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Human in interdisciplinary focus

Jeremy Kendal, Jamshid J. Tehrani and John Odling-Smee

Phil. Trans. R. Soc. B 2011 366, 785-792 doi: 10.1098/rstb.2010.0306

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Phil. Trans. R. Soc. B (2011) 366, 785–792 doi:10.1098/rstb.2010.0306

Introduction Human niche construction in interdisciplinary focus

Jeremy Kendal1,*, Jamshid J. Tehrani1 and John Odling-Smee2 1Centre for the of and Culture, Department of Anthropology, University of Durham, South Road, Durham DH1 3LE, UK 2School of Anthropology, University of Oxford, 51/53 Banbury Road, Oxford OX2 6PE, UK Niche construction is an endogenous causal process in evolution, reciprocal to the causal process of . It works by adding ecological inheritance, comprising the inheritance of natural selection pressures previously modified by niche construction, to genetic inheritance in evolution. Human niche construction modifies selection pressures in environments in ways that affect both , and the evolution of other . Human ecological inheritance is exceptionally potent because it includes the social transmission and inheritance of cultural knowledge, and material culture. Human genetic inheritance in combination with human cultural inheritance thus provides a basis for gene–culture coevolution, and multivariate dynamics in cultural evolution. Niche construction theory potentially integrates the biological and social aspects of the human sciences. We elaborate on these processes, and provide brief introductions to each of the papers published in this theme issue. Keywords: niche construction; gene–culture coevolution; cultural evolution; human evolution

1. INTRODUCTION Evolution entails networks of causation and feed- Niche-construction theory (NCT) originated as a back in which previously selected drive branch of that emphasizes the environmental changes, and -modified capacity of organisms to modify their environment environments subsequently select for changes in and thereby influence their own and other species’ organisms. evolution [1]. The defining characteristic of niche con- NCT provides both a philosophical shift in the way struction is not the modification of environments per we view and understand evolutionary processes as well se, but rather organism-induced changes in selection as a testable scientific theory. While the effects of niche pressures in environments [1]. The effects of niche construction on the evolutionary process have often construction have been documented across a wide been neglected in the past, it is also important to range of species including animals manufacturing note that many aspects of NCT are already incorpor- nests, burrows and webs, and plants modifying nutri- ated in standard theories of evolutionary biology, ent cycles. The papers presented in this special issue , and the human explore the phenomenon in Homo sapiens, for whom sciences. However, rather than aiming just to relabel endogenous causes of evolutionary dynamics are or reclassify established theories such as gene–culture impossible to ignore. coevolution or engineering, NCT is put to NCT differs from standard evolutionary theory better use when formulating new hypotheses, or build- (SET) in recognizing that the evolution of organisms ing a more general evolutionary framework within is co-directed by both natural selection and niche con- which other theories can be subsumed. NCT provides struction. While genetic variation is subject to natural mechanisms by which currently disconnected bodies selection through differential survival and reproductive of theory, such as evolutionary and developmental success, the selective environments themselves are biology (‘evo-devo’) [1,3,4], or human cultural evol- partly determined by modifications made by niche- ution and structuration theory ([5], see below) can constructing organisms. Hence NCT recognizes be united [6]. natural selection and niche construction as reciprocal Here, we review the fundamental principles of con- causal processes in evolution, and treats the adapta- temporary NCT, initially in the context of biological tions of organisms as products of both processes [2]. evolution, before showing how the theory incorporates cultural evolutionary processes, and how it provides a framework for consilience between the natural and * Author for correspondence ([email protected]). the social sciences [7]. Where appropriate, we give One contribution of 13 to a Theme Issue ‘Human niche short summaries of each of the contributing papers construction’. in this theme issue.

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786 J. Kendal et al. Introduction. Human niche construction

2. FUNDAMENTAL PRINCIPLES OF NICHE environments, natural selection pressures. SET is CONSTRUCTION also fully consistent with the traditional view An early advocate of the niche construction perspec- expounded by Mayr [11], who regarded natural selec- tive, Lewontin [8], neatly summarized the differences tion as the ultimate cause of phenotypic characters between standard evolutionary theory and NCT in [12]. It is a view that devalues so-called proximate two pairs of coupled differential equations. His first causes, including developmental processes such as pair (equations 2.1a,b) summarizes SET: learning, and human cultural processes in evolutionary biology [13,14]. In SET, niche-construction effects dO ¼ f ðO; EÞð2:1aÞ caused by developmental or proximate processes can dt be regarded as the expression of phenotypic plasticity and [15], or sometimes as extended phenotypes [16], but ultimately they still have to be explained by prior dE natural selection [17]. ¼ gðEÞ: ð2:1bÞ dt For many purposes, SET is sufficient, but it is insufficient when genetic selective environments are In equation (2.1a), evolutionary change in - modified as a of phenotypic variation derived isms, dO/dt, depends on both organisms’ states, O, from so-called proximate processes such as learning. and environmental states, E. In equation (2.1b), For example, human cultural variation, depending lar- environmental change, dE/dt, depends exclusively on gely on differential social transmission of information environmental states. In general, organisms are not through social learning, may result in cultural niche- treated as the cause of any evolutionarily significant constructing practices that modify the natural selection changes in their environments, with the exception of of some human genes. As the selected genes may also cases such as frequency dependent selection, influence human cultural practices, the assignment of selection, maternal inheritance and coevolution. ‘causation’ becomes complex. NCT replaces SET’s Instead adaptive evolutionary change is assumed to dichotomous proximate and ultimate distinction with be governed exclusively by a single ‘causal arrow’, ‘reciprocal causation’. of organisms natural selection. depend on natural selection that is modified by niche However, SET underestimates the significance of construction, and niche construction that is selected the fact that, to stay alive, organisms must be active by natural selection [1,2]. In this light, niche construc- as well as reactive relative to their environments. tion is a mechanism of endogenous causation, reciprocal Organisms must gain resources from their external to natural selection in the evolutionary process. environments by genetically informed, or in animals, NCT asserts that, as a consequence of ancestral possibly brain informed, fuel consuming, non- niche construction, offspring inherit not only genes, random work [3]. They must perturb specific com- but an ecological inheritance, in the form of modified ponents of their environments, often at locations local selective environments relative to genetic fitness. chosen by the organisms themselves, and they must Overall, each offspring actually inherits an initial excrete to their environments throughout organism–environment relationship, or ‘niche’, from their lives [1]. Organisms are, therefore, compelled its ancestors such that: to modify some natural selection pressures in their environments by the accumulating consequences of NðtÞ¼hðO; EÞ; ð2:3Þ their activities. Lewontin captured this point by his second pair (equations 2.2a,b) of equations that, in where N(t) represents the niche of a of effect, summarize NCT: organisms O at time t in an environment E. The dO dynamics of N(t) are driven by the interaction ¼ f ðO; EÞð2:2aÞ dt of both population-modifying natural selection pressures in E, and by the environment-modifying and niche-constructing activities of , O [3]. dE In effect, this innovation replaces an ‘externalist’ ¼ gðO; EÞ: ð2:2bÞ theory by an ‘interactionist’ theory of evolution [10]. dt A niche is a neutral explanatory reference device. It In equation (2.2a) change in organisms, dO/dt, is again can capture reciprocal causation in evolution without assumed to depend on both organisms’ states and imposing any bias either in favour of natural selection environmental states, but in equation (2.2b) environ- and against niche construction, or vice versa [3]. Con- mental change, dE/dt, is now assumed to depend on ceptually, it permits differential natural selection to be both environment states, and the niche-constructing treated no longer as a function of external environ- activities of organisms. Therefore, equation (2.2b) ments, but, where appropriate, as a function of introduces the second ‘causal arrow’ in evolution that organism–environment interactions. Odling-Smee [9] called niche construction. In humans, much niche construction is influenced The philosopher Godfrey-Smith [10] highlighted by socially transmitted behaviour This observation the same distinction between SET and NCT by provoked Laland et al. [18] to propose a triple inheri- describing SET as an ‘externalist’ theory of evolution. tance evolutionary framework, delineating genetic, SET is externalist because it seeks to explain the cultural and ecological inheritance systems. Either internal properties of organisms, their adaptations, genetically or culturally influenced behaviours can exclusively in terms of properties of their external modify an environmental that subsequently

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Introduction. Human niche construction J. Kendal et al. 787 contributes to a human ecological inheritance across natural selection generations. In Laland et al.’s models the inherited environmental resource was originally assumed to be time a material or energetic resource previously modified by cultural niche construction. The inherited resource might then affect either a human cultural process with- niche construction out having any effect on human , or it could affect the natural selection of human genes including, semantic information* sometimes, the natural selection of genes that sub- ecological genetic and sequently influenced the expression of human inheritance inheritance cultural processes [18–20]. physical Riede [21] in this issue argues that this triple inheri- resources** tance system (genes, culture and ecology) can provide natural selection an effective framework to study archaeological data, and he reviews a variety of cases where provides signatures of human niche construction activity within this system. A problem with using the archaeological record is that it can be difficult niche construction to distinguish causal relationships between niche- constructing traditions and traditions selected as Figure 1. A schematic diagram of NCT. *Includes cultural knowledge; includes material culture. consequences. To overcome this, Riede advocates ** the use of phylogenetic comparative methods used to study correlated evolution in biology. He demonstrates the potential of this approach in a case study investi- of informatic and energetic/physical resources. The gating the causal relationships between traditions for delineation assumes a working definition of semantic reindeer economies and dog use in Late Palaeolithic information to be ‘anything that reduces uncertainty populations of southern Scandinavia, using a cultural about selective environments, relative to the fitness phylogeny derived from artefact traditions. interests of organisms’ [3, p. 184]. In addition, the Recently, Odling-Smee [3,22] suggested that inherited can include epigenetic infor- Laland et al.’s triple inheritance system is unnecessarily matic and physical resources that lie internal to the complicated and constraining. Instead, the original organism, such as the epigenetic inheritance of DNA cultural and ecological inheritance systems can be col- methylation patterns or the cytoplasmic inheritance lapsed into a single ecological inheritance system of nutritional resources [14,25]. These ‘evo-devo’ con- consisting of informatic as well as physical material siderations at the cellular level are beyond the scope of and resources (figure 1). This simplification is this theme issue. consistent with the idea that an individual can inherit Systematic changes in developmental environments both a social and a physical niche that can include cul- can also result in systematic changes to the phenotypic turally transmitted knowledge and behaviours, as well expression of developing organisms [26]. For example, as material culture, providing both culturally modified the construction of a developmental niche may modify sources of information and culturally modified phys- the shape of the relevant norm of reaction by reducing ical resources, in an individual’s developmental the range of developmental environments to which environment [23,24]. juveniles are exposed [2,6]. Animal burrows and Semantic information in an individual’s inherited nests typically buffer variation in environmental vari- niche might take the form of a behaviour, demon- ables such as temperature and humidity, while strated by peers or elders, acquired through social human habitation and clothing provide similar roles. learning, such as subsistence practices or social Constructed human social environments may also norms. Inherited physical resources could refer to affect behavioural development. For instance, activi- aspects of material culture, for example, nutritional ties such as play and teaching can provide scaffolding resources or tools, created through hunter–gathering for learning [27]. Sterelney [28]inthisissueargues activity or farming. Many inherited niches obviously that the construction of developmental niche has been consist of both informatic and physical resources: for critical for the evolution of behavioural modernity in instance, farmed livestock and crops are not just nutri- humans. In particular, he asserts that in the context tional resources, but also a source of public of demographic expansion in the Upper Palaeolithic, information concerning subsistence practices. the construction of structured learning environments, While the proposed two-track human inheritance which result in apprentice learning, allows high fidelity system is consistent with Laland et al.’s [18] original cultural transmission of skill sets across generations, triple inheritance model, it is founded on general prin- resulting in the behaviourally modern cultures. ciples that should apply to all organisms. Informally, evolution based on the transmission of adaptive semantic information or ‘know how’ requires energy 3. EVOLUTIONARY CONSEQUENCES OF and material resources to pay for its physical acqui- NICHE CONSTRUCTION sition, storage (whether it be in RNA, DNA or There has been considerable use of mathematical neurons etc.), use and transmission. Thus, there is a models to examine the evolutionary consequences of natural delineation between the ecological inheritance niche construction. These studies are often based on

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788 J. Kendal et al. Introduction. Human niche construction a two-locus population genetic framework, where a activities, used by small-scale human societies, to pro- genetic (or cultural) trait at one locus affects the selec- duce and raw material resources from wild flora tive environment for recipient genetic (or cultural) and fauna. He highlights how the scheme distinguishes traits at the second locus [1,18,29–34]. The research particular characteristics of wild taxa that make them has revealed interesting evolutionary dynamics such likely targets for niche construction, as well as the as momentum effects (populations continuing to proactive impact that humans have had on their own evolve in the same direction after selection has stopped subsequent resource selection as a function of yields. or reversed), time lags and inertia in response to selec- Rowley-Conwy & Layton [50] in this issue contrast tion, and sudden catastrophic responses to selection the stability of constructed niches by hunter–gatherers [6,19,29,30,35,36]. The findings have also been con- with constructed niches during the advent of agricul- sistent with quantitative genetic analysis of indirect ture. Considering a wide variety of plant manipulation genetic effects and maternal inheritance [37–40]. and hunting activity, they show how hunter–gatherers Cultural niche construction can affect either genetic can proactively alter both the and or cultural evolutionary dynamics (or both), depending evolutionary dynamics of the affected species. The in part on the relative intensity of selection. Theory authors examine the role of niche construction in suggests that human gene–culture coevolution will typi- the development and geographical expansion of both cally occur if a genetic selective environment remains cereal and livestock agriculture, and highlight the stable across sufficient generations for natural selection feedback effects of population expansion on niche to act on human genetic variation [1]. Human evolution instability. may be unique insofar as our cultural capacities and Gerbault et al.[51] in this issue review the current adaptive cultural niche-constructing activities reinforce understanding of perhaps the most well-cited case of and amplify each other [1,19,41,42]. Thus, the gene–culture coevolution, that of lactase persistence capacities for social, technical or cultural intelligence, and dairy farming. Their paper takes an interdisci- such as language and cooperation, have apparently plinary approach, considering new genetic data, coevolved with the cumulative cultural evolution of tech- archaeological evidence and simulation modelling to nologies and social conventions that these capacities explore how this coevolutionary process took place. afford [43–45]. Focusing on the European Neolithic transition, includ- In the current issue Rendell et al.[46] use a cellular ing the spread of animal domestication and uptake of automaton model to explore local and global spatial dairy farming, Gerbault et al. synthesize these data to effects of cultural niche construction on gene–culture give a contemporary explanation for the observed distri- coevolutionary dynamics. Similar to runaway sexual bution of lactase persistence, highlighting the role of selection, they explore coevolution through ‘hitchhik- both demography and niche construction. ing’ between cultural transmission of a behaviour The role of gene–culture coevolution on the evol- (equivalent to the mate preference) that modifies the ution of human sociality is explored in this issue by local selective environment of a genetic trait (equival- Gintis [52]. This paper provides a formal argument ent to the preferred trait), even when there is an that culture is not a by-product of genetic evolution, inherent cost associated with either the cultural trait but rather that culturally constituted aspects of the or the genetic trait. They also examine the unique social environment have driven the genetic evolution spatial influence on the evolution, through secondary of predispositions for cognitive features such as proso- hitchhiking, of a genetic trait that affects the capacity cial emotions and moral cognition. The paper draws for cultural niche construction, but bears an inherent on both theoretical and experimental literature to cost. The findings show the potential importance of support the case for the impact of gene–culture coevo- cultural niche construction influencing, for example, lution on, for instance, the internalization of norms, genetic evolution of disease resistance and hominid altruism and character virtues. brain size. The theme of human sociality is continued in this Gene–culture coevolutionary dynamics are likely to issue by Ihara [53], who develops a mathematical have been particularly important in recent human model to examine how culture-dependent discriminate evolution by influencing processes such as global dis- sociality could have evolved by gene–culture coevolu- persal and migration, language evolution, behavioural tion. Using the scenario of a Hawk–Dove game to modernity and sociality, the advent of agriculture, elicit resource , Ihara shows how a culturally and the evolution of human and domesticate diseases transmitted trait can alter the selective environment to [20,47,48]. This is consistent with evidence for favour the genetic evolution of culture-dependent dis- recent and rapid genetic selection, affecting character- criminators that exercise either in-group favouritism or istics including skin pigmentation, body shape, prestige bias as a function of the cultural trait distri- dentition, brain function, metabolic efficiency and dis- bution. Ramifications for the evolution of discriminate ease resistance [20,47]. The impact of cultural niche sociality include the intriguing possibility that the construction and gene–culture coevolutionary evolution of this capacity in Homo sapiens,butnotNean- dynamics on both human technological and social derthals, contributed to their contrasting fates during the evolution are considered in the current theme issue. Middle to Upper Palaeolithic transition. One of the most powerful examples of such changes Ihara notes that his model is also consistent with a is in humans’ exploitation and modification of natural cultural practice influencing the cultural, rather than resources. In this issue, Smith [49] draws on a wealth genetic, evolution of discriminate sociality. In general, of fascinating examples, largely from North America, it is probably more common for cultural niche con- to develop a classification of niche-constructing struction to result in a cultural, rather than a genetic

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Introduction. Human niche construction J. Kendal et al. 789 response. Feasibly, trans-generational cultural niche from uxorilocal to virilocal marriage practices in early construction can modify environments in ways that twentieth century Tawain, they develop a mathemat- favour ever-more culture, causing cultural niche con- ical model to explore the interaction between social struction to become ever-more powerful structure and cultural ideas. Lipatov et al. show how [1,18,23,43,54]. This process may have led to the a disjuncture between practice and belief can be accumulation of complex social norms and complex affected by changes in the economy, specifically the cultural knowledge in behaviourally modern humans. proportion of the population wealthy enough to pay Fast cultural responses to a culturally modified the observed brideprice in the virilocal system. niche can also render genetic responses unnecessary Shennan [59] in this issue chooses to focus on the [19]. For example, human-induced pollution may inheritance of material wealth, such as property, as a provoke new to remove environmental constructed niche or resource afforded by private contaminants, thus counteracting the change in the property rights that develop with agriculturalist and genetic selective environments for species across rel- pastoralist societies. Shennan explores the influence evant . Here, the initial detrimental of this niche on variation and stratification of activity could be due to ‘inceptive’ niche construction, reproductive strategies, and in addition, uses McNa- while the cultural response is likely to depend on mara & Houston’s [60] model for non-genetic ‘counteractive’ niche construction. Similarly, drug inheritance of phenotypic quality to bring insight to treatments to prevent diseases may relax genetic selec- the importance of inter-generational transfers of land tion for disease resistance or susceptibility. For wealth on long-term reproductive success. instance, Boni & Feldman [55] develop a mathemat- ical model to examine how antibiotic use, favouring selection of resistant bacterial strains, can result in cul- 4. INTEGRATING HUMAN, BIOLOGICAL tural selection for the avoidance of antibiotic use. This AND SOCIAL SCIENCES example is also notable as a case of interspecific niche Human and social scientists have been reticent to construction, as the cultural and genetic evolution of make use of evolutionary theory in the past for several antibiotic use and bacterial strains, respectively, reasons. One is that human scientists are predomi- modify the selective environments of one another. nantly interested in human behaviour and culture, NCT may be particularly relevant to the dynamics rather than genes, and as a consequence they have of cultural traits because the theory can incorporate little use for a standard theory of evolution that is the effects of cultural backgrounds, or environments, exclusively driven by natural selection acting on gen- as components of constructed niches, affecting selec- etic variation. A second reason is that an tion between cultural variants [56]. This point is adaptationist account of behaviour derived from illustrated by theoretical studies of fertility control SET, for example in evolutionary , is and the demographic transition. Ihara & Feldman regarded somewhere between oversimplification and [31] examined the effects of a preference for a high misrepresentation [61]. or low level of education on the evolution of small NCT addresses both these issues by accounting for family size. They assumed that the average level of the proactive role of human development and cultural education can affect the degree to which traits are processes in human evolution through the modifi- transmitted obliquely rather than vertically, for cation and ecological inheritance of selective example, from teachers rather than from parents, to environments. The inherited selective environment pupils. They found that a preference for small family can pertain to any form of ecologically inherited size can evolve if individuals with few offspring are semantic information, including culturally inherited more likely to transmit their fertility preference to the information, as well as physical environments. This offspring generation than are individuals with a high enables human scientists to explore human phenotypic number of offspring. This study revealed the classic variation from the perspective of genetic, ontogenetic niche-construction characteristic of a time-lag between and cultural processes operating at distinct, but the increase of the average level of education and a richly interconnected levels [62], as exemplified in subsequent decline in fertility: a pattern that is consist- many of the papers in the current issue. ent with, and may partially explain, a typical Another source of reticence is the perception among demographic transition. In a related study, Borenstein social scientists that evolutionary theory cannot et al.[34] developed a cultural niche- account for cultural diversity, but only for pan- construction model where the frequency of a trait, human traits. It is, therefore, unable to offer social such as the preference for a high level of education, scientists any insights into the phenomena that primar- affects the construction of a social interaction network, ily interest them. However, NCT provides a through which other cultural traits may percolate. framework for the quantitative examination of cultural They found that local between-population cultural diversity through modification of cultural and social niche construction could account for the spread of selective environments to affect local cultural histories reduced fertility preference across countries occurring and promote additional cultural diversity, where it is at ever-lower levels of development [57]. unnecessary to consider genetic fitness consequences Lipatov et al.[58] in this issue distinguish between a [63]. Furthermore, it is now well established that a social niche, referring to the structure of expected quantitative evolutionary model can be of great utility social roles, and a cultural niche, referring to a set of to study cultural diversity [56,64–66]. socially transmitted symbolic or meaningful ideas. There are also some parallels between the evol- Drawing on rich ethnographic data relating to a shift utionary framework offered by NCT and some recent

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790 J. Kendal et al. Introduction. Human niche construction developments in social theory that have tried to trans- 100 000 years or so, humans have become increasingly cend the classic dichotomy between structure (the reliant on physical and semantic resources that have rules and institutions of societies) and agency (the been shaped by the cultural activities of preceding gen- intentions, motivations and performances of individ- erations—from domesticated animals and tool-making uals). So-called ‘structuration theory’ [5] is based on to writing, the built environment and even religious a similar premise to niche construction in that it cosmologies. Such inventions have in various ways emphasizes that social structures are both the context both depended on and then subsequently shaped the and the consequences of individual actions. Cultural evolution of genetic and other cultural traits. Niche meanings, moral orders and the distribution of econ- construction provides a unique paradigm for studying omic and political power constrain agency and these relationships that explicitly recognizes the inform its goals, but they also depend on it for their reciprocal influences of cultural evolution, cultural . Humans’ ability to assess the effective- evolvability and gene–cultural evolution on one ness of their behaviours (‘reflexive monitoring’) another. allows them to manipulate and occasionally even to Empirical investigations will not be easy, however. transform structure, which can have intended and The eco-evolutionary feedbacks generated by cultural unintended consequences for their and others’ future niche construction are typically complicated, and behaviour. they are likely to demand multi-disciplinary The importance of evaluative and purposeful approaches [70]. At present, different disciplines, ran- agency has clear implications for our understanding ging from population geneticists, ecologists and of human niche construction, as Lansing & Fox [67] molecular biologists to anthropologists, archaeologists discuss in their contribution to this issue. They and economists, contribute different datasets and describe how the engineered landscape of Balinese different theoretical interpretations to different arcs rice terraces is governed principally by local farming of these feedback cycles. In future, a better under- associations responding to the ecologically inherited standing of human evolution may only be achieved conflicting interests of water availability and pest con- by closer between-discipline cooperation, and the trol. The development of rice cults has played a crucial mutual sharing and integration of different bodies of role in coordinating farmers’ behaviours, but has also theory from different disciplines [20]. This promises generated a wide range of new cultural representations to be an illuminating, though challenging, enterprise. and paradigms, including the agricultural calendar, The papers represented in this issue constitute an cosmological system and religious consciousness. encouraging start.

5. CONCLUSION This theme issue is borne out of an interdisciplinary meeting We can gain a richer understanding of evolutionary on human niche construction hosted by the Institute of processes by accounting explicitly for phenotypic Advanced Study and the Department of Anthropology at modification of selective environments that can result Durham University, in association with AHRC Centre for in the ecological inheritance of semantic and physical the Evolution of Cultural Diversity. We thank them for resources (figure 1). Thus, niche construction pro- their financial and academic support, which was vides an endogenous causal role in evolution that is fundamental to the development of this theme issue. We reciprocal to that of selection. In recent human evol- also thank Kevin Laland for his helpful comments on an early draft. Finally, we thank all the contributors to this ution, the most potent human contributions to theme issue for their enthusiasm, hard work and human ecological inheritance have probably occurred cooperation. J.K. and J.T. are funded by RCUK Academic through cultural inheritance. If so, that requires the Fellowships. investigation of human evolution in the context of a theory of gene–culture coevolution that explicitly includes cultural niche construction. One potential advantage of combining NCT with gene–culture co- REFERENCES evolutionary theory is that it should make gene–culture 1 Odling-Smee, F. J., Laland, K. N. & Feldman, M. W. 2003 Niche construction: the neglected process in evolution. coevolutionary models more empirically tractable by Princeton, NJ: Princeton University Press. including NCT’s mechanisms of niche construction 2 Laland, K. N. & Sterelny, K. 2006 Seven reasons (not) and ecological inheritance, both of which are open to to neglect niche construction. Evolution 60, 1751– investigation [1]. 1762. (doi:10.1111/j.0014-3820.2006.tb00520.x) Niche construction provides a powerful framework 3 Odling-Smee, F. J. 2010 Niche inheritance. In Evolution: for bringing together different perspectives on the the extended synthesis (eds M. Pigliucci & G. B. Muller), role of culture as a selective force in human evolution pp. 175–207. Cambridge, MA: MIT Press. that have developed in recent decades [68]. When 4 Laland, K.N., Odling-Smee, F.J. & Gilbert, S. F.2008 Evo- framed in terms of SET, these approaches often devo and niche construction: building bridges. J. Exp. Zool. seem to be in conflict with one another—for example, B 310, 549–566. (doi:10.1002/jez.b.21232) 5 Giddens, A. 1986 Constitution of society: outline of the over the question of the extent to which culture is theory of structuration. Berkeley, CA: University of ‘adaptive’ [69]. In NCT, these differences largely dis- California Press. appear. Rather than focusing on the adaptiveness of 6 Laland, K. N., Odling-Smee, F. J., Feldman, M. W. & cultural behaviours relative to an external environ- Kendal, J. R. 2009 Conceptual barriers to progress ment, NCT recognizes that culture is a crucial part within evolutionary biology. Found. Sci. 4, 195–216. of our ‘ecological inheritance’. Thus, over the last (doi:10.1007/s10699-008-9153-8)

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