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ANTH 3500 Module 6 Assessment Laland et al. and Fuentes

Answer the following questions with your group in essay format, using specific details that demonstrate that you engaged meaningfully with the readings.

Fuentes A New Synthesis: Resituating approaches to the evolution of human behavior

1.How does Developmental Systems Theory (DST) allow us to transcend narrow, neo-Darwinian approaches to understanding variation? What are its main theses?

2. Fuentes introduces seven proposed guidelines for creating a framework for understanding the evolution for human behavior. What are they? Using the example of the co-evolution of humans and their pets provided in the text, assess whether you think this guidelines help create a more accurate and nuanced understanding of evolution.

Read the following and answer the 2 questions

A new synthesis

Resituating approaches to the evolution of human behaviour

By all means let us seek a way of embracing human history and culture within a wider concept of evolution: not, however, by reducing history to a reconstructed phylogeny of cultural traits but by releasing the concept of evolution itself from the stran glehold of neo-Darwinian thinking, allowing us to understand the self-organizing and transformational dynamics of fields of relationships among both human and non-human beings. (Ingold 2007)

The success of humans as a species can be attributed largely to our tendency towards extreme alteration of the world around us. We not only construct material items, we engage in the creation and navigation of social structures, space and place in a manner unequalled by other organ isms. Most anthropologists would agree that humans can be seen as constructed by, and involved in the construction of, a conflux of biological, behavioural and social contexts. However, many anthropologists refuse to acknowledge a significant role for biological features and biological his tories (evolution) in human action, sensation and engage ment. Anthropology can and should counter this trend by incorporating theoretical and research perspectives that have constructivist evolutionary elements and placing them in transaction to improve both our toolkits and our significance (Schultz and Fuentes 2007, Schultz in press). Following Tim Ingold (2001, 2007) and Jon Marks (2004), I suggest that if anthropology in general becomes time. more familiar with the offerings of ‘post-neo-Darwinian’

evolutionary perspectives, we will see that there is indeed a significant space for mutual enrichment and overlap between evolutionary and anthropological approaches. I agree with Jon Marks’ call for a true Darwinian anthro pology; ‘[a] Darwinian anthropology should be as cultur ally sensitive as any other kind of anthropology; and if Darwinism is a missionary activity it needs to be regarded as of a piece with other missionary activities’ (Marks 2004:

191). However, I go one step further and want to discard the labels of ‘Darwinian’ and ‘neo-Darwinian’ for many of the evolutionary theories of interest, and recognize that there is an expansive body of research and theory that is insufficiently captured under those headings. Basic neo-Darwinian theory prioritizes natural selection and sexual selection as the prime factors in evolutionary change and the emergence of adaptations. Natural selection is the process by which certain phenotypes (morphology and behaviour) that are most effective at reproducing them selves (and thus their genetic basis or genotype) in a given environment become more frequent in a population across generations. Sexual selection is the over-representation of specific phenotypes across generations as a result of mate choice or intrasexual competition. Those traits that lead to the success of particular phenotypes and become the predominant traits in subsequent generations are termed adaptations. These traits, and the individuals possessing them, are then said to be more ‘fit’ (better at reproducing themselves into subsequent generations) than other indi viduals in the same population with less successful traits. The basal neo-Darwinian paradigm holds that most sys tems will strive for optimality, with the result that the most ‘fit’ phenotypes and their associated genotypes will rise to a majority status within the population over evolutionary

Without discounting the presence and important role of natural and sexual selection in biological systems, I want to emphasize that we have expanded on Darwin’s contri butions, and I hope to illustrate theoretically where and how this is happening as it most relates to the interests of anthropologists, especially those who prioritize a con structivist orientation. Anthropology, in a general sense, should be concerned with evolutionary theory, and incor porate relevant perspectives well beyond those that focus

1. In this context

‘epigenetic’ is used to refer to physiological/biological process of the body at a level of organization above the DNA but still within the body proper.

2. Jablonka and Lamb actually make the case that some signing chimpanzees and other apes are participating in a symbolic system as well. However, the intraspecific transmission of such a system is currently debated.

3. Here broadly defined as those animals that share human space and place, are considered distinct from other types of animals by the individual and/or local culture, form integral parts of the human social network, and are the subject of a strong, and potentially lasting, emotional and physiological bond between human and non-human animal. See Olmert (2009), Fuentes (2006), Haraway (2003), and Mullin (1999) for more detail. I have coined the term Audrey effect to refer to this system. 4. In the neo-Darwinian paradigm altruism, or acts that have a net loss of energy (ultimately fitness) to the actor but a net gain in energy (ultimately fitness) to the receiver, does not make sense if organisms benefit

by maximizing their own fitness. However, the idea of kin selection (proposed by William Hamilton) offers a simple equation that predicts when an individual organism might behave in a manner that looks altruistic. This equation is r x b>c, where r = genetic relatedness between the actor and receiver of the behaviour, b = the fitness benefit to the receiver and

c = the fitness cost to the actor. If the individual who receives the benefit from a behaviour that costs fitness to the actor is a relative, then a certain percentage of the actor’s genotype (depending on the degree of relatedness) also benefits from the action. Reciprocal altruism (proposed by Robert Trivers) acts to explain apparently altruistic acts between unrelated individuals. The basic model is that unrelated organisms can enter into relationships that can be characterized as fitness value exchanges. Using a simple Mendelian genetic system as a basis, Trivers presents a mathematical equation that outlines the relationship variables between an actor and a recipient in a series of reciprocal exchanges, using a prisoner’s dilemma-style ‘payoff matrix’.

exclusively on the action of natural and sexual selection as the prime drivers in evolutionary change. Here I review emergent perspectives in evolutionary theory that may be unfamiliar to many anthropologists, and offer a framework for asking questions about human behavioural evolution that seeks to provide space for evolutionary perspectives in anthropological enquiries.

Emerging evolutionary theory: A mini-primer

Three recent trends in evolutionary theory – multi inheritance systems theory, developmental systems theory, and niche construction, offer fertile arenas for anthro pology (see Table 1). The biologists Eva Jablonka and Marion Lamb (2005) call for a renewal in evolutionary theory at the start of the 21st century, a ‘new’ new synthesis in how we model evolution. They argue for recognition of ‘evolution in four dimensions’ rather than a focus on just one – their main point being that practitioners of traditional neo Darwinian approaches focus on one system of inheritance, the genetic. Because of this the majority of hypotheses proposed for scenarios of the selection and adaptation of human behaviour (as a phenotype) rely on, or are derived from, perspectives based on explanations of causal fac tors at the level of the DNA, some proxy for genic effect, or an assumption of some unidentified link to genetics. Jablonka and Lamb argue for a new perspective taking in three other inheritance systems which may also have causal roles in evolutionary change. These other systems are the epigenetic, behavioural and symbolic inheritance systems. Epigenetic inheritance 1 is found in all organisms, behavioural inheritance in most, and symbolic inheritance occurs only in humans. 2 ‘Information is transferred from one generation to the next by many interacting inheritance systems […] Variation is also constructed, in the sense that, whatever their origin, which variants are inherited and what final form they

assume depend on various “filtering” and “editing” proc esses that occur before and during transmission’ (Jablonka and Lamb 2005: 319; italics in original). This view moves beyond standard neo-Darwinian approaches. Many organ isms transmit information via behaviour; thus acquisition of evolutionarily relevant behavioural patterns can occur through socially mediated learning. This transmission of information occurs without any link to genetic systems that can be targeted by natural selection (in a neo- Darwinian view). Symbolic inheritance comes with language and the ability to engage in information transfer that can be temporally and spatially complex, contain a high density of information, and convey more than material descrip tions. This allows for the acquisition and reproduction of a variety of behaviours, perceptions and beliefs potentially beneficial for human populations but having no genetic basis or connection. In the quest to understand human behaviour, the behav ioural and symbolic inheritance systems are obviously of great interest to us. This possibility that ‘instruction’, the passing of non-genetic information or structure across gen erations, can influence evolutionary patterns, changes the way we can envision human evolution. Models using this system will by necessity become more complex than the generally linear genetic models of neo-Darwinian behav ioural theory (such as kin selection, the favouring of close relatives on the grounds of their high degree of shared gen otype, for example). However, such models may be better attuned to the actual interactions of systems. Specifically in terms of anthropology, this perspective imposes an evolutionary concern with the way in which bodies and behavioural and symbolic systems construct and interact with social and ecological niches and how, in turn, these systems interact with epigenetic and genetic systems. This perspective blurs any clear prioritization in inheritance systems and requires a move away from approaches that are limited to either social or biological focuses.

Table : 1

Evolution in Four Dimensions


Systems Theory

Niche Construction

Focus of selection

Gene, epigenetic systems, behavioural

Outcome of complex between genic,

interactions epigenetic and

Individual or group and local ecologies (niche)

interaction with


behavioural factors

Main underlying causes for

Combination of genic, epigenetic, behavioural

Constantly constructing constructed by –

– and being demography, social

Tri-inheritance vision human behaviour results

model (TIV): from

evolution of human behaviour

and symbolic inheritance systems

interactions, cultural manipulation of the

variations and environment

information-acquiring levels: population genetic

processes at three processes,

in intra- and intergroup


ontogenetic processes

and cultural

in addition to the biological and

developmental, ecological factors


throughout the course

of life history

Basic premise

There is more to heredity than genes

Some hereditary variations are non random in origin

Some acquired information is inherited

Evolutionary change can result from instruction as well as selection

Evolution is not a matter of organisms or populations being moulded by their environments but of organism-environment systems changing over time. This involves joint determination by multiple causes, context sensitivity and contingency, extended inheritance, development as construction, distributed control and evolution as construction

Forces other than neo-Darwinian selection at play

Epigenetic, behavioural and symbolic modes of inheritance


Epigenetic and developmental systems, ecological and cultural inheritance

Genetic, epigenetic, behavioural and cultural

Developmental systems, behaviour, physiology

Ecosystem engineering Organisms modify their own, and other, organisms’ selective environments Ecological inheritance, including modified selection pressures for subsequent populations Is a process, in addition to natural selection, that contributes to changes over time in the dynamic relationship between organisms and environments (niches) Niche construction interacts with, and modifies, patterns and strength of natural selection Genetic, behavioural, cultural, ecological

Fig. . 3

5. My personal choice not

· to have children aside (or not considered by most neo- Darwinians as relevant).

6. To paraphrase the quote,

· it states that one would jump in a freezing river to save

· ne’s brother over a cousin, and a cousin over a stranger. This is due to the closer genetic relationship between close kin and thus fitness is enhanced relative to the degree

· f relatedness, losing value with distance to the point of it becoming maladaptive, in a strict neo-Darwinian sense, to save a stranger.

4. Drawn directly from

1. Emily Schultz and Agustín Fuentes (2007), ‘Constructivist Evolutionary Anthropology: Beyond Nature versus Nurture’. Invited session co-sponsored by the Biological Anthropology and the General Anthropology sections of the American Anthropological Association 2007 annual meetings, Washington, DC.

5. I am a biological

1. anthropologist by training, in that much of my background includes a focus on evolutionary, physiological and ecological elements in human and other primate behaviour. However, I retain the idealistic perspective that regardless of specialist focus, we can see

1. ne another as anthropologists under a ‘big tent’ of shared disciplinary goals.

Bourdieu, Pierre 1977. Outline of a theory of practice, trans. Richard Nice. Cambridge: Cambridge University Press.

— 1990. The logic of practice,

trans. Richard Nice. Stanford, CA: Stanford University Press. Boyd, Robert and Richerson,

Peter J. 2005. The origin and evolution of cultures. New York: Oxford University Press. Dawkins, Richard 1982. The extended phenotype: The gene as the unit of selection. Oxford: W.H. Freeman and Company.

Dobzhansky, Theodosius 1972.On the evolutionary uniqueness of man. In Dobzhansky, T., Hecht, M.K. and Steere, W.C. (eds) Evolutionary biology, vol.

6, pp. 415-430. New York: Appleton-Century-Crofts. Farmer, P. 1999. Infections and inequalities: The modern plagues. Berkeley: University of California Press.

Moving from multiple systems of inheritance to a focus on development, developmental systems theory (DST) is proposed as an alternative to what Susan Oyama (2000) calls ‘developmental dualism’. Oyama states that ‘[t]he developmental systems approach allows us to redraw the overly restrictive boundary around the genes to include other developmentally important influences.’ Traditional neo-Darwinian approaches view development as com bining some aspects driven mainly by ‘internal’ causes (genes) and others driven mainly by external causes (envi ronment, or cultural variants). The dualistic approach has led to the proposal of two systems of natural selection, one to carry culture and one biology, with the two mir roring one another in their mode of change (selection), mutually co-evolving, occasionally impacting one another (gene-culture co-evolution; see Boyd and Richerson 2005, Richerson and Boyd 2005). This approach casts ‘culture’ as a unit or end product of selection (as are genes), on the grounds that ‘[c]ulture is adaptive because it can do things that genes cannot do for themselves’ (Richerson and Boyd 2005: 145). As an alterna tive to this approach, DST seeks to exchange discrete dual channels for synthetically inter acting systems whose processes give rise to successive generations. DST, then, is an approach that attempts to combine mul tiple dimensions and interactants using a systems approach to understand devel opment, in the broadest sense, and its evolutionary impact.

Oyama et al. (2001) lay out a set of main theses of DST:

Joint determination by multiple

causes suggests that explanations assuming the primacy of ‘genes’, their competition for propagation, and their interactions with an envi ronment are not the only arenas for enquiry into the evolution of human behaviour.

Context sensitivity and contingency

mandates that assessment of the evo lution of human traits/patterns must take into account all developmental processes, including experiential and social contexts. Single aspects of human behaviour or morphology cannot be seen as independent, in an evolutionary sense, from any others.

tionary patterns and is especially character istic of humans; it includes the memory and experience of group members, the previous manipulation of the area in which the group lives, and the patterns of cultural interaction extant in that population.

Extended inheritance affects evolu

Development as construction posits that traits are

evolution is not a matter of organisms or populations being moulded by their environments but of organism environment systems changing over time. Human evolu

made – reconstructed – in development. Humans do not inherit pre-made phenotypes. Development is a lifelong process; the phenotype is an ongoing constructive event interactively affected by physiological, social and ecolog ical factors simultaneously during the life course. Distributed control suggests that focusing on natural selection and genic inheritance while ignoring other dimensions of inheritance is unlikely to offer an adequate explanation of the full range of human behaviour and evo lution (as per Jablonka and Lamb above). Finally, evolution as construction is the concept that

tionary patterns are constantly constructing – and being constructed by – constituent elements of demography, physiology, reproduction, social interactions, cultural variations, complex information transfer, local ecological change and manipulation of the environment in intra- and intergroup contexts throughout the course of life history. Beyond multiple modalities of inheritance and the proc esses and systems of development, niche construction is also proposed as a major force, alongside natural selec tion, influencing the evolution of organisms. Building on work of Richard Lewontin (1983) and earlier perspectives proposed by Ernst Mayr (1963) and Conrad Waddington (1959), and even drawing from the ‘extended phenotype’ concept of Richard Dawkins (1982), F. John Odling-Smee, Kevin Laland and Marcus Feldman (2003) formalized and proposed niche construction as a significant evolu tionary force. Niche construction comprises the building and destroying of niches by organisms and the synergistic interactions between organisms and environments. Niche construction creates feedback within the evolutionary dynamic, such that organisms engaged in niche construction significantly modify the evolutionary pressures acting on them, on their descendants, and on unrelated populations sharing the same space (Odling-Smee et al. 2003). Niche construction affects energy flows in ecosystems through ecosystem engineering, creating an ecological inheritance and, like natural selec tion, contributes to changes over time in the dynamic relationship between organisms and environ ments (niches). Niche construc tion is inherently a constructivist argument, which we may be able to use to reflect synthetic uni ties of ecological, biological and social niches rather than treating them as discrete spheres (Flack et al. 2006, Fuentes 2009, Odling Smee et al. 2003). Odling-Smee et al. (2003) explicitly state that ecological inheritance via material culture, and niche construction in general, can occur via cultural means. They state that humans are the ‘ultimate niche constructors’ and that adding niche construction to attempts to understand human systems makes such attempts more complicated (bypassing more simplistic neo-Darwinian adapta tionist accounts). They see cultural proc esses as providing a particularly robust vehicle for niche construction, and propose a specific model for human genetic and cultural evolution which they term a tri-inheritance vision model (TIV). As understood in TIV, human behaviour results from information-acquiring processes at three levels: population genetic processes, ontogenetic processes and cultural processes (see Odling-Smee et al. 2003, figures 6.1c and 6.2). Niche construction in humans emerges from all three of these processes, each of which can influence the patterns, contexts and structure of natural selection. Odling-Smee et al. state: ‘[m]uch of human niche construc tion is guided by socially learned knowledge and cultural inheritance, but the transmission of this knowledge it itself dependent on preexisting information acquired through genetic evolution, complex ontogenetic processes, or prior social learning’ (Odling-Smee et al. 2003: 260-261).

Fig s

Flack, Jessica C. et al.

2006. Policing stabilizes construction of social niches in primates. Nature 439: 426-429.

Fuentes, A. 2002. Patterns and trends in primate pair bonds.

International Journal of Primatology 23(4): 953-978.

— 2004. It’s not all sex and violence: Integrated anthropology and the role of cooperation and social complexity in human evolution. American Anthropologist 106(4): 710-718.

— 2006. The humanity of animals and the animality of humans: A view from biological anthropology inspired by J.M. Coetzee’s Elizabeth Costello.

American Anthropologist

108(1): 124-132.

— 2009. Evolution of human

behavior. New York/Oxford: Oxford University Press. Goodman, A. and Leatherman, T. (eds) 1998. Building a

new biocultural synthesis: Political-economic perspectives on human biology. Ann Arbor:

University of Michigan Press.

Harrison F.V. 1998.

Introduction: Expanding the discourse on ‘race’.

American Anthropologist

100(3): 609-631.

Hermann, Esther et al. 2007. Humans have evolved specialized skills of social cognition: The cultural intelligence hypothesis. Science 317: 1360-1366. Haraway, Donna 2003.

The companion species manifesto: Dogs, people, and significant otherness.

Chicago: Prickly Paradigm Press.

Ingold, Tim 2007. The trouble with ‘evolutionary biology’. Anthropology Today 23(2): 13-17.

— 2001. From

complementarity to obviation: On dissolving the boundaries between social and biological anthropology, archaeology and psychology. In: Oyama, S., Griffiths, P.E. & Gray, R.D.

(eds) Cycles of contingency: Developmental systems and evolution, pp. 255-279.

Cambridge: Cambridge University Press.

Jablonka, Eva and Lamb,

Marion 2005. Evolution in four dimensions: Genetic, epigenetic, behavioral, and symbolic variation in the history of life. Cambridge,

MA: MIT Press.

Anthropology and the new perspectives

While offering useful refinements and enhancements to existing evolutionary theory, these perspectives do not meld together without contradictions. The discrete por tioning of inheritance systems (multi-inheritance), the rejection of the ability to truly differentiate single systems in development (DST), and the use of a tripartite biology/ experience/culture construct (niche construction TIV) do stand somewhat at odds with one another. The various proposals also diverge in their view on the role of tradi tional natural selection, the relationship between selection and their processes of change, and the types of datasets amenable to analysis under their proposals (see Table 1). However, I do not believe that these discordances negate the potential for applying aspects of these and other per spectives (including elements of the neo-Darwinian view) to anthropological questions about human evolution and behaviour. We need to seek a synthesis, drawing from diverse perspectives, to have the best chance of elucidating patterns and processes.

Most anthropologists would agree that humans are constructed by, and involved in the construction of, con texts that are simultaneously physi ological, behavioural, historical and social. It is these contexts, the way in which they interact, and their pat terns and modes of processing that I suggest offer the greatest scope for substantial overlap between theo retical perspectives in social anthro pology and those presented here. Take, for example, the perspec tives contained in Bourdieu’s (1977, 1990) conceptualizations of habitus and doxa, often drawn upon by anthropologists, and their interrela tion in his ‘structured structures pre disposed to function as structuring structures’ (1990: 53). This view on human behaviour and processing resonates strongly with the elements of niche construction, ecological and social inheritance, social attention and symbolic inheritance discussed above. One can see how anthropolo gists interested in understanding the processes underlying the creation of habitus, and the initiation and perpet uation of doxic relationships, might exploit and integrate these emergent evolutionary concepts to good effect when thinking about the histories, trajectories and patterns in human behaviour. Anthropological enquiries are frequently concerned with who/what has agency in human behaviour. Current answers to these questions in the anthropological literature include elements from phylogenetic histories, social and symbolic histories, evolutionary processes, economic-political pres sures, gendered relationships, the dynamic human place circumscribed by local context (ecological, cultural, eco nomic, environmental, familial, etc.), and experiential moulding of individuals (physiological, behavioural and perceptual), just to name a few. This plethora of poten tial agents precludes simplistic or reductionist (universal) explanations for human action and thought. However, integrating such questions and explanations (at least occasionally) by exploiting elements of the per spectives summarized here may bring us closer to the anthropology envisioned by Ingold and Marks. Obvious areas of application for the type of approach I propose

here are illustrated in the recent explosion in the biocul tural anthropology of health and adversity (Farmer 1999, Goodman and Leatherman 1994, Panter-Brick and Fuentes 2008) and the integrated anthropology of race (AAA race project – see http://www.understandingrace.org/home. html, Harrison 1999, Jablonksi 2007). However, I hope that it might also be evident that a much wider swath of human behaviour and social patterns may be amenable to analyses incorporating these emergent evolutionary perspectives.

Framing questions

In the light of this overview, I would like to propose a possible framework for resituating some approaches to the creation of anthropological models and hypotheses for the evolution of human behaviour (see Fuentes 2009). This framework draws heavily from a diverse array of prac titioners and perspectives, and many anthropologists are already practising aspects of what I list here.

1) Human behavioural evolution must be seen prima rily as a system evolving, rather than a set of independent or moderately connected traits evolving. While we might create limited and trait-based hypotheses to explain specific behav ioural patterns, they must be explic itly connected to broader themes and contexts. Human behaviour itself is not a set of individual actions but rather a combination of action, expe rience and innovation. 2) Niche construction is a core factor in human behavioural evolu tion. The ability of humans to modify their surroundings is central to any explanation of human behaviour.

These surroundings include, at min imum, the social, the material and the ecological. Understanding humans requires assessing the interactive and mutually mutable relationship humans have with their social and structural ecologies. We must accept the possibility that evolutionary pressures can be modified as they are occurring and that human response to such challenges need not always fit the standard neo-Darwinian or social constructivist models.

3) Ecological and social inher itance are core to human behav iour. Humans almost always exist in a place where there have been humans before them. Behavioural parameters are influenced by the previous generations, and individuals in succeeding human genera tions inherit a much larger amount of information than do any other organisms. This area also requires some attempt at integrating morphological, physiological, social and historical avenues of enquiry. 4) Enhanced communication and information transfer is …