Changing Ideas About Cells As Complex Systems

NING 5CIENCE: VISIONS OF CEtt BIOTOGY RINE BIOLOGICAL LABORATORY REFLECTIONS INSPIRED BY COWDRY,S td by Jane Maienscheín GENERAL CYTOLOGY r for the new University of Chicago press Edítedby )very at the Marine Biological Laboratory. ngoing role the Marine Biological Labora_ KARL S. MATLIN, JANE MAIENSCFIEIN, emination of science, in its broader histor_ ractice and future potential. Each volume O.Nd M.ANFRED D. LAUBICHLER udes work about the MBL and its science :ntists; work that begins with workshops, :ollaborations made possible by the MBL; for, inspired by, and otherwise related to liscovery by the community of MBL scien_ ronographic, while others will be collabo_

: new ideas and approaches that find their :st did summer research, with a small NSF 6; it led to my first edited volume inspired oeople have been similarly inspired, and :ogether ourworks into a collection of re_ noting discovery through its exceptional aside. AIENSCHEIN íes Editor r of the Centerfor Biotogy and Society, itate Uniuersity

9 i o I o g i c a I L a b o r a. t o ry

THE UNIVERSITY OF CHICAGO PRESS

Chicago and London :rl<: Oxfortl Urrivcrsity prcss. CHAPTER 2 produced: 2oo7. Herediq) At the Crossroctds of 87o. Carnbriclge, MA: Ml.I prcss. CHANGING IDEAS ABOUT rbilclung in der Iliolog.ie. Jena: custâv Fischer. ir-r Heredity." CELTS AS COMPTEX SYSTEMS aiolog.ical Bulletirt 4(5)t 41-_So. e: ein Grundriss cler Lehre vom Leben, .lena: Gustav Jane Maienschein Tradition , und Evolutíon z.¡uischen lAlno- und finished the third ancl final eclition of 'nt and Inhcrítatrce. New yorl<: Macmillau. As Ednuncl Beecl-rer Wilson writing I H e r e dity. ecl. yorl<: prob- 3rcl New Macmillan. orig. his The Cell ín Development and Heredity, he notecl that in the future theritance in rB96; znd ecl., r9oo. ably no single author could write such a ceìl biology text. The subject hacl become too complex and required too many different kinds of expertise to clo it justice. About the same titne, Wilson joinecl Edrnund Cowdry ancl other leading biologists in a workshop at the Marine Biological Laboratory (MBL), where the clistinguishêcl group clivicled uP the topics to write tlÌe col- lective 1924 volume G eneral Cytolo gy (Wilson r9 z5; Cowclry 792 4)' Cowdry then convenecl a larger ancl even more diverse group to produce two vol- urnes on,specíøl cytolog (cowdry 1928; cowclry 1932). Far from being of merely antiquarian interest, these volumes reveal unclerlying assumptions that both reflected ancl informed the directions of scientific research. "lhe t9z4 Cowdry volume focused on the architecture and activities of individual cells, not primarily as building blocks of living organisms but also as the fundamental units that were themselves living, ancl the authors emphasizecl the value of studying these living cells in cletail. The group clearly saw the beginning of a new fielcl of cell biology emerging, one that might lose the coherence of a single approach but gain from different points of view, using clifferent techniques to ask different questions about complex cells and their activities. This chapter looks at the context in which Cowdry's volume appeared, a context constructed on the foundation of the first cell theory of 1839 ancl subsequent developments. The story leads to questions about what the Cowdry volume tells us about the science of understalÌding cells more generally. It looks at what the tgz4volutne offers in seeing cells as having gained autonomy, integrity, and biological itr-rportance as complex living systems in their own right. In addition, some of the chapters and reviews focus on understanding that the individual cells work together to make up complex organisms, such that organization arises through their connections. Yet the chapters mostly remain focused on the individual cells themselves rather than on how they communicate with each other and work as a whole. The Cowdry volume presents an American story, focused on Woods ein, and Manfred D. Laubichler 'r5 Hole at a time after worrd war I when scientists in the pired united states as- to scientific leadership. To gain perspective on the contributions of cowdry's vorume, it is usefur to start ori.ny at a prominent discussion the centennial at ofthe celr theory for a broader view rooking back, and then to move on to Generat Cytotogy itself.

Reflecting on the First Century of Cell Theory In his introduction to the centenniar volume entitled The ce, and protoplasm in rg40, editor Forest Ray Mourton noted that ciation the American Asso- for the Advancement of science was pubrishing the book of a series' It grew as part out of a symposium, herd in 7g3g, tocerebrate the centenniar of Matthiap schreiden and Theodor schwann,s introduction scientific cell of the theory. Because of the rich h. to that time, ,,in Mourton fert that a sensejii]J,;iï:ïi::::îï:llï:i he continued, "In another serse the cell Theory is arways new, for every discovery respecting this prirnary and essential unit of both pla't riving organisms, and a'irnar, h^as raised more questions than it has answered and has always widened the fields of inquiry,;fruroulton rg4o,,,Foreword,,). volume The set out to show both what was old and we, estabrished ancl was new a century what after the originat idea of celìs. By rg4o, discussion of cells usually separated plant, microbial, and animal cets' For plants, discussion typically invorved rooking dictable at such pre_ topics as celr walrs, whte discussion of animar celrs looked more closely at derineation of individual cers; contents of celrs, including cleus' cytoplasm, nu_ and organelres; und both internal and ternar to each cet' AIong "nuiron-ents, ex- the way came considerations orbioc¡emisüy cell physiorogy. and More.surprising in the rg40 volume are the less standard chapters on microbiology, viruses, ,n rri"'",hormones, and vitamins. The choice of topics and of contributors -uL", åtear.¡ust how much remained in ry40 to be discovered about ce's and especialry about teract with the ways they in- each other as we' as with their ånvironment. realized The contributors that they still knew relatively little about how the individual cells add up to an organized, whole, "o_pl"* organism, though they recognized that the process of making coordinited understanding "Jm¡inutions of parts was key to living systems and organisms. we see this emphasis on individual cets in textbooks of the For example, time as welr. Lester W. Sharp,s very widely used,Fundamentals of Cytutogy of rg43 (as welr as other editions) laid out the way that cleus cytoprasm and nu- work in the celr, Iooked at different t in¿, or cells, and recognized t6 Jane Maienschein : I when scientists in the United States as_ but had relatively little to say about the latter 'o cells make up organisms, gain perspective on the contributions that of Sharp noted that in addition to the cell theory focused on the cells r start briefly at a prominent ooint. discussion at other researchers had a different view of living organisms that ' for a b¡oader view looking th"-r"lu"r, back, and then on each organism as a whole. The two different :self. supported an emphasis perspectives have to come together in some way, Sharp recognized, for "in norrnul mass of protoplasm, whatever its growth pattern or degree First Century of Cell Theory "u"ry the many diversified activities are so coordinated that it nnial volume of differentiation, entitled The Ceil and proto- behaves as a consistent whole, or individual, from the beginning of devel- 'Moulton noted that the American Asso_ such harmony there obviously could be no organ- Science opment onward;without was publishing the book as part ism', (Sharp tg43, zo). Yes, but how could this harmony be achieved? r-sium, held in rg3gt to celebrate the cen_ The connections were not yet clear. Some biologists continued to look d Theodor Schwann,s introduction of the at organisms as organized living systems that happen to consist of cells, he rich history of thinking about , uf while others looked at aggregations of cells as making up organized living r a sense the Cell Theor ".ff At issue was partly a matter of emphasis, but also paftly a matter of the celr rheoryt, systems. causai efficacy. Do cell divisions and actions cause organisms, or does some ¡ and essential unit"ì;:';;"ïïJ;i of living orgurir_r, integrated wholeness cause cells to behave as they do? What drives the inte- more questions than it has answered and gration of the whole organism? These were still questions in the r94os, and rquiry" (Moulton r94o, ,,Foreword,,). The the discussion shows that biology had not embraced a single "cell theory" was old and well established and what to explain the basis for all living systems. I idea of cells. Sharp explained that cell structure and function affect or perhaps even ;ualþ separated plant, microbial, and effect the organism, but it remained unclear just how that happens. In an n typically involved looking at such pre- earlier picture of cells as structural units or building blocks, it was easier discussion of animal cells looked more to treat them like bricks or stones that combined into a larger organism I cells; contents of cells, including nu_ through forces outside the cells themselves. But if cells were each living nd environments, both internal un¿ units in their own right, then how do all those sepârate cells relate to the me considerations of "*_ biochemistry and organism as a whole, and how do they make up that whole? How could new the r94o volume are the less standard research resolve persistent debates? In particular, by the end of the nine- Inzymes, hormones, and vitamins. The teenth century, in the face of increasing knowledge about protoplasm and makes clearjust how much remained internal workings of cells, which theory about life should hold? ,,The pro- and especially about the ways they in_ ponents of the cell theory stressed the cell as the primary agent of organi- r their environment. The contributors zation, while adherents of the organismal theory insist upon the primacy þ little about how the individual cells of the whole, cells when present being important but subsidiary parts" lex organism, though they recognized (Sharp t943, zt). Furthermore, looking at evolutionary relationships by rted.combinations of parts was key to comparing studies of different organisms suggested that a different kind ganisms. of protoplasm might serve to connect cells and might thereby help to bring I cells in textbooks of the time as well. together coordinated whole organisms in somewhat different ways and videly used Fundamentals of Cytotogy not necessarily in exactly the same processes and patterns for each kind of I out the way that cytoplasm ,ru_ organism. Many questions remained in r94o, including questions about erent kinds of cells, "n¿ and recognized what cell theory was and how it had changed over time. lnschein Changing ldeas about Cells as Complex Systems .rt Agai', this work of trre micr-twentieth ce'tury rei'forcecr just how hacl been learnecl about trre cletairs 'nuch of ce's, ancì yet rrow rittle trrat knowledge revearecr about trre ways cers work tog'ether in a coorclinated way in more comprex animals. It is worth rooki'g at how trre science rrad gotten to this poi't' The history of cet biorogy ,^or", a first stage of trrinkiig. about, cells as the structural units of Iiving árga'isms, fbrtowecl by a stage of think_ ing about ce's as trremserves ,,agent[s] rnore n"arly the of orga'ization,,, as sharp put it' Reflecting on the deveroime't of ce' theory took srrarp, cowdry's group' ancl tar

Mosr orus have hedrcr.so-"iillr:;;;iïîI".,. story orschreiden and schwann and the cell theory that they inventecl, which is recountecl book in text_ after textbook-except that we clo not know the historicary accurate story, because the textbool

~~, e m s ? A . u o n ii ll ;; jffi ",,, Iï liJ, : ff : ïi#:ï,îil notyet been answered. Jr",ï;i*ï;;~~

Cells as Structural Units of Living Organisms For many who saw Schleiden and Sctrwan th o s e two gave c e, rh e o ry a n a m e. ;"; ;: :å:: are ; Ï:ä ïili ::î,liliL""î; constituents of living organisms and, second, ,lru, tn" help to expìain the indiviãuañ,t tiräry might organisms "t-"*i'*rn'u ""mplex that consist of ä,jÏi::iÏ;ii:î-:::t: '"lt*'lg *'"* ¿"ur ornããìtionar ob- i'both-,"-,.oo"lnJ,",Tili:::ili:::"JJ,ï;:îlJîî*å:ru*::Xl: some detail' In studyi'g cers, what one can see ancr it are of crucial how weìr one can see importance; and rnaking sure that others can thing is especiaty important. see the same Better Ienses reduced and better chromatic aberration, fixing, staining, and sectioni'rg L"thoa, the preparation brought consisrency to of specimens (Bracegirii. ,gzsl. richer The whole srory is much than rhis' ancr a numb". hi;;;;;ns of science have taken pects of earry cet theory- "f up as- In addition, as schickore of this Jutta explains in upt", volume, the technical innouation" "t a i'rerprerario,'. ¡, rh"-"mpr'asi s here,, *ii*¿iîJmlu:.jï: cells that informed the "" :l t9z4cowary voru'", to tribution herp us interpret the con- and impact of that volume. In 7834,I(arl Erns asesrases,.-n,"nr""tr",in::T,1ff each reriabry .i,:,lr*nïïï::;:ïi;ffi divicring i' the sarne urri , terns' -u, the same basic pat_ or what were rater calred lineagås of ce,",,owing divisiãns within ism' von Baer's images ,n.-urrnrr_ were arso taken as showing in the that the full materiar initial egg divides into more His.'rusrrati";"";;;;r"pportedacraimrr",ï:'#i::ri"îîrrilffurrA _or" makes up the developing organism-not iJ; intercerular connections cellular materiar (von or non- Baer rB34)' Trre i-;;;, were raken as representing division into separate structurar ,nitr, unifor those who herd that units were cells, they played those an important structural role. 20 Jane Maienschein lo, in fact, play this central role, thereby re_ cells as having such a role, especially those yet yet not everybody accepted Itheory. Moulton's question from ig4o focused on organisms as a whole. Historian Marsha Richmond points /es serve as the drivers for development who and Henry Huxley as a leading critic, and she argues that he re- rd the results of to Thomas the process of organismal pre- iected cell theory in part because it seemed to assign the cells a sort of way: To what extent, and in what ways, are role, as if the celts themselves cause development and body n simply the structural ?ormationist units of living sys_ Huxley held a more epigenetic view As Richmond notes, Hux- uction structure. of the cell theory, this question had ley insisted that cells are "not instruments, but indications-that they are no more the producers of the vital phenomena than the shells scattered in orderly lines along the sea-beach are the instruments by which the gravi- I Units of living Organisms force of the moon acts upon the oceans. Like these, the cells mark I Schwann þtive as the beginningforcell theory, only where the vital tides have been, and how they have acted" (Richmond re. They declared, first, that cells exist anï zooz, citing Huxley r8S3). Richmond further discusses the debates of the isms and, second, that the theory might time, which make clear that pell theory was not a clearly defined' unified, or lf more complex organisms that consist of universally accepted idea. (See also Whitman 1893 on what he called "the ving brought a great deal ofadditional ob_ inadequacies of the cell theory.") t. Those years also brought improvements In fact, one main alternative idea persisted, affirming that some sort of :opic techniques, as Hughes discussed in protoplasm lies outside the cells and connects them. Huxley put forth such rat one can see and how well one can see ideas. Adam Sedgwick was still invoking this idea through the end of the naking sure that others can see the same nineteenth century, as Baker discusses at greater length (Baker r988, r75; ter lenses reduced chromatic aberration, Sedgwick 1894). The idea of a reticulum, or syncytial connections, proved at- ctioning methods brought consistency to tractive because it seemed to offer an explanation for how the cellular parts rcegirdle r978). The whole story is much might work together as an organismal whole. Physical connections make Ihistorians ofscience have taken up as_ the parts into a network. The same reasoning held for the nervous system. n, asJutta Schickore explains in chapter 4 At the end of the nineteenth century, researchers argued aboutwhetherthe vations brought additional questions of nervous system is there from the very beginning in a sort of reticulum that e is on the emerging understanding of then grows largerwhile maintaining its structure (Billings r97t). This idea lry volume, to help us interpret the con_ of a protoplasmic reticulum could explain how the complex structure arose te, and was maintained. In contrast, the neuronal theory held that individual Ld presented observations offrog cleav- neuroblast cells then develop nerve fibers that grow out and make connec- hat later biologists saw as cells dividing, tions. Gradually, they extend, develop connections, and make up the ner- way and following the same basic pat_ vous system, :ages of cell divisions within the organ_ Some researchers, such as Ross Granville Harrison, could easily imagine :aken as showing that the full material how such a complex system can arise from the interactions of individual and more cells, which remain separate. cells. His study of individual cells led him to develop the first successful claim that the collection of cells is what tissue culture and the first stem cell research, with transplanted neuro- intercellular connections -not or non_ blasts (neural cells). Harrison's work was taken by many as having resolved 'he images were taken as representing the question in favor of the action of individual cells working together. rits, and for those who held that thosã Certainly Santiago Ramon y Cajal agreed. In contrast, Ramon y Cajal's co- ]rtant structural role. recipient of the Nobel Prize, Camillo Golgi, and others never gave up their ¡ienschein Changing ldeas about Cells as Complex Systems 21 convictiolls that the system was inextricably interco'nectecr fro'r trre be_ gi''i'g. The discussions were part of persistent clebates ar¡out whetr.rer cre_ velopment is more pref'o'natio'ist, that is raicl out frorn trre very begi''ing ir :r prefo'necl way, or epigenetic, that is arising o'ry gracluaily over ti're. 'I'l-rese clebates rrave bee' criscussccr i' cletail ersewrrere, ancl we neecl not repeat the entire story here (Maienschein t9B3; Maienschein r99r). It is worth remembering that wrren cliffere't people loolá at ce,s, some saw thern as newly arisen objects up an organism, ancr sone saw theln rnore as proclucts 'raki'g of celr crivision from past ceils. Both are partry true, and it realry depencls on how one cloes the rooking ancr wrrat one is looking for. It depencls, as is so often the case, on perspective. The fbrr-nôr, epigerretic view requires an expranation for how the ìndivid_ ual cells arise ancr how trrey make up a whore organism. wrrere does the organization and where croes the life come from if the separate and incrividuar cells corne togetrrer onry gradualry over time to make a whole? The tempta- tion was strong to invoke some fo.n of tereological or vitalistic principre or force to off'er sucrr an explanation ancr crrive the process. ¿ristotie had given us two and a half miilennia of trrinki'g in sucrr graduar, epigenetic terms, and his insti'cts fit with those of many other obseryers. while later thinkers discarded Aristotre's enterechy, they hacl to fincl expranatiors for the emer- ge'ce of form and function in sorne other way (see Maienscrrein zor r). In contrast' if cells arise o'ry fron"r ,,life,,ancl other cells, trren the the be- ginni'gs of the fbrm are in some sense arreacly there from the begi'ni'g. Thus, whe'Robert Remak srrowecl that cells divide a'cr give rise*to other cells, and Rudolf virchow farnousry declarecl that,'omnis cerura e cerurø,,, their assertions that celrs come fiom other celrs were crear ancl direct. The claim was neither entirely new, however, nor was it universa,y accepted at the tirne. Asserting that cells come from otrrer ceils pushes back the expra- of where trrey come 'ation from in the first place as wen as the question of how they come to be "arive." If ceils are tne fundamental unit åf life, ancl rnore tha' just the structural builcìing blocks of rivi'g things, then how so? And what follows for our unclerstancling of biological processes? (See Maienschein zor4, chap. r.)

rhis very quick rook ",,""ol"jlio1'":i:lfri:n the end of rhe nineteenth celìtury and to the work of Trreodor Boveri, oscar Hertwig, and Eclmuncr Beecher wilson. A number of researchers hacr demonstrated that trre celr has structure, with a clistinct bounded nucleus, liquid orgel-like cytoprasm, 22 Jane Maienschein tricably interconnectecl frorn the be- the mitochondria and Golgi boclies, with ancl other structures inclucling persistent debates about whether roles during cell cle- spi'clle fibers, asters, and centrosomes playing illlportallt Lat is laicl out from the very cletail, ancl the rich beginning clivision. Again, historians have coverecl this period in tt is arising only graclually over tirne. historical work of cell biologist Sir Henry Harris at the Ur-riversity of Oxforcl n detail elsewhere, ancl we need not provicles the best moclern account in The Birth of the Cell, which appearecl ein r983; Maienschein r99r). in t999. :n clifferent people lookecl at cells, Harris goes over much of the same grour.rd as Baker ancl Hughes, but s making up an organisrn, ancl some with co¡siderably n-rore interpretive subtlety. He has reread the origipal ision from past cells. Both are partly sollrces, ancl furthermore has the benefit of an additional half-century of e does the looking ancl wl-rat one is biological discovery ancl reflection on ollr unclerstalìcling of cells. Harris ìe case, on perspective. frames his work with a selected quotation from the French microscopist s an explanation for how the individ_ François-Vincent Raspail: "Give me an organic vesicle encìowed with life whole organism. Where does the or_ ancl I will give you back the whple of the orgatlizecl world." Making the e frorn if the separate and individual claim that the Gerinan story came to dotninate-ancl perhaps to clistort- tirnc to llral

Gelts as Complex Living Systems: Wi lsons The Cell Edmund Beecher wilson built on the r,iorì< of Herhvig, Boveri, and many others' For w'son, the celr prays a foundational rore for for rife ancl therefore arso the srudy of rife rhrough biorogy. wilrån', work influenced generarions biologists ;fi:ï:::"ïetopmental because of the way he r,rouirritog"trr", cerandirscrranges"i"T,ii"i'iï:î:ffi entitled ;:::;î:ff ,:ïî,,i"ïï:*; The cett in Deveropment and tnheritinceand decricated to Boveri, wil_ son opened his introduction bypointingto schleiden and schwann: ,,During the half-century trrat has erapsed ,in."ih" Schleiden of the cell_theory by ancl Schwann, in r83B_39, "rrunciation it has become ever more ent that rhe clearly appar_ key ro aìl ultimate biotog.icat probtems be ;r;;,;; ål;;;l,rutyrir, sought in the celr' " Furthe.more, ;'No other biological generarization, only the theory of organic save evolution, has brought so phenomena many apparentry diverse under a common point ofvi".," o, ,-ru, n".omplisrred more fbr unification of knowledge. the The cer-theo.y-rr, tt evolutio'-theoryas be praced besicle the one of the foundation "r"fore stones of lnoclern biorogy,,(wilson 1896' r)' He saw his task in part as bringing the two together, showing the role of cells in deveropmenr herediry, t; Tg -;r tnut -ud" evorurion possibre. By the third and final edition in rgz5,rilro' acknowledged that a great deal had changed-the volume nua grorvn from 37t to 7232pages had undergone reconceptualization and whire seeking approach' to retain its synthetic For that last edition, he opened witrr a srightly different tone: of modern ,:il:å:ji:il:"#:ï", scientinc prosress the ceu_theory of commandingruna-u,r:,:iil"ïï,:ï:"1i'.ï,ï:îî::;:i::;i:""_,î:

24 Jane Maienschein succeeded cf chromosomes after divisions, a "rude sketch" and that it nonetheless but they their ideas werejust that study of living organisms' and re- :less. Observing carefully both naturally ,,opening a new polnt of view for the in plan of organization that ived examples, Boveri added immeasur- ,n" of a fundamental common "udines (wilson t9z5' t)' ivision with his experimental work. "T"Ui, their endless external diversity" ;;;;it"t periods since the ¡avidson have suggested that Boveri was "";;;;; third edition, wilson poinred to three rough focused on the basic ideas terms about the cell and the roles of its of the initial idea of cells: the first ;;;;; and ce¡ divi- 'isionary able to imagine something con- and rheir rores; the second looked at development Tiä."rr, of heredity, which ex organisms guided by gene regulatory the rhird brought in the chromosome theory i* third period 'ught of them as determinants on chro- ""0 explanations of the causes of cell division. This äir"Jr""o Wilson rtyet have a concept ofgenes specifically more a matter of biochemistry and metabolism, t uã -"¿" hereclity l'tt4). Wilson pointed to several key phenomena tt,ougl't (Wilson tgz5, that ,,we are still without adequate un- ,t u, ä_ui". d.puzzles,concluding g relations between nucleus and cytoplasm Systems: Wilson's The Cell ä"ir,urrai"g of the physiological Le is concerned in the operations of work of Hertwig, Boveri, and many oth- ]nã or rr," Ãurrrr". in which t'he nucleus repair' and in the determination ndational role for life and therefore also lorrr,r,r.,iu" metabolism, of growth and de- ¿. may be said of our present knowledge of Wilson's work influenced generations ãf t traits. The same "r.aituty localization"' Further, s because ofthe way he brought together u.lop-"rlt, above all in respect to the problem of appearance of hereditary traits in regular nt pieces ofevidence about parts ofthe he asked, "What determines the first 1896 edition ofhis classic textbook, orderofspaceandtime?Howaretheoperationsofdevelopmentsocoor- system?" (Wilson tgz5, ttt5)' [nheritance and dedicated to Boveri, Wil- dinated as to give rise to a definitely ordered ,,During terms of the "or- tingto Schleiden and Schwann: And how can a proper understanding in physicochemical toward vitalism rce the enunciation of the cell-theory by ganization" of the organism push away any temptations , it has become ever more clearly appar- that he found in some of his contemporaries? ;;ical problems must, in the last analysis, Inthethirdeditionalso,evenmorethanintheprevioustwo'heended 'No other biological generalization, save bypointingtothesuccessesofrecentyearsinmovingforwardonallthree we has brought so many apparently diverse contributions, while acknowledgingthat many questions remained' "If a formidable ar- ¡fviewor has accomplished more forthe are confronted still," he wrote in the final paragraph, "with the certainty that eory must therefore be placed beside the ray of problems not yet solved, we may take courâge from been Ltion stones of modern biology,'(Wilson we shall solve a great number of them in the future' as so malry have nging the two together, showing the role in the past" (Wilson rgz5, tttS)- and in ways that made evolution possible. senior scholars today recall buying this last edition of wilson's book instructed to keep the 725, Wilson acknowledged that a great reading it for one or another class, as well as being or The ceII, d grown from 37t to rz3z pages and volume nearby for reference. with his series of three editions the r while seeking to retain its synthetic Wilson provided a compendium of existing knowledge about cells and each cell rpened with a slightly different rone: ways that they reflect the processes of life. His message was that of life scientific progress the cell-theory of is a fundamental living unit, useful for understanding the processes those processes' in r83B-39, stands forth as one ofthe as well as the structure of living organisms. Interpreting and evolution' :eenth century." yet he went on to note though, required bringing together heredity, development, ienschein Changing ldeas about Cells as Complex Systems 25 wilson provided a view of the ceil as an individual, complex, riving system' The year before his first edition in 1895, he had pubrished The Atrq.s of Fertílizatíon and Karyokinesís of the ouum,which included a set of beau- tiful print copies of photographs taken of the early stages of fertirization and celr division. coraborating with photographer Edward Leaming, wilson sought to show his readers the complex parts of the celr and how thcy cha.ge during those early stages. By rg96, he provided considerably more detail about later roles of the cell as we'. His poi't in the Atras was precisely to provide an atlas, a sort of collection of maps of structures. The Cell added function and development. In the process, the cell came alive' cells sti'went through stages oidevelopment, but wirson sought to capture more than the standardized stages characteristic of normar tabres. He wanted to understand more about what it meant to be arive and espe- cialþwhat it meant to be organized into an individual organism with integrity aird autonomy. what wilson did not quite see yet, despite his crarity of vision and depth of understanding, *ur,h" importance of understanding the ways in which cells interact with other celrs and the comprexity of the regulatory processes that reside within the inherited materiar but go be_ yond each individual cell itself.

Edmund Cowdry, General Cytology As wilson had acknowledged, and despite his attemlts to provide a summary update of the field with his own third edition, by tgz4 thecharenges of understanding the cet had already grown beyond what any singre searcher re- could grasp. Indeed, the very brief summary of ideas taken to be important leading up to Tgz4has focused especially on parts of the story about study of celrs' other researchers were rooking more intentry at physiology, biochemistry, and other areas that fed into the study of cells, es_ pecially in later periods. The point here has been to put us at reast partly inside the thinking of those who gathered to produce the volume edited Cowdry. by By 7924, the biorogists who gathered to produce the edited vorume agreed that it was time for a cooperative approach, which Edmund Cowdry coordinated at the Marine Biological t uboratory in woods Hole (see arso the introduction to this volume). cowdry noted that, because contributors had worked in the MBL facirities, "the vãlume, as it stands, is to be considered, to some extent at reast, as a contribution from the Marine Biorogical Laboratory" (Cowdry t924, v).

26 Jane Maienschein he cell as an individual of Chicago, which published the volume, has a folder of :dirion in rBe5, n" il'oiil,orfJå';;ir^::,:: The University to book. One s of the ovum, and letters related the is labeled "t7. A Textbook of which included a set of ¡"uu_ reviews rhs taken cytology. By Frank R. Lillie, et al." and summarizes the proposal of the early stages of fertilization General ng which Lillie apparently presented to the press. Claiming that with photographer Edward f,"u_i.rg, for thebook, cers total would not exceed 65o pages (the actual was 7s4pages), the press the complex parts of the cell the and hoî of The press By calculated a net investment $3436.75' estimated sales of one .tju8"r: 1896, he provided considerabþ 'the cell copies at $5 each' Those numbers probably look astonishingly as well. His point in the lttas w; thousand crt to today's publishers, who would nonetheless be reassured to note that of collection of maps of structures. low Jevelopment. the costs included $toto of "overhead." In the process, the cell came tges of development, but Wilson sought to THE ized stages CONTRIBUTORS characteristic of normal ,ãU1".. about The contributors to'Cowdry's volume all had close ties with the MBL, which what it meant to be alive ana place ed into "rp"_ was a prime gathering for biological research by the rgzos (figs. z.r an individual organism with in;g_ did not and z:z). They each had a home institution, but they came together at the quite see yet, despite his clarity Jf MBL in the summers to discuss their shared interest in cell biology. It is ng, was the importance of understanaing worth getting a sense ofthe people involved, and a short biographical sketch vith other cells and the complexity of the gives a sense ofthe group. within the ofeach Yet they each had independent research inherited material but go be_ careers, and so their biographies remain separate and largely not overlap- ping beyond their collaborations at the MBL. lry, General Cytotogy EdmundVincent Cowdry d despite his attempts to provide a sum_ ,wn_third Edmund Vincent Cowdry was born in Alberta, British Columbia, in tggg, edition, by tgz4the challenges 'eady grown the same year that the MBL opened its doors. Cowdry received his Bach- beyond whar any singleîe_ elor's degree from the University of Toronto a phD very brief summary and from the University of ideas taken to be of Chicago. He moved to the Johns Hopkins University in anatomy and in focused especially on parts ofthe story r9r6 married Alice Hanford Smith, going to the MBL for summer research. rers were looking more intently at physi- A year later, eas that the China Medical Board of the Rockefeller Foundation re- fed into the study of cells, es_ cruited Cowdry to establish and lead an Anatomy peking L here Department of has been to put us at least partly thered Union Medical College in Beijing, and he and Alice moved there in r9t7. ro produce the volume eaiiea bi With the birth of their first ctrild in t9zo, Cowdry returned to the United thered states to the Rockefeller Institute in Newyork and studied a range of topics, to produce the edited volume including anatomy, cytology, parasitic diseases, and aging. He continued ltive approach, which Edmund Cowdry to spend al Laboratory many summers at the MBL and to establish his editorial creden- in Woods Hole (see also ,wdry tials. cowdry took up an academic position at washington university in noted that, because contributors st. Louis in r93o. There, he moved increasingly to studies of aging while con- re volume, as it stands, is to be consid_ tinuing ntribution to focus on cytology, looking at cell degeneration in particular. As from the Marine Biological HyungWook Park has shown, Cowdry became a leader in gerontology and organized a conference on agingat the MBLin tg3T,whichwas supported by ienschein Changing ldeas about Cells as Complex Systems 27 GENERAI, CYTOLOGY A ]-EX-t Booli o¡' cIi¡.t,tiÌ-rilì s.ntt;c].uRl T.,\ùl_ti oll coNltrÌN1.s 1\NI) NL]NC'I'ION I,()Iì S1'UDIÌN.I'S OF Iì¡ol.()ct AND i\t¡:t)t(.tNt,. tU,,¿ r. v*n,, ciS*nln Uut""".ly, ^-r, rt.r Cir. tl :oùr U, .fp1r j$r.rs 4¡ Ùr¡.ûr¡vr-rR! (ì:nc Àtt ¿t ^p t'. .\tail:a,,. u"r, , ,ry .r , i". i,,,,.i¡ ß¡ ì:ij,i ì, ,,,. nu. trl I'r!\ir¡r.r¡r\ ¡+ rrrr CfÚ rñ t)¡Ìr¡!,\,,S1!s¡^\.',,, trdt. il. r.v.Lt. u¡.:v,F;,r (r r, ,...i""-,,",, i.i,i..ì,r,r.,.,, iii¡I,",'"r*" ;iär¡,lifir' $1hniJ i...,. IV. RF^crnny or rn¡j C¡r¿r lllì,i..i l; ,1'.: ìi;;i;ìì: i,^ìl,ill:* tdrr, s. 1_ilû, Nct¡ Rcs.:"1 i.**r"ìy, ù^ñ,r(¿r tr rr:wrJ rrrorr^* r ,rti,ÀN ir."iu, olio ¡1t)\f¡tNÞ u s..Þ, ¡. \vt¡ (ôv tr,rr tr I,no¡o!¿¡cx,ls Drr¡:Fxr,-¡:D Drssr.rros]l:1..'.::'..^, Á\Þ Iñrù,ô\ ¡y )fr R,. f ¡, / (r,,,i¡¿r. I oijr.jt Unn, r.irr L.i . rt (i.Iqc, N." v".i Cr," VL llrMs +¡"r¡, (iorLr.hÌ.rtsrn.,.\\D (irp^!rur1r. SurçÌrsili tnilru, t t.. ( r.^Lr..Iht IDMUND V. COTII)IìY Ror\rftjLr Iì{ i'¡lc lor Rtrer¡(l" \{\ Yo¡L ciry '\tcJtnl Vll. n¡.IrH^À,,r c'rs rr- t]ss|r (,u,.ruR¡i rvÒail n. t.¿ß n\¿ .v t¡,.ta r i,..,,. C,,,.*¡" rqJ \1'rdr¡{tuu. U,F¡rjn,nr"A tu,,j,u,iou ,i r,".f,r.,,ts. |-rrirror., ¡t¡Dl.,r VIIL ITTÂrr¡¡z{ros t ¡ril({ ¿,r;., U,rivr:ft;ottll.ro. r1,..,ç,., r,, r.. ¿. /6r, ìfo\nrJ , "ffib Univil.ilr, tvi.hi_Ârd¡, D.c. '.,, lX. C!ûùüR Drrr¿Ârivnrro"- u¿.iil c. rrt Cot*tr,,ptitccto u"i,",,lr, l.;,"..", i"*,i^", X. T¡rri W Õ'ko!cso!L trroRy Dr Irrr1,,¡v u"r'"i','¡ i'"''"n,"¡, @ "íi::,¡,""'" ' 'r.i'.,u.i,*,", Xl. Mr\DxüÀr Itr.Frr{rr ru,,,,n.u.,"",,i"),^;ì,,";l;;j;ìi:lrùr;,,,r"- nrr lrrui ro (\: r,, , . , . . iJ7

~~THE UNIVERSITY OF CHICAGO PRESS CHICAGO. ILLINOTS d~~

~~Figure z.r. The title page and table ofcontents f cowd rv's G e n e ra t cvto t o gv' a s p u br ished,^, Jli~~

~~theJosiah MacyJr. Foundation ancr is considerecr the first sucrl scientific meeting (park zoo8; zor6).~~

Ecrmund Beecher *,r,",tÍ,i{i{!r'::f;{!;ïriorogist or the cray ancr senior statesman. Bol a rearni'guboutnutuiïi,T"'"lii:Tillî;.iîliil,åJ.'::î:,iî;?ff versity I in ß75 and proceeded to the newJohns Hopkins university f.or PhD under wilriam I(eith a Brooks. After a visit to ce'nany Zoological urra tn" Nupt", station, wirson spent a year at w'riams corege, then visited at MIT' where he wrote a biology rexrbook with his fe,ow ate wi,iam ñoor.inl gradu- secrgwicr<, and in rg'5 he took a position as head of the bior_ ogy department at the movedtocorumbia""i"""i,Hif"î;:Hi,{::î:ä."X.J ji;iii::; z8 Jane Maienschein IY TABLE OF CONIENIS I. INnoDUmroN t\ unyÅ a.wilþs, cot\ñ¡¡" i¡e"";ty, t"* V.* b¡ly r II. So!¿ CENE¡^L o! mE ftrErírsny o! CILLq Attdt p. ' UothúB,UnlvcÁty^sprs of Cin(jnuri, CiÌcinn¡ì¡, Ohto 'r IL P¡¡¡rE^lrrrrr or rrm CrrL ro Dffusr.'o SuDs¡i.'oa ìIdt¿ il. tüoät, UD¡vcßily or peûyld, rlit"àapt,r, i,.r.. e7 sylvmt¡ lV. Raøvn o¡ m C¡f,,, . ,6s xdrp¡ S. ¿ili¿, Neb RR;.¡ LUo.ory, Cl"*Lo¿, Oh; V. 1\G Prß¡w Snuøur; or p¡mplÁs Æ DEBI¡Nþ sr lf¡oo. Dt6lb¡oñ INJEø¡ON . ^ñD Robat chñb.,!, com.rrJni*"¡ty uci¡-¡ ðol"g", ñ"" ïo.¡ cu, 'r, Vl. Àf¡eroNDnu, cokr ÁÞÀutus, GEolrDrÀL Sos N.. y. ^m Añnd C@dry,hc Rdcfê¡¡cr l6rituh for Àtcdj*i Ã***, "'. Ns York City UI. ß&vror o! CELß n t$@ CuLil¡E tl'ô,ilil râs ,t .H. ail N;;;; ¡.-¡*¡, ä.s" nui u,å" oi $,ùúinsþn, Dryrhom of [nbr¡olory, ¡¡rìr"q U"Virm' VIIL FErrrdarrox 4e F,alÊ R. Litu,uúvcßiy u óliq", òlr.g., irm"i, ¿ ¿. Jßr, Homd Utr¡vcB¡ry, W4biryron, D.C.

UC CE[@r D¡n¡rEW l¡oñ , pr¡iccton 5J7 E¿o¡ñ C. CùLtíì, Unn"rity, ¡rinem, lt* J:*; X. hE Csror@lD Iiro¡y o! I{s¡Ð¡¡y tu aûaec z. u"ciuas, t ;;u¡s-; it"ryl*"¡, ltir,¿jp¡¡rl Pcrylv¡nii )aL Mm¡u^N IIEnÐrrr rx REkitoN rc Cìftlæy thût A. L|øroi, Cotwbi¡ Udvcrily, Ncw yo¡k CO; -' ItDÈ,....,ii|

~~rf contents from Cowdry,s 1eneral Cytology, rlished in 1924.~~

~~rd is considered the first such scientific~~

~~BeecherWilson~~

leading cell biologist of the day and a Illinois, in r856, young Wilson enjoyed phB ) received a degree from yale Uni_ he newJohns Hopkins University for a After a visit to Germany and the Naples Figure z.z. The contributors to ceneral Cytology, as pictured in the tgzos. Edmund cowdry, bers, I year ât Williams College, then visited the editor, is in the ce nter. Others include (clockwise from the top leftl Robert Cha m Margaret Lewis, Warren Lewis, Frank Lillie, Ralph rxtbook with his fellow Hopkins gradu_ Edwin Conklin, Merle Jacobs, Ernest Just, Lillie, Clarence McClung, Albert Matthews, Thomas Hunt Morgan, and E. B. Wilson. lmages he took a position as head of the biol_ reproduced from the website "Historyof the Marine Biological Laboratory" rwr College for Women. In rggr Wilson (http://h ps repository.as u.ed u/). re he remained for the rest of his career, aiensche¡n spending almost a, his surnmers at the MBL. His The and Atras of Fert,ízation Karyokinesz's, ancl The ce, setthe sta'crard, as cliscussecl earlier (w'son rB95; r896; Morgan r94o).

Arbert p. Marhews was orrrrlt!{iäKtl:*:"He receivecr gree from a Bacheror,s crel MIT, stuclying biology uncler william sedgwick, and then phD from columbia university. a Heiaught first at rufts and college Meclical school, then in rgor movecl to the university of chicago, where he remainecl for fifteen years ancr was promoted ,o oroLrro, nnd eventuarìy chairman the p'ysiorogy Department' of In 1916, he rnouecr to the university nati as carnegie professor of cincin- of Biochemistry, and chemistry servecr as heacl of the Bio_ Department unt' he retired i,-, ,g3g. Thougrr his work fbcused on chemistry, he also pursuecr physics in relation to rife on gravity, matter, ancr wrote worr< space_time, ancl such tc rooking at the biochernistry orsecretio",, ;ï::#Hiä i;iiliî;ilîï r a i n e d r h a r,, jÏ wo o d s H o I e wa j I ;i "xp s u s, .r., ir u ror. u Jj,î:ï:;:" interests, ancl " " " the group benefiteâ immensely his u,r¿ ,ti_ullad frorn 'ew onBuzzardsu"ro*llirr;t":îilff :ilJå".,äîffi across the srreet, conklin *î:î*:,,m; a'cl Jacques Loeb nearby,;_";; Harvey reca'ed, ,;;;; orhers. "I can_stit see himiarking.briskly*i,r, g*ír the streets of r,.io"s arong WoocÌs Hole, with his heacl i ing rook in rris brue "r",, o, irhe were universe. His convict. "r"":ilTiiil1iï:åi,",""ffi: rr:;u*tîr;:äi::iii;:iiiiiiîtfi îJ1åf T;:ï,î"î;îîi*

Merker H. racobs *u, oïl f::Å:r::,#lli,,o,.r, pennsyrvania. ceived He re_ his Bacheror's ancr phD .reg.ee, from the university of pennsyrvania, and after ayear in Berlin, he retuinecl to tt urriu".sity i'protozoology. spent the waryears in the sanitation " He corps ancl then retur'ed to penn rgzr' remaining trrere in unt' he retirecr i' rg55. A memoriaÌ Love reported that ,,he by warner E.

p ri n c i pi e s, d r. p ; ;iiîi : iï:, " " " l'åi'i,lji :ï ïjiï: ïîå., arouncl him, he was ab, ï å,ji pubrishecrhi,""r,b";i;:'iå,H,"llJJJlíl;lj]å1*iåîL"":i.r# the MBL' He had been a member of the MBL corporation since rgrr a'd 3o Jane Maienschein the MBL. }lis The Atlas of Fertílizøtion r in tgz5-tgz6 and then director from 7926-7938, laêfâme âssociate directo standard, was a lively time for as discussed earlier (Wilson :,:,;,;; directing the physiology course. The rgzos with significant growth of the phvsical facilitl¡s.'1rl^t:::"tt"t ä;;;, out research' In contrast' the Ïr.*;; attencting the courses and carrying Mathews r93os brought serious challenges thatJacobs had to navi- Olnr"rrt". "fthe to con- Chicago. He received a Bachelor's de_ íï",.keep the institution afloat. That he nonetheless managed er William many to enter the field Sedgwick, and then a phD ä;. ;i, studies of cell permeability and to inspire first at Tufts College Medical School, the dedication of a special issue of the.lournal of Cellular -an,dcomparatívePltysiologytoJacobsinrg56(Harveyr956).,, from rsity of Chicago, where he remained ""i0"", professor and eventually chairman of re moved to the University of Cincin_ Ralph StaYner Lillie nistry, and sewed as head of the Bio- RalphStaynerLillie,wasborninTorontoandreceivedhisBAfromtheUni- d in t939. of Chicago. After Though his work focused u.,,'., o' in 1896 and his PhD from the University :s in relation to life and "o'onto Laboratory in Cleveland, the Carnegie wrote work ,t n, fr" *ott ed at the Nela Research ch topics. then at Clark University Mathews began his career lnrtitution of Washington at Johns Hopkins, and |ns, but soon moved also at the University of Nebraska' to studyof living u, prof"rro. of biology, with positions just "Woods Hole was the place for a Harvar¿,JorrnSHopkins'andtheUniversityofPennsylvania.Inrgz4,Lillie lhe group benefited as professor of general physiology immensely from moved back to the university of chicago group, activity of or- living near Mathews's house until he retired in rg5z. Lillie was ir-rtrigued by the dynamic mas Hunt Morgan next door, Wilson ganisms,asreflectedinhisGeneralBiologyandPhilosophyoforganism.Like s Loeb nearby, among many others. ii, broth", Frank, Ralph spent many summers at the MBL as a researcher lking brisklywith great strides along ancl as a trustee (Ralph S. Lillie Papers)' ead held high and a keen penetrat_ about to lay bare the secrets ofthe RobertChambers parents served as and his ideals high" (Harvey r958a, Robert chambers was born in Erzerum, Turkey, where his and received in his highly idiosyncratic essay in missionaries. He graduatecl from Robert's college in Istanbul Canada' Af- an MA degree from Queens University in l(ingston, ontario, ter earning his PhD in Munich' studying cell physiology and embryology' staff' Iacobs he spent time at the MBL, first as a researcher, then on the teaching L pennsylvania. first Harrisburg, He re- and then directing a summer cell biology laboratory at the MBL' His years' iom the University of pennsylvania, academic position was in cincinnati at the Medical college for three he retired r the university in protozoology. He then at cornell university Medical college in New York city until orps and then returned to penn in in t949. His student Irene P' Goldring noted that Chambers encouraged she in 1955. A memorial by Warner E. her to go to the MBL embryology course' which she did in 1948' and lpurpose, very hard working, high- later wrote, ,,My introduction to the life of that community enabled me to He spoke ill of no one. To those hear at first-hand, anecdotes and somewhat apocryphal tales of the legend ,ove 7977t r6). ShortlyafterCowdry that Robert Chambers had become." Chambers told Goldring that during cobs became the third director of his own introduction to embryology, he had said, "Dear God, I believe in the r MBL Corporation since lgrr and chromosomes, I believe in the spindle, I believe in the asters, now help me rchein Changing ldeas about Cells as Complex Systems 31 to fìncr out what they actu:rily are" (Gorcrrir lg rg7g, tzTt). cham,ers as an MBL seruecl t*lstee erncr regarclecl sumlncrs i'woocrs l{ore ¿rs trrc nrost inr_ portant tirne of his ve¿rr.

warre"Harmonr:î:'",i:'::i::;';"",:'{üì"o,co',,ecricur,u,,.r,oo,, lnovecl to crricago. He receivecl his BS fiom trre u'iversity ancr remainecl of Michigan as a.'assista't ayear. In 1896, he ellterecr trre Jorrns Hopr

M(trgqret Reed Lewis Margaret Reecr Lewis ceivedherBAo.s*;J;,::;;ili:':ïä:ï"îiJiî:;î;ï:iïi::;. lege, Columbia U'iversity, ancl abroacl aízutrich,paris, ancl she did not Berlin, though receive a graduate clegree. u"r rtuai", amphibia's incrudecr regeneration in ancr crayfish, a'.r she servecr as an assistant to Thomas Morgan at both Bryn Hunt Mawr ancr corumbia. srre taugrrt zoology as an assistant in at Bryn Mawr in 1go7-zancr at t'" N"- york Medicar women in physiorogy, coììege fbr and trren servecr as a recturer and later ut surru..r ;àrr"*., trained nurses at Hopkins. Johns In 1910, she marriecr warren Lewis and bega. a long a'cr fruitful "ur"., of colraboration in celr biorogy, 32 iane A4aienschein lìolclring t97g, tzTr). Chambers servecl efllbryology,anclrelateclstuclies'Thatworkbuiltonherlgo8visittoBerliu' lrìlners in Woocls guinea pig bone marrow into a solution of nutri- Hole as the l-nost im- Tí.rå tn. ,tansplantecl in an experiment later cited as the first successful ;;;;;culturing'tissue wellt on to clevelop highly suc- #; of cells. The Lewises Lewis 'rammalian 'armon ..''*'approaclrestoculturit-rgtissuesinclifl'erentgrowtlrmediaandpro. LBTo in Suffield, of the process (Landecker 2oo4)' In r9r5' Lewis Connecticut, ancl soon äT*O tt"O*ssive vicleos Institution of washington Depart- s BS from the University of Micl-rigar-r ;;;;. o ,"r"u"h"' at the carnegie year. In 1896, he entered the still new ment of BmbrYologY' ichool, where he became fascinatecl by stuclied experirnental enbryology with FrankRattraY Lillie grapher ,,A born in rBTo in Toronto, and was brother to Ralpll George Corner put it, sum- Frank Rattray Lilrie was Toronto' where he to open the eyes ofhis young associate ;;;.t lilliå. He receivecl his BA from the university of 'imental cytology" (Corner by ernbryology' This lecl him to the MBL where Charles 7967, 326). ,..utlr" intrigued lineage in the freshwater Reed, whom he married in r9ro, and ãii, wr.it-un recruited Lillie for stuclies of cell d Lillie to graduate study at Clark Univer- Warren Lewis workecl closely in their mussel U¡zio. Whitrnan enticed of Chicago' where Whitman moved : and cell culture studies designecl to ,-Ç ir', tS9, and then to the University )nts of zoology' Lillie received his PhD under different conditions. They in" take up the first directorship "egrto course instructor for the os of developing cells, ancl continued ã.g-r.. two years later' At the MBL, Lillie became Lillie held positions at the Uni- . In r9t7, Warren Lewis movecl to the .,inryofogy course when it began in rB93' 's Department of Embryology, which versityofMichigananclVassarCollege,thenreturnedtotheUniversityof and remained there through- fhe study of living cells eventually lecl Chicago as assistant professor of embryology Department of Zoology' and ells behave in different culture condi- out his career as professor, chairman of the until he retired' He was a taì- ;. Lewis led a quiet life of devotion to dean of the Division of Biological Sciences sustain both the University of ldom seen apart. Theirvacations were ented administrator' who helped shape ancl a textbook onThe Deuelopment t the Mt. Desert Islancl Biological Lab- Chicago and the MBL. His research included and studyof freemartins by observations and experiments on of theChíclcinrgoB, studyof marine invertebrates' regarded as his most importa¡t ¿). ir-, Ctti.ago. His stucly of fertilization was It is this work' contribution, despite disagreements about interpretation' ?eedLewis developedinProblemsofFertitizationintgtgandupdatedinrgz4withhis the Cowdry volume' That rnnillg, Pennsylvania, in rgBr. She re- stuclent Ernest Everett Just, that they present in hypothesis" that a sub- rllege, then studied at Bryn Mawr Col- work involved what Lillie referred to as "a working and sperm' Lillie con- I at Zurich, paris, and Berlin, though stance, "fertilizin," contributed to the joining of egg (Willier r957)' Her studies included regeneration in tinuecl to support his stuclents, especiallyJust vecl as an assistant to Thomas Hunt mbia. She taught as an assistant in Ernest Everettlust Carolina' and was lt the New york Medical College for Ernest EverettJustwas born in r8B3 in Charleston, South in Meriden' New ed as a lecturer at Barnarcl College, sent to a boarding school, the l(imball Union Academy t9o7 magna cum rkins. In r9to, she marriecl Warren Hampshire. He graduated from Dartmouth College in at Howard University reer of collaboration in cell biology, laud.e asa Rufus Choate scholar. He began teaching nschein Changing ldeas about Cells as Complex Systems 33 i. washington, DC, ancl soon became crrair of the zoorogy clepartment.Just began goir-rg to the MBL in r9o9, wrrere he workecl with Frank Lilrie, stucry- ing the process of fertirizatio' as the starting point for i'criviclual creverop_ ment' Just received his phD from trre university of chicago fbr this work. It is not surprising that Lillie woulcl invite Just to ioin hirn as coauthor on the chapter for trre volume. Biographer I(en.eth R. Manni'g rras writte'a definitive and provocative inter¡rretation ofJust,s scientific contributiors and his place in the history of biological sciences as well as in the culture and life of academic society more generally (Manning r9g3).

Edwin Grant Conklin Edwin Grant conkrin had a large presence i' celr biology ancl at the MBL, as seen in stories in an interview taped just days before his cleath (conklin rg5z)' conklin was born in rso3 in walcro, ohio. The famiry rived on a farm, and conklin worked wh'e attencring a country schoor with one room and one teacher. He studied naturar rrisàry at orrio wesleyan university and received his cregree in rBB5. while teaching at trre college for blacks, Rust university, -irionury frorn 1BB5 to rBBB, he met and married Adkinson' Belre They hacr three chirdren. conkrin phD receivecr his from Johns Hopkins in r89r. He studied embryology, cells, and relatecl topics, while also working to reconcire his Methodist cànvictio's with evorutànary bior_ ogy' while atJohns Hopkins, williarn I(eith Brooks sent conkrin ancr other students to the us Fish commission station in woocrs Hore. very quickly, conklin learned about the just MBL across the street, where he met whitman and began his own cell lineage stucries under whitman,s encourage- ment' His study of ascidian eggs became his dissertation work, and his completed work made vp z26pages, 9 plates, ancl ro5 colored figures in theJournal ofMorphology. conklin enjoyecl expraining how his dissertation very nearry bankrupted the journal. His work demonstrated how cers divide, step by step, ancr acquire differentiation in their different locations within the embryo. conklin buirds on thatwork in his essay in the cowdry volume' At first Brooks had been skeptical of conkrin,s proposar to study cell lineage, but in the encr Brooks saicr, "weil, we give stucrents degrees for countingwords in crassics, so I guess we can give you a degree for c"ounting cells" (Harvey r95B).

Clarence E. McClung clarence E' Mcclungwas born in ß70in clayton, carifornia. He receivecr a Bachelor's and phD degree from the university of I(ansas and then became 34 Jane Maienschein rme chair of the zoology department.Just of medical school there' In 7972'lne went to the nrofessor and dean the vhere he worked with Frank Lillie, he re- study- "un'i""rrity or pennsylvania as director of zoology and remained until the starting point for individual of the develop- ,, ,ôqo.n"also chaired the Division of Biology and Agriculture the University urrà of Chicago for this work. Council through World War I, serving from L972to .^927' Ñutionut n"tearch rld invite Just to join him as coauthor on lfterretiring,hespentoneyearattheUniversityoflllinoisasactingdirec- lpher I(enneth R. Manning has written a toroftheDepartmentofZoologyandthenbecametheactingheadofthe 'etation ofJust's scientific contributions O"purtrrl"ntofBiologyatSwarthmore'HisobituaryintheNeuYorkTimes logical sciences as well as in the culture that his one hundred or so students had honored his "profound generally (Manning r983). ffirr"a with biological re- irrnu"n". on individuals and organizations concerned led him to hy- ,"ur.h" (New YorkTímes tg46). McClung's study of heredity irant Conklin of X chromosomes determines pottresize that in grasshoppers, the number presence in cell biology and at the MBL, .h. ,"* of an indiyidual organism. Males lack a second x chromosome' taped just days before his dearh (Conk- and provided early which led to the idea of a sex-determining chromosome i3 in Waldo, Ohio. The family lived on a a definable set of hereditary evidence that a particular chromosome carries :tending a country school with one room of heredity stimu- units and theieby shapes inheritance. His discussions ral history at Ohio Wesleyan University lated others, such as Thomas Hunt Morgan' /hile teaching at the missionary college t885 to 1888, he met and married Belle Thomas Hunt Morgan r. Conklin received his phD from Johns ThomasHuntMorganwasborninrS66inLexington,Kentucþ'intoafam- ryology, cells, and related topics, while BS degree from the ily with deep roots in us history. Morgan received his rdist convictions with evolutionary following biol- University of Kentucl

Initially, Cowdry's volume was to include an essay by Jacques Loeb on E. McClung physical chemistry, with a special focus on proteins' Loeb was one of the in 7o Clayton, California. He received a luminaries of late nineteenth- and early twentieth-century biolory. He University of Kansas and then became asked challenging questions and posed provocative interpretations about ¡ienschein Changing ldeas about Cells as Complex Systems 35 the nature of lifr rrre,'ecrr¿r¡ri.,";;Xri,1ll;:'.',Jil",,;"Tiä:::ïï':ilî:ïffi contribtttors hacl surely :îï:ï,,î be'efitecl fronr Loeb,s presence years. at the MBL over Unf'ortunately, I_oeb the became ill ancl cliecl in ry24 (pauly ry57).

In his i'rrocru*io.r r" rr]. ing "rr;;tï;r:;H,rted ro rhree periocrs of srucry- ceils as rre ìracr witrr his "t"u'br;i.;;ur in sornewh¿rt clifferent terms. earlr, larselv r,ru.*rn,, ;J"",,äiil",:rÏl,the cerr rrreory, whire rrre seconcr

i a n'ìr e re c, i rv, :, :j: '| " ".li :ff :î':,îï study of trre clera'sofiii::"Ï the ;*:*** *:; ceris. This 0".ìão reçluirecr bri'gi'g morphology ancl physiology, rogerher celr biophysics ancl . and biochemistry, embryology, genetics, ancl all tog"th", _;"i"n;j r ar b i or ogv "'i,h i,'.." u, i'g c o op era rt :i;:ï :ri;" researchers (Cowclry " " lïllil;i *il,î :-924, ro).Wilson no arrive ar comprete u'irv in a work sentingwideìy 0."*Jå",i'r'1"î;iî.i',xfiîx:::':ï: cliverse fieìcls of research. such a group, bring to their however, ca¡r at ìeast task a broacrer ur-r.r n'u." .rr,r"u, tlran knowreclge of the subject any sing.ìe writer can at this clay hope to corrmancl,, (Cowclry tgz4, This cornme'tary by the preemin*, tt). .",i tiotogist set up the vorume way trrat alrowecr each in a autrror consi¿"raure incrividual Rather .orrtrol ou". t_ri, topi.. than provicling a review or chapter, it is useful the approach of some "u"h to reflect on of the chapters ancr the resulti'gwrrore. chapters seem Ma'y of the rather surprising i' mocrern ;;"i:::":::i: ï:t Ma t h ews e n t i tr e s h i s n rs t s e c t i * : ,,m i-ilii riHi the ch emist of en ",,, " " ", i åii,îï ï I lî; ry tality,, un¿ uitu ^f for"",, ilÏ;iJ-H:ru::'-*""";;;;;;,.":iïffi :Ï:älJlY,îå;Ï:: rivingiscrear".rr.;;'l:"nïirr*iî,îi,î:;:"Jtrj:*nff in his eighty-page chapter, i;î:lj he arso .ou"r, u lot of molecular of co'temporary cliscussion chemistry, even if his chemistry colleagues would recognized some of not have it. Mathews crid not have tÌr" when ,nort ior*"ra_rooiirrg he concluclecl his chapter bt "r", chemistry li*re was *nown abour rhe of genes and that existing "Jil;;;r u"rri,"ä*" therefore seemecr against the gene theory. to weigtr Marhews showecl ""r"in.i"i, that, while the bio_ I eave m u crr oo ;;; u rion, n it was e s s e n riar :iì:iî,i;,:ï ::,i]*, "; "s ror

¡6 Jane Maienschein ed a physicochemical interpretation of In contrast, Jacobs's discussion of cell permeability explicitly acknowl- 's work was always stimulating, and the how little was known and yet how itnportant it would be to know rom edged Loeb's presence at the MBL over the every- about the process of crossing cell membranes. Cells clo not coutain ,ill and died in ry24(pauly tg}fl. thing that they will ever need from the beginning, so they must have permeability. But how that permeability works and what controls the diffusion of VOLUME materials across the membrane remained unknown. ViÌson pointed to three periods of stucly- Ralph Lillie asked about how cells react to their environments. Under- ook, but in somewhat clifferent terms. stancling their reactions requires knowing about both the stimuli and the ;tructural, cell theory, while the seconcl responses. As Lillie's chapter explores possible chemical, mechanical, mbryology. The third involved Mende- electrical, and other possible factors influencing reaction, it becomes clear : analysis of cell phenomena and rnore that here, too, remained many open questions. Yet Lillie introduced the i period required bringing together cell idea that cells do react and are not cornpletely autonomous or insular. The rysics and biochemistry, embryology, ability of cells to reâct to stimuli from outside is what makes cell-cell inter- : leading to a new, ,,many-sided cellu- actions, as they were called later, actually work; it allows individual cells ration among approaches and among to work together as whole coordinated and organized orgauisms. Ralph ;on noted that ,,it is hardly possible to Lillie's chapter really just points to the interactions, which became much 'oduced by several collaborators repre- more important in later decades. rch. Such a group, however, can at least The first four chapters, incluclingWilson's introduction, all raised many rore critical knowledge of the subject more questions than allswers. Clearly, a cotnmuuity of researchers had hope to command" (Cowdry tgz4, tr), come to recognize that cells are themselves complex and are also parts of tt cell biologist set up the volume in a complex systems, requiring diverse kinds of methods and questions to in- rrable individual control over his topic. crease understanding. 'each chapter, it is useful to reflect on In his seveuty-page chapter, Chambers provided a much more definitive s and the resultingwhole. Many of the report on the results of microscopic techniques for establishing the physical odern terms. In discussing chemistry, structure of protoplasm. Chambers acknowledged that there was more to be remistry and psychism" and discusses learned about asters and other details ofthe cleavage process' but also that forces as well as such mystifying ideas researchers had already learned an astonishing amount about both the cy- invokes Sir Oliver Lodge's etherions, toplasm and nucleus. Whereas Mathews, especially, had veered toward the rroviding an explanation for how the theoretical and abstract, Chambers grounded his discussion in concrete ob- Despite the rather remarkable quirks servatious. The same is largelytrue also of Cowdry's own seventy-page chapter vers a lot of contemporary discussion on the cellular parts-mitochondria, Golgi apparatus, and chromidial sub- :hemistry colleagues would not have stance. Thus, we see a diversity of methods and approaches as well as topics. rt have the most forward-lookingview In their sixty-two-page chapter, Warren and Margaret Lewis introduced lting that little was known about the experimental approaches, looking at cells in tissue culture. As the acknowl- :nowledge therefore seemed to weigh edged authorities on this topic, their work focused on laying out the tech- retheless showed that, while the bio_ nique and observing how cells behave as a result of being moved to artificial rpen for question, it was essential for media. Theirs is the only chapter in the volume with photographic plates, which had become a standard way to demonstrate the results of tissue rnschein Changing ldeas about Cells as Complex Systems 37 culture experiments. They discussed workwith a number of different kinds of cells and concluded the chapter with a short section on cell cleath in culture' In brief, cells in culture eventually die, they reported, and they did not knowwhy. In fact, it took several more decades before researchers began to sort out factors leading to the death of cells under normal conditions as well as in the artificial conditions of tissue culture. Fertilization was a more familiar topic by the 192os, and it is worth noting that cowdry included Frank Lillie and Just rather than Loeb on the eighty-five-page chapter on the topic. Liltie and Just, on the one hand, and Loeb, on the other, had rather different interpretations ofwhat happens at fertilization and of the extent to which the process is strictly mechanical and chemical-physical (as Loelr said) or involves a special substance called fertilizin (as t illie maintained) (pauly r9B7; Manning r9B3). while Loeb had worked for a number of years at the MBL, Lillie served as the second and long-term director of the MsL. It is therefore not surprising that cowdry included Lillie's and Just's interpretation ancl did not discuss the controversies. Perhaps because the debates had continued for a number of years already, this chapter comes across as more specific and established than some of the others. The same is true of conklin's forty-eight-page chapter on cellular dif- fere.tiation. conklin was a leader in examining the cell lineage in several invertebrate organisms and then pursuing the causes and patterns of differentiation in each case. Differentiation takes cells from a more general to a more specialized state and constitutes development, conklin explained. The process of differentiation occurs because of changes in both the nucleus and the cytoplasm. For conkrin in tgz4, it was not the genes that drive development, however. He held a common view of those focused on cells and embryology that "the genes or Mendelian factors are uncloubt- edly located in the chromosomes, ancl they are sometimes regarded as the only differential factors of developrnent, but if this were true these gelìes would of necessity have to undergo differential division and distribution to the cleavage cells, as weismann maintained. since this is not true, it must be that some of the differential factors of development lie outside of the nucleus, and if they are inherited, as most of these early differentiations are, they must lie in the cytoplasm" (Cowdr! t924,6or). That sounds mis_ guided, or at least over-simplified, to us today, but it made sense at the time in the face ofexisting evidence. Two chapters followed conklin's and focused on the nucleus and its contents. Mcclung's seventy-nine pages on the chromosome theoryof heredity

¡8 Jane Maienschein I work with a number of different kinds pages on Mendelian heredity focused on the con- and Morgan's forty-two of liv- ¡ith a short section on cell death in cul- ,"*, nucleus in germ cells. Look inside the cell fbr the driver ally die, they reported, and they did not "t*" approaches said' Chromosomes and chromatin carry ag Oro."rt"s, these ore decades before researchers that chromosome theory as began t.iåat,y-romehow. McClung concluded "the th plants and of cells under normal conditions as iirruni, is logical, consistent, ancl generally applicable to both tissue culture. ityet stands as one of the highest achieve- animals. admittedly incomplete, topic by the 7g2os, and it is the most promisingguide to further advances" worth not- ments in biology and offers llie and Just rather than Loeb on the (Cowdrytgz4,68z).Morganemphasizedthateventhoughthenucleusand :. Lillie and on the to driving what happens in the cell, the Just, one hand, ancl hereditary genes might contribute rent interpretations need, Morgan suggested, tnore in- of what happens cytoplasm remains essential as well. We vhich place in the chro- the process is strictly mechani- fårmation about the "physiological processes that take the book. said) or involves a special substance mosomes and in the cytoplasm" (Cowdry 7924,728). That ended J) (Pauly r987;Manning 1983). White ThroughOut, the.authors notecl the need for more information, more complex rrs at the MBL, Lillie served as the sec- understanding, and more sucóess in puttingtogether a picture of the are in BL. It is therefore not surprising that organism and its interacting parts. Cells are organic units, and they nterpretation living organ- and did not discuss the a real sense alive. But they are not the sole factor in making up ebates had continued for a number of isms and are also responsive to enviroumental conditions and to changing ross as more specific and established internal conditions. Most of the authors went on to further studies that expanded on their summary review approaches here, and some of the ideas ty-eight-page chapter on cellular dif- here were left behind with time. Yet the overall picture is one of growing examining the cell lineage in several understanding of the neecl for multiple approaches, perspectives, and :suing the causes and patterns of dif- interpretations of cell structure, function, and interactions. Many questions :ion takes cells from a more general to remained, with many opportunities for further study, some of which are tes development, Conklin explained. picked up in other chapters in this volume. s because of changes in both the nu- i,n in tgz4, it was not the genes that REACTIONS TO GENER.AL CY'TOLOGY a common view of those focused on Overall, reviewers responded enthusiastically to the edited volume. They s or Mendelian factors are undoubt- recognized the challenges of having thirteen authors with relatively short i they are sometimes regardecl as the contributions on each topic (though they seem relatively long as chapters). ent, but if this were true these genes And, as always with edited volumes, they liked some sections l¡etter than [ferential division and distribution to others. All acknowledged that the authors were all leaders in their respec- tained. Since this is not true, it must tive fields. And several noted that the volume could not have been written by rs of development lie outside of the any one person alone. It took a group to provide what they all acknowledged most of these early differentiations as an authoritative, comprehensive, and overall very impressive laying out )owdry tgz4,6or). That sounds mis- of the contemporary study of cells. s today, but it made sense at the time A review in ttfle Nationbegan by noting that "the summer capital of biology in America is at Woods Hole, Massachusetts. A couple of years ago some d focused on the nucleus and its con- dozen ofthe leaders ofthis scientific convent decided that there should be n the chromosome theoryof heredity a new book about cells" (Thone 1925). Another review by Raoul M. May in nschein Changing ldeas about Cells as Complex Systems 39 the history of science journal,1s.1,s, reviewecl cowchy's volume ancl wilson,s ,,too thircl eclition together. He concluclecl that rnuch goocl carìnot be saicl of these two great contributions to science. while, lÌowever, wilson's book is a rnilestone, the combined studies of the Arnerican i¡vestigators which togetlrer to.'.''t Gener(tl cjtotogy are a stepping-stone. wilson has mainly elaboratecl, as in the previous eclitions of his book, on c¡uestions concenì- ing cellular morphology, while Generar cytorogy incrvcres a great creal wrrich is physiological in nature. The two books together are a sple'clicl mise au poínt of all that is known concerning that most funclarnental of all living structures-the cell." wilson's was a classic volurne, rnarking the state of a fielcl, while Cowdry's also pointecl to new icle¿rs ancl clirections for future stu dy (May z t9 5, z t 4). Another review by Arvilra Meek Taylor in the c hic ag o Evening Post'Literaty Revielu surnmarized the book ancl concluclecl with a call for ,,for more such collaborative projects, nothing will clo more to acl- . vance the cause of science as a whole than such efforts as this" (university of Chicago Archivcs). other reviews offered similar views, though a number of them did find parts of Mathews's chapter on chernistry clecidedly ocld. cytologistJ. Brontë Gatenby provided a long and detailecl review, i'which he pointed out wrrat he found missing or misleacling in places, though he applaucrecl the volume as awhole. He referred to Mathews's cliscussion of chemistryancl psychism, in particular with regard to understanding how life emerges. It is worth reflecting on this point more closely, because cowclry, as editor, had allowed this part to rernain. The persistence of the views Mathews espoused shows that in 19z4 biologists were notyet clear on how celrs gai'rife, nor how they make up living organisms. Mathews wrote, ,,It is in fact the lu- miniferous ether which has macle things alive, for ether is the storehouse of energy; it is itself nothing else tha' spâce ancl tirne; energy and time,, (Cowdry t924, l^B5). Today, Gatenby's response seems reserved in its cri_ tique: "It is impossible for a working cytologist aclecluately to comment on such passages. They lnay mean something to the metaphysician, but one prof. cannot help feeling that Mathews'views on the relationship between cell lipins and cell proteins, or on the biochemistry of clevelopment, would have been more useful" (Gatenby t gz5, :.g6). A few critics we't further, suggesting that Mathews's chapter ought not to have been included at all. wilcler B. Bancroft, writing a six-page review for theJournal of Plrysical chemzsrry noted that the Mathews chapter was decidedly the weakest. After quoting passages related to psychism and the soul of

4o Jane Maienschein LS,-reviewed Cowdry,s volume and Wilson,s luded ,,too concluded that "this sort of speculative metaphysics may that much goocl cannot atolrìsr Bancroft ¡e saiã but should not have been allowed in a to.science. While, in a popular article; it however, Wilson,s booì< Ue;ustina5t. ¡dies of the nt" this', (Bancroft rgz5, ro7). Fortunately, all the reviewers seem to American investigators rvhiclt ¡oåt are a stepping_stone. agreedthat the other chapters ranged from very useful to excellent. Wilson has mainly have litions of his Indeedl Yetwhat Mathews shows, alongside the collection of chapters, is book, on questions concern_ zneralCytology ideas about cells thatwere available in r924. While researchers includes agreat deal which therangeof vo books learned agreat deal, much remained to be learned. Cowdry's volume together are a splend id had míse au stone. And the steps forward lead to more study of ning that most funda¡ was, in fact, a stepping hinted at in the Cowdryvolume but became increasingly as a crassic vorume, ,"ï:lä1:,iJ:llîj:i areas that are only ed to new important, such as cell-cell interaction, cell signaling, gene transcription ideas and direcdons for future ziew blArvilla and regulation, and so on' including the range of topics discussed in other Meek Taylor in the Chicago in ourvolume' narized the book and concluded essays with a the cells themselves, not interac- rojects, ,,for nothing Deeper underStanding.of if of their will do more to ad_ ole tions, began to appear in the 19z8 and tggz editions of Special Cytology (of than such efforts as this,,(University twoand three volumes, respectively) that Cowdry edited. Here, too, Cowdry ews, though brought together a collection of authors. He explained that the purpose was a number of them did find nistry decidedly to present in more detail knowledge about the different kinds of cells. "The odd. CytologisrJ. Brontë e.d review, book," he explained, "is to be regarded as supplementary to an earlier vol- in which he pointed out what ,,1:"r, though ume called G eneral Cytology. " There, the authors looked at "the fundamen- he apptauded rhe volume r discussion tal principles of architecture and activity which cells possess in common" of chemistry and psychism, standing (Cowdry r928, vii). Special Cytology looked instead at the characteristics of how life emerges. It is worth cells. The thirty-seven chapters allowed room for a variety of rly, because Cowdry, specialized as editor, had al_ :stence of types of cells as well as some of the methods used to study them. the views Mathews espoused lotyet clear on how cells gain life, nor Conclusion s. Mathews wrote, ,,It is in fact the lu_ rings This brings us back to the r94o celebration of a century of cells in The Cell alive, for ether is the storehouse an space and Protoplasrz. That 2os-page volume included relatively short chapters and time; energy and time,, ''s response by very distinguished researchers on cells, protoplasm, cell walls, chro- seems reserved in its cri- cytologist mosomes and genes, enzymes, molecular structure, plant hormones, vi- adequately to comment on physiology, viruses, ething to the tamins, differentiation, microorganisms, techniques, metaphysician, but one vs'views and a chapter by Charles I(ofoid on "Cells and Organisms." on the relationship between biochemistry While these researchers had acquired more knowledge about details of development, would by this 1940 volume, is papers 5, 186). symposium and it striking how many of the ng thatùtathews,s again include acknowledgements about how much remained to be learned. chapter ought not Bancroft, Conklin noted that "the mystery of mysteries is not the mechanism of evo- writing a six_page review for lution, the of the cells and protoplasm that the Mathews but evolution mechanism bywhich chapter was decid- :s related came to have the organization that has resulted in 'the promise and potency to psychism and the soul of of all life.' This is the great problem which is sure to occupy increasingly

~~'rschein~~

~~Changing ldeas about Cells as Complex Systems 41 tlre attention of biorogists in trre f'ture" (Mourton t94o, tB).Richarcl Gorcl- schmicrt poi'tecl to the neecl to get past thinr~~

42 Jane Maienschein 'e" (Moulton t g4o, tB). Richard Gold- Acknowledgments Arizonâ state university, and the webster rast thinking in terms of individual the National science Founclation, ii**;" Richard Creath' and l.ote generous and timely support' and to Karl Matlin' I on connections ancl chromosomes, r,ounclation for for valuablc suggestions' and not fully understood yet. But the ,ti".c"t¿ :d by hiding the head in the sand, or References ry calling names" (Moulton tg 66). Man and His science. prir"rceton, Nf : 4o, iu"r,ä"r** rg7B. Tìromas Hunt Morgân, the ck of completeness, or ". Prcss' to remaining Pri nceton UniversitY rB34' "Die Metamorphose des Eies der Batrachiervor der Erscheinung :ytology in r94o was still in its early U"er, nrtiU^ttvon' Erz e.U.gu,t'g." Müller'sArchiv "'äî,'Ooo und Folgeruugen aus ihr für die Theorie der physiotogie, wissenschaftlíche Medízin' äì t n.atomíe, und 481-5o9' Mirsky edited The Cell: Biochemistry, ,.The Cell-Theory: A Restatement, Hisrory, and critique-pårr I." ionn onn¿al. rg48. """q-iånury (r): (All parts are :w to five large volumes and showed "o,lJr, o.fMicroscopical sbience, series 3, Bg ro3-25. ^ruiew I had expanded. They recognized the reorintecl in Bal

rschein Changing ldeas about Cells as Complex Systems 43 Latrllichicr, Ma.l'¡ctl l)., artcl IÌric IL I)avitls''. 2oori.,,Bovcri,s (Jrchin r,or.ì!.Ììx'cri,rcrìt: sciì Mcrogollcs ¿utcl thc listaltlishllrcnt of tl.ìc l(ole of,Nuclc¿u.(.ìhrontosotncs ill l)cvclo¡rn.rcnt." I)t:vclo¡:tnr:tttn! lliokryy 3r4 (r): I r l. Liiric, Irrarrl< n.I'robrL:ns of'r"crtíriz(ttion. ig, g. ,,rri".g,,, tJ.ivcr.sity of (ìrricag. r)r.css. ar,ri,lO,r,,,t.hcl)avalopntctttul't:hcoÌtícl.:A,t,l:rotlut:t.iotttotintbryology.tti,*v,,rt.,'",,,.y Lillir:, rìal¡rh ¡t' r945. {ìt:ttL:r.r Iliorogy und Phiros.plty o.r.orgztnisnt.orric.g.: urrivcr.sity.f (llìic¿ì.qo lrress. valious ¡rrr¡rc|s. Rarprr s. ì,iilie -' Papcrs. univcrsity or,orricirgo Archivcs. ât httl)s://www.lib.'chicago.cclu/e/scrc/fi'clirgairls/vicw.'hp./c¿ì(li(l_I.U.st,.t,onrinc .CtìMSiU). l'ove'walrrclrì r97r'"Markcr FIcnry.J¿ìcobs"inthcrePort .t rortg7<>. Iliorog.it:ur.I]uilcr:in. 4t(t ): t5 .t7. Maic'schcirl'.f aiìc' 19ti3' "lìxpctinrentnIIìi.logy irr'Ì'rarsitio.: H¿rrriso.,s ìlrr.rbry.l.¡¡y, 1 895. j 9 ro.', J ¿ u d i c s i n I:t i s to _ ry o I. n io I o g-y 6: r-o7 z7 '' rgcl t .'I'rans.ritrntittg't'ratritions i., Antcrícan Iliotogy, t BBo.l.g r.5. Iì¿ìrtirììorc, Mì): .lrtlllls I lo¡rkins Urrivt,rsity l,r.css.

zo r 1 - -- ' ' "'orgtr.iztrtiou' Ns sc'ttirui llo.nclarics ,f Ircliviclual I)evcl.'rn crtt.,, 't'ltcory 6 (t):73_79. I) iolq.icctl '---' 20 L4' riìttbryos uttcrer trtc lvlicrosco¡tc: 1)ivcrg.i,g.Mcan.ittg-s of r,írÞ.canrbricrgc, M,4: llarv¿rrcl University l,rcss. Mannirrg, I(cnnctrr r<. 1983, Btocri Apoilo oJ scicttca:,I,rtc t,íJ'e or.tirncst ïvcrctL.Just.Ncw yol.r<: Oxfirrcl Ur.r ivers i ty l:rcss. May, Iìaoul M. 7()26. "rìo.k rìevicw: Ge ncrctr cr)tor'gy, ,r,rte rì. V. oowcrry; oail ín l)eve ropntcttt atrl H c re d í ty, Icl muncl Ì1. Wi lson.,, 1s¿s S f l, r rî-. r r. Morgan, 'l'honr¿ìs Hunt. r ,,tìclnrund 94o. Bcecher tlils I,tot)íosrc,thícat tvrentoírs, 31s-42.Natio.ar Acacrc.ry o'scie'ces. #il,iili;,lliri" Ac¿rclcr.nics Press.

Mrrrrltor.r, lk)rcst ll¿ìy, ccl. r g40. 'I'he cc, ancr l,roto¡tr.ttsr,.wasrringt.., rxr: Associ¿rtion fbr thc Aclvancelneltt ofscicncc, 1,hc Sciencc press. ^r.eric¿ìrì New YorkI'íme s. r946. Obittrary f.or Cla¡:cncc Iì. McClung, Nobcl prize' z.or. Jalìu¿ìly 19. "rt'rre Nobel r)r'izc in r)hysi.rogy or Mccriciue.,, .nobelprize.or.g/nobcl_¡rrizcs/r.r.rc.clicine/lauiclat o'ri'c at rrtQr://www csl zoo t l. Hyu..qwoor<. zoog. "'.cl'ru¡cr vinccnt r;owcrry n'cl the Mirl

44 Jane Maienschein son. 2oo8."llovcri,s LolìgExperirrcltt: Seâ ia/t-nicroscopicalres4Tschwl pargei oÍrline at http://wwwbiocliversitylibrary.orgi 9 ent of the Role of Nucle¿rr Chrotnosor.nes in ¡r4þ): t-tt. irnoclelruP' Aclarn. r894. "Oll tl.re Iunclecluacyof the Ccllularl'heoryof Developlìlent, aìlcl t9. Chicilgo: Seclewick, Urrivt,rsityof Chicago prcss. Nerue ancl the Early DùvclopnìL'nt of Nerues, Particularly of the'l'l-rircl of the An Introduction Ã. to Entbryology. Ncw york; Henry ii," of Microscopical Science, series 2, Syrrrpatl-retic in Elastnobranch Qua.rtcrtyJournal rílosophy of organisnz. 37 U4Ðt87-1'12' Chicargo: University of 1943. Futldcunentals oJ'Cytology' New York, McGraw-Hill' Shaip, l-ester W. HulltMorgan, 1866-1945." htBiographicalMemoírs,2B3- Sturtevarlt,A. H. 1959."I'hotnas rs. University of Chicago Archives. online National Aczrclemy of Scieuces. Washiugton, DC: Natiollal Ac¿rclet.nies Press. linclingaicls/view.php?eadicl=ICU.SpCL 325, Monument in Biology'" Natiott, Aptil r,36r' Thorre, Franl<. 1925. "A Special Collections of the Ulriversity of Cl-ricago Library, University of Chicago Archives, s" ir-r the report lor t97o. Biological Bt¿llctín. boxtzT,folcler r. Thanlcs to I(arl Matlin for findir-rg ancl sharing this infbrrnation. Whitman, Charles Otis. 1893. "The Iuaclecluacy of the Cell-Tl-reory of Development," rlogy in'l'ransition: Harrison's Ernbryology, otu'nal of M orp ho lo glt 8: 639-58' &to7-27 .l Wíllier, ts. H. rg57. "Franl<,Rattray Lillie, t87o-t947." Ríog'raphical Memoirs, TT9-236. zrican Biology, tBBo-tgt5. Baltimore, MD: National Acadelny of Sciences. washington, DC: Nationâl Acaclelìlies Press. Wilson, E.B. 78g5. AnAtlas of Fertílizatíon ancl Karyokinesis of the Olul/?. New rclaries of Inclividual Developrn ent.,, Bíological york: Macmillau..Available online ât http://www.biocliversitylibrary.org/bibliography 16244# lsurnnttry' Díverging Meanings of Ltfe.Car-nbriclge, MA: 1896. The Cell in.Development and Inherítance. New York: Macmillan. The Cell inDevelopment andHerediô).3rd ed. NewYork: Macrnillan. Orig. :ience: The Life of Ernest EuerettJusl. New york: -. pub. asThe Cell inDevelopment (Ind Inheritance in 1896; znd ed., r9oo. iJ)tology, E. V. Cowclry; The Cell in Developme nt t):2t3-rS.

~~:her Wilson, 1856-1939.,, h.t Bíographícal iences. Washington, DC: National~~

'otop las m. Washington, DC: American ce,1'he Science press. ì. McClung,January 19. logyorMeclicine." Online athttp:I lwww Lureates/zootl, )owdryancl the Makir-rgof Gerontologyas a ìited Srâtes."/o urnal ofthe History ofBiotogy

~~s t srand Thcir B io so c Visio ial ns, 1 g oo- 1 96 o. oeb and the Engíneeringldeal in Biology.New tme de chimíe organique, fondé sue des lière. Quoted in Harris rggg,32.~~