from his paper may be read in multiple ways.

What if Chetverikov only turned to nature because he technically or practically couldn’t do what he wanted in the laboratory? This seems unlikely given historical work on the Russian school of genetics, but regardless, the point is that at this conference a value judgment was being made about Chetverikov. He was first and foremost a naturalist and thus was (or should be) an important contributor overcoming the deficiencies of the experimentalist tradition, and thus was a good evolutionary biologist. That’s Coleman’s point.653 And it may have been made about many other reflections of the synthesis architects themselves, such as

Dobzhansky’s comment that H. J. Muller was “rather naive” in his pronouncements about evolution because he had “no familiarity with organisms as they live outside,” or Stebbins’

1942 recipe for making an evolutionary biologist, in which the necessary first ingredient is an appreciation of nature in the form of “teeming tide pools through green woodlands, lush ponds, and orchid-filled mountain tops.”654

652 Another reason why above I have said that the naturalist tradition is better represented as being opposed to human disturbance. 653 Mark Adams’ rejoinder is on the mark: “It may well be that Bill’s [William Coleman’s] point is absolutely on target. Chetverikov was neither a pure experimentalist nor a pure naturalist and that’s one of the reasons why he’s been pointed to as one of the founders of the synthetic [theory of evolution] .... (Laughter)”. 654 See Dobzhansky’s discussion of Muller above and the footnote accompanying that discussion.

!309 Just as with D. melanogaster and D. pseudoobscura, the classifications biologists and historians make of their predecessors according to the artificial-natural distinction involve complicated normative-descriptive dimensions.655 These classifications, in other words, go hand-in-hand with scientific authority and legitimacy. The artificial and the natural is used to endow certain people, objects, practices, and processes with scientific authority and validity.

655 For future work, one might look into Hacking’s differences between classifying things and classifying people. See Hacking 2002, Chapter 6, “Making Up People.”

!310 APPENDIX 2 What is a Natural Experiment?

There is a sense in which the natural experiment is an oxymoron. Experiments by common definition are artificial, not natural. Consider again historian Garland Allen’s definition of experimentalism: “Experimentalists are interventionists, the experimenter deliberately altering the natural condition to observe whatever the consequence might be. [...]

A common assumption behind experimentation is the conviction that intervention into the workings of the organism, although artificial, still reveals something about natural or normal states of nature.”656 Historically, this is how many have understood an experiment. The natural experiment is an attempt to see things, in a way, less anthropocentrically.657 The proponent asks, mightn’t the types of situations we construct in the lab be found naturally occurring? The answer will be aided by considering that these different types of experiments really grade into one another. Let us think a little more about the characteristics of experiments.

Robert Brandon offers two contrasts that are helpful: (i) that between experiments and observations, and (ii) that between experimental and descriptive work. The key to the first

656 Allen 1994, 90. See Chapter 2. 657 An interesting question arises here. What is experimental about a natural experiment? One answer to this question would be to focus on the rhetorical value of describing something that is not experimental as an experiment. Experiments are seen as rigorous science, as we saw above, and so it might be suggested that Diamond’s methods have little to do with experiments at all. He is, it might be suggested, calling them experiments to exploit the positive connotations accompanying that word. This might be further supported as well by the fact that Diamond calls his approach “natural experiments,” whereas Simberloff frequently refers to Diamond’s work as nonexperimental or observational. Some ecologists after Diamond see his calling comparative methods, natural experiments, unfortunate because they do not see them as experiments (see for instance, Lubchenco and Real 1991). Usually this is based on an assessment of experiments as necessarily involving intention on the part of the experimenter, or extensive human-mediated control. Neither of these I see as necessary to experiments. I agree with Brandon’s assessment given below.

!311 contrast is manipulation: in an experiment the experimenter manipulates nature in some way, whereas in observation the experimenter observes nature as is. Brandon uses manipulation in a restricted way: a manipulation involves the deliberate alteration of phenomena by human experimenters and not just “handling.”658 As Brandon puts it, “If our response variable—that which we measure and record in an experiment—is thought of as being a function of some number of independent variables, then changing and/or controlling the values of some or all of the independent variables is what constitutes a manipulation.”659 “Manipulation” is thus a category that covers both holding background conditions, such as temperature, constant and performing particular experimental perturbations, such as introducing a competitor species into an enclosure.

The key to the second contrast—that between experimental and descriptive work— corresponds to a distinction between testing hypotheses and measuring the values of parameters: experimental work involves the former, descriptive work the latter.660 As

Brandon notes, this contrast is orthogonal to the first, giving us four possibilities for evolutionary study (if one thinks of the contrasts as dichotomies): manipulative hypothesis testing, nonmanipulative hypothesis testing, manipulative description, nonmanipulative description.661 Since both of these contrasts admit of degrees (i.e., are not strict dichotomies) the possibilities for evolutionary study are better accommodated by the following figure

(Figure 3).

658 The advantage to this is that the preparations that go into dissections or gel electrophoresis do not could as manipulations. 659 Brandon 1994, 61-2. 660 That the role of experiments is to test hypotheses is a somewhat simplistic view of experiments (see Weber 2012). However, for the main purpose of this chapter I can abstract away from these details. 661 Brandon 1994, 63.

!312 !

Hypothesis test

Laboratory zone Field zone Natural zone

Experimental

No test

! Not manipulate Manipulate Figure 7.3: Experimental Types. Modified from Brandon 1994. (© Springer-Verlag) ! In the upper right corner we would find studies that test hypotheses and that are highly manipulative (e.g., laboratory studies), whereas in the lower left we would find studies that do not test hypotheses and are non-manipulative (e.g., descriptions of fauna). In between lies everything else.

This characterization is helpful because it highlights an important difference we have already noted between the types of experiments to which Diamond refers. In the above diagram, I have indicated where these types of experiments can be found by specifying

“zones.” The borders between these zones are intentionally vague. Let us start with the laboratory types. These are the most artificial because of the degree of experimental manipulation involved: in these types of experiments biologists seek answers to questions

(Brandon’s “test hypothesis”) by manipulating the natural world in terms of controlling some variables and perturbing others. Near this end of the spectrum we might find so-called

!313 “bottle” experiments. For example, Thomas Park’s famous studies on competition in the flour beetle—an experimental system he tellingly referred to as his “organic machine”—would find a home in the laboratory zone.662

In the middle, involving less manipulation, we find field experiments. Finally, at the top left of the diagram, involving the least manipulation, we find natural experiments. The lesser degree of manipulation involved in field and natural experiments does not render them non-experimental; they are not the same as descriptive natural history (the bottom left corner).663 Natural and field experiments are still experimental and thus take place in and require controlled circumstances (as do all “good” experiments), even if this control is (i) less extensive and (ii) not directly provided by humans, that is, not manipulation in Brandon’s sense. The controls, and sometimes even the particular experimental perturbation, are provided instead by the way in which natural sites are arranged. It is this sense in which natural experiments push us to see experiments less anthropocentrically. When Diamond used natural experiments to understand community structure, he took advantage of natural systems that provide the type of arrangement amenable to experimental work—otherwise they would not be very useful as experiments. In his 1975 paper, for example, he lists the properties required to test his hypotheses about community structure, and shows how his

New Guinea “field situation” meets these desiderata.664 To reiterate, field and natural experiments are in the ideal case analogous to laboratory experiments in every way except that the former have been set-up by humans and involve human manipulation, whereas the

662 Park 1962, 1369. So-named after studies in community ecology that used small arthropods kept in bottles. 663 In fact, early in the 20th-century they were part of an approach called “scientific natural history” (Kohler 2002). 664 Diamond 1975b, 348.

!314 latter are natural systems discovered by humans and depend on “Nature” doing the manipulating (or at least this is often the goal).665

665 Interestingly, the authors of a famous laboratory test of the so-called “founder effect” in speciation using Drosophila simulans referred to the population cages used to house the fruit flies as “virtual islands” because of the reproductive isolation between cages; see Ringo et al 1985. The discourse of islands thus works its way into laboratory practice just as the discourse of test tubes works its way into field practice.

!315 APPENDIX 3 Diamond on Experimental Tradeoffs ! This appendix offers a more detailed account of Diamond’s tradeoffs. In a review well-known to every evolutionary biologist and ecologist, Jared Diamond (1986) outlined the advantages and disadvantages of laboratory (LE), field (FE) and “natural” (NE) experiments in community ecology. Diamond split natural experiments into two types: natural trajectory experiments (NTE) and natural snapshot experiments (NSE). The former are “comparisons of the same community at various times before, during and after a witnessed perturbation by nature or by humans other than ecologists.” The latter are “comparisons of communities assumed to have reached a quasi-steady state with respect to the perturbing variable.”666

Diamond’s goal was to demonstrate that different experimental approaches had inherent advantages and disadvantages, and thus to see one approach as the correct important was shortsighted and misguided. His paper was an attempt to specify some of the important tradeoffs between different types of experiments, but since laboratory experiments were the methods par excellence of physics and molecular biology, to which he was reacting, much of the originality of his review is in his developing the important advantages of the other types of experiments. Diamond claimed that there were “eight axes for trade-offs” between different types of experiments (summarized in Table 2; the ninth I have added from a more

666 In a recent analysis Mary Morgan has drawn a distinction very similar to Diamond’s with regard to natural experiments in economics (Morgan 2013). She refers to “Nature’s experiments” and “natural experiments.” The former are like Diamond’s NTEs whereas the latter are like his NSEs. I think there is, at least in ecology, less of a distinction between these types than Morgan suggests. Thus the advantages and disadvantages of these types that she outlines in her paper I would argue are much less absolute than she suggests.

!316 recent article). Above I only dealt with a small subsection of these. More precisely, they can be explained as follows: ! (1) Regulation of independent variables: What we measure and record in an experiment is called our response variable. It is a function of a number of independent variables that we change or control. We attempt to determine which of the independent variables caused a change in the response variable by, e.g., holding all but one constant. Diamond calls this the regulation of independent variables. It is active and performed by a human experimenter. Diamond argues that LE far outweigh FE and NE on this axis. FE regulate only a few variables, whereas NE regulate none. (2) Site matching: To compensate for lack of regulation of independent variables, we often try to minimize intersite differences between experimental replicates, that is, between control and treatment sites. This is done through three methods: replication of sites, selection of sites that have the same values of unregulated variables, and randomization of control and treatment. Diamond argues that matching is least successful in NE, and most in LE. FE again lies in between. (3) Ability to follow trajectory: We often follow trajectories in treatments after the initial perturbation. All three types of experiments allow for the following of trajectories. (The “maybe” entered for NE in Table 1 is because sometimes NEs are just comparisons or “snap-shots” of a particular time.) (4) Maximum temporal scale: To follow the trajectory of experiments it is required that we exert ongoing effort to maintain the experimental system. LEs and FEs cost time and money to continue for long periods of time. Diamond argues that NEs will outweigh the others in this respect. (5) Maximum spatial scale: The physical size of the experimental system. Diamond argues that LEs suffer the most in this regard (the size of a controlled laboratory room is quite small); FEs can be larger; NEs can be very large, basically unlimited. (6) Scope (range of perturbations): The particular perturbations that can be applied to experimental systems. Giving, e.g., practical, legal and ethical constraints, Diamond argues that LEs and FEs are outweighed by NEs on this axis.

!317 (7) Realism: Are there any natural communities to which the results can be extrapolated. If yes, then the experiment possesses realism. LEs are unrealistic by intent. FEs and NEs possess high realism. (8) Generality: The number of natural communities to which the results of the experiment can be extrapolated. Same as Realism for LEs. FEs do better but are far outweighed by NEs in terms of generality. (9) Exploratory/discovery: This advantage comes from Diamond 2001. There he contrasts types of experiments with regard to whether they can give rise to interesting questions not yet considered and that were outside the bounds of the initial research question. He argues that NEs have this as an advantage as well. ! Table A3.1: Full table from Diamond 1986a comparing the advantages and disadvantages of different types of experiments in ecology. See Diamond 1986a for details.

Type of experiment

Axis LE FE NTE NSE

1. Regulation of independent Highest Medium/Low None None variables

2. Site matching Highest Medium Medium/Low Lowest

3. Ability to follow trajectory Yes Yes Yes No

4. Maximum temporal scale Lowest Lowest Highest Highest

5. Maximum spatial scale Lowest Low Highest Highest

6. Scope (range of perturbations) Lowest Medium/Low Medium/High Highest

7. Realism None High Highest Highest

8. Generality None Low High High

9. Exploratory/discovery Low Medium High High ! One point is worth emphasizing: Diamond’s analysis is about community ecology.

This is not a static description of the tradeoffs between experiments. It is about the tradeoffs between experiments in the context of a particular science that has to deal with a specific set of its own questions and its own natural systems.

!318 To condense Diamond’s arguments, laboratory experiments provide great advantages in terms of (1) the regulation of independent variables and (2) site-matching. These virtues, following common practice in experimental design, pertain to internal validity, which we considered in some detail in the last chapter, as well as in the above. Internal validity is thus about determining a cause (or more generally, a “result”) in a particular experimental context.

An experiment will fail to be internally valid if, for example, the perceived correlation between X and Y is due merely to chance, or, if other uncontrolled or unknown variables are correlated with X and are the actual causes of Y. Problems such as these can be avoided through the active regulation of independent variables.

Natural experiments, on the other hand, provide advantages in terms of (4) temporal scale, (5) spatial scale, (6) scope, (7) realism, (8) generality, but are less rigorous in terms of

(1) and (2). Field experiments are useful because they are intermediate: they offer the ability to control independent variables somewhat, but possess realism and may have greater generality than laboratory experiments. Diamond concluded his analysis by claiming that the experimenter must accept that performing a particular type of experiment involves a trade-off in virtues along each axis; although it would be desirable to maximize all virtues in one experiment, this will not be possible. The question “which is the best type of experiment?” is thus a silly question. “NEs tend to have advantages over LEs for studying species interactions among whales; the advantage is reversed for yeasts.”667

In a recent article, Diamond adds another interesting axis to those already given (axis

9 above).668 This is presented as being about an “exploratory” or “discovery” advantage that

667 Diamond 1986a, 19. 668 See Diamond 2001.

!319 natural experiments have over other types. A more interesting way to characterize this is in terms of “surprises.” Since we have not set-up the experiment actively in a natural experiments, we may find among the results we were looking for (or outside of them) surprises we did not expect. Diamond writes, ! Professors of field biology urge their students to avoid “bird watching” not driven by hypotheses formulated in advance, and to go into the field only after designing a well-controlled experiment to discriminate between competing hypotheses. All too often, the sad result is that the student succeeds in answering that original question, and thereby fails to notice some much more interesting question at the same field site. Unplanned natural experiments create ecological communities that we would never have dreamed of creating, or that laws, moral scruples, or practical obstacles would have prevented us from creating even if we had dreamed of them.669 ! The epistemic role that surprises play in scientific practice, and especially in discovery, would be a fascinating study, which I have no space to pursue here. Diamond is arguing that natural experiments also provide more surprises than other types of experiments, and that this is epistemologically advantageous.

669 Diamond 2001, 1848.

!320