Review of General Psychology 2013, Vol. 17, No. 2, 184 –189
© 2013 American Psychological Association 1089-2680/13/$12.00 DOI: 10.1037/a0032932
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A Developmental Evolutionary Framework for Psychology Robert Lickliter
Hunter Honeycutt
Florida International University
Bridgewater College
Evolutionary psychology (EP) was founded on the metatheoretical assumptions of the modern (or neo-Darwinian) synthesis of evolutionary biology, which dichotomize internal from external sources of causation. By prioritizing the former, EP has promoted a preformationist view of individual development, which effectively divorces developmental from evolutionary analysis. The authors argue that these assumptions about development are in need of revision in light of recent advances in genomics, epigenetics, and developmental science. The authors outline a developmental evolutionary framework for psychology, a relational metatheory that integrates the study of developmental and evolutionary mechanisms within one explanatory framework. They argue that knowledge of the dynamics of developmental processes are necessary to illuminate mechanisms of evolutionary change and that the psychological sciences, particularly comparative and developmental psychology, are ideally positioned to contribute to this endeavor. Keywords: evolutionary psychology, unifying psychology, theory of psychology, developmental processes, dynamic systems
The modern synthesis is primarily a theory about genes, and for the better part of the past century, evolution was narrowly defined as “a change in the genetic composition of populations” (see Dobzhansky, 1937; Mayr, 1982). Evolution came to be defined in this way because of the special privileges assigned to genetic factors. As we have noted elsewhere (Lickliter & Honeycutt, 2003, 2009), genes were thought to be the exclusive means by which information is transmitted across generations and that genes encode programs or instructions that predetermine the development of traits. Moreover, genetic information was not thought to modifiable by experiential or environmental (i.e., proximal) factors encountered during the lifetime of the organism. From this view, phenotypic novelty could only come about through nondirected, chance events like genetic mutation and recombination. Natural selection thus plays the primary creative role in evolution by filtering out nonadaptive variants, and in so doing, shapes the frequency and spread of adaptive (genetically determined) traits in populations. Proponents of the modern synthesis largely focused on the evolution of physical traits, ranging from enzymes to morphological features (e.g., beak size). Behavioral traits were thought to evolve in a manner similar to physical traits, but little empirical attention was devoted to this issue in evolutionary biology (but see Roe & Simpson, 1958), likely because behavior was not thought to contribute to evolutionary change (Bateson, 2004). Sociobiology, which gained prominence in the 1970s, was the first popular attempt to explain behavior using the modern synthesis (Wilson, 1975). Sociobiologists argued that many social behaviors (e.g., infanticide, incest avoidance, male promiscuity, and altruism) of animals could be understood as having evolved by means of natural selection because they served (and continue to serve) to increase the frequency of an individual’s genes (directly or indirectly via kinship relations) in subsequent generations. EP emerged in the 1980s, focusing specifically on human behavior. EP modi-
The Modern Synthesis Applied to Psychology and Its Shortcomings Because all modern life forms evolved from earlier, now extinct forms of life, it seems evolutionary perspectives are well suited to provide a metatheoretical unity of understanding and discovery within and across the life sciences. However, when most readers hear the terms evolution and psychology paired together, they likely think of evolutionary psychology (EP), a subdiscipline in psychology that combines cognitive psychology with the general principles of the modern (or neo-Darwinian) synthesis, the prevailing theory of evolution during the 20th century. Some have claimed that EP could serve as a metatheoretical basis for psychology (e.g., Duntley & Buss, 2008), but we are doubtful in part because the modern synthesis itself has failed to unify even the biological sciences. For example, both developmental biology and ecology advanced largely independently of evolutionary biology throughout the last century (Pigliucci, 2009). Recently there has been a growing discontent with the modern synthesis among life scientists, resulting in a call out for a broader synthesis of ideas in understanding evolution (Pigliucci & Müller, 2010). Although this newer synthesis has yet to be formalized, we believe it has the potential to unify much psychological research. Likewise, we believe psychology can play a constructive role in the formation of this new synthesis.
Robert Lickliter, Department of Psychology, Florida International University; Hunter Honeycutt, Department of Psychology, Bridgewater College. The writing of this article was supported in part by NSF grant BCS 1057898 awarded to Robert Lickliter. Correspondence concerning this article should be addressed to Robert Lickliter, Department of Psychology, Florida International University, Miami, FL 33199. E-mail:
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fied the scope (e.g., an interest in cognition and social behaviors) and explanatory basis of sociobiology. A central claim of modern EP (see Barkow, Cosmides, & Tooby, 1992; Buss, 1995; Pinker, 2003) is that natural selection operates on the underlying psychological mechanisms that generate behavior. Specifically, EP argues that there exists a large set of domain-specific cognitive modules in the minds of modern humans that evolved by means of natural selection as adaptations to specific ancestral environments during the emergence of Homo sapiens. Despite major changes in living conditions and lifestyles since the emergence of our species, EP claims there has not been enough time for selection to change the genetically determined, modular nature of the human mind. Thus, to understand modern behavior and cognition, EP argues that psychologists must look to the distant past to consider the adaptive challenges faced by our early ancestors to identify the specific (now innate) cognitive solutions that evolved to solve these problems (Tooby & Cosmides, 1990). EP has been criticized on numerous grounds, ranging from its particularly narrow view of adaptation to only ancestral (and largely unknown) conditions, its appeals to genetic determinism, its falsifiability, its lack of evidence for extensive modularity of mind, and failures to replicate key findings (see Buller, 2005; Richardson, 2007; Rose & Rose, 2000 for reviews). As we see it, many of the shortcomings of contemporary EP can be traced to its ongoing commitment to the metatheoretical assumptions of the modern synthesis of evolutionary biology. These metatheoretical assumptions split the internal world from the external world, and because it places priority (or “realness”) on the former, it promotes a predetermined and nondevelopmental explanation of physical and behavioral traits (Overton, 2006; Oyama, 2000). The special privileges assigned to genetic factors effectively divorced heredity (as genetic transmission) from development, and developmental influences (viewed as proximate causes) from evolutionary processes. These divisions in turn fostered the view that developmental analyses could add no explanatory value to evolutionary theory. As a result of this type of thinking, the modern synthesis focused on population genetics during most of the last century, moving away from earlier evolutionary concerns with embryology and developmental biology (Amundson, 2005; Gottlieb, 1992). This population genetics approach concentrated on the traits of adults in populations and virtually ignored questions about how these traits were actually realized during the course of development. However, empirical and conceptual advances in the biological and psychological sciences over the last several decades have combined to establish a more integrative, epigenetic account of the stability and variability of phenotypic development. The evidence and rationale behind this epigenetic alternative to the Modern Synthesis has been a major focus of research across the life sciences in recent years and is bringing together genetics, molecular, cellular, and developmental biology, neuroscience, developmental psychology and psychobiology, and evolutionary biology to reconsider the nature of the links between development and evolution (e.g., Gottlieb, 2002; Hallgrîmsson & Hall, 2011; Pigliucci & Müller, 2010; Robert, 2004; West-Eberhard, 2003). Collectively, this work shows that the passing on of a genome in reproduction, although certainly necessary, cannot serve as a sufficient explanation for the achievement of any phenotypic out-
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come. Gene activity and expression is determined and regulated by historically construed, systemic factors in and above the level of the cell. As a result, genetic and environmental factors cannot be meaningfully partitioned, and perspectives that favor notions of prespecified or predetermined phenotypic traits are not up to the task of making sense of the dynamics of the developmental process and its varied outcomes (see Lewkowicz, 2011; Spencer et al., 2009, for examples in the development of perception and cognition). Moreover, there are multiple, experience-dependent inheritance systems beyond the traditional genetic system (see Jablonka & Lamb, 2005, for review). For example, the epigenetic system affects the expression of the genome of an individual during its development and can influence (directly or indirectly) the development of offspring without altering the coding sequence of genes. To date, the majority of such transgenerational epigenetic effects that have been identified deal with exposure to toxins, drugs, restricted diets, or temperature changes, but studies with rodents have found transgenerational effects associated with social stimulation such as maternal behaviors (levels of licking and grooming), abusive caregiving, and experimenter-provided handling (Champagne, 2010; Crews, 2008; Harper, 2005; Honeycutt, 2006). These modern advances challenge the foundational dichotomies (genetic vs. environmental; innate vs. acquired; ultimate vs. proximate; heredity vs. development; biology vs. culture) of EP and other nativist approaches throughout the history of psychology. These divisions are artificial. Not only have they outlived their usefulness in describing or explaining behavior, they currently serve to hinder rather than promote advances in the behavioral sciences. To describe a behavior pattern as innate (or genetically determined) is in fact a statement of ignorance about how that trait actually develops. As we have outlined in more detail elsewhere (Lickliter & Honeycutt, 2003, 2009), the genocentric view (as with any preformationist view) has several serious shortcomings, not the least being that it assumes as a given the developmental outcomes that actually require a causal developmental analysis. What is needed (i.e., what the data demands) is a different metatheoretical framework that can more adequately deal with the dynamics of behavioral development across multiple time-scales. In what follows, we outline such an approach and discuss its implications for psychological research.
Metatheoretical Assumptions of a Developmental Evolutionary Framework We propose a developmental evolutionary framework for psychology. We believe that psychology needs such a framework to address how to characterize the developmental dynamics involved in generating, maintaining, and transforming behavior within and across generations. The overarching goal of this effort would be to fuse approaches that emphasize developmental mechanisms and those that focus on evolutionary processes. In this sense, our developmental evolutionary framework shares much in common with the assumptions associated with other modern developmental theories (see Lerner, this issue; Michel, this issue; Overton, 2006). Although space limitations prevent a full description and development of our framework, below we describe several metatheoretical assumptions that should contribute to the course of psychology’s future:
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• Individual development takes place within a hierarchy of embedded relational systems (gene– cell–tissue– organ system– organism–proximate environments– distal environments) that reciprocally influence each other. What happens at one level in a system (e.g., genetic transcription) is causally connected to what is happening at other levels of the system (e.g., cellular environment, sensory environment, social context, and so on). • Because of the multiplicity of levels, factors, and interactions, and because of its dynamic and historical nature, the control of developmental processes is distributed relationally throughout the organism– environment system and cannot be attributed to any single resource, level (e.g., genes), or experience. Internal and external developmental resources operate as coequals, not as competing alternatives. Development is situated, contingent, and context-sensitive. • Attempts to identify traits that are innate versus acquired are both meaningless and invalid. A belief in innate traits reflects a commitment to preformationism and ultimately, mysticism. • There are numerous interrelated systems of biological inheritance beyond the genetic system. At the moment of conception there is already a structured organism embedded in a structured, multilevel environment. • Each generation uses a set of heritable developmental resources. These are reconstructed in each organism’s development through a self-organizing process that does not rely on a “central” information source. • Evolutionary changes are the result of developmental changes. The process of development both constrains phenotypic diversity as well as introduces phenotypic variation upon which selection can act. • The introduction of phenotypic novelty is not necessarily random or arbitrary. Variations of the phenotype are produced by the transactions between organisms and the environment during development. In the most general sense, the assumptions of our developmental evolutionary approach are consistent with a relational metatheory of development (Overton, 2006), with its emphasis on process, activity, change, emergence, and self-organization. When applied to psychological analysis, our framework emphasizes an empirical concern with how behavioral and cognitive traits are generated and maintained in developmental processes (see Caporael, 2003; Quartz, 2003, for similar perspectives). Given that all phenotypes, including behavior and cognition, have a specific developmental history that explains their emergence, a developmental mode of analysis is required to fully explain the structures and functions of maturing and mature organisms. Our framework recognizes the need to understand how combinations of genetic, hormonal, neural, physiological, behavioral, and social mechanisms act synergistically as a system from which behaviors emerge and are maintained within and across generations. For example, explanations of alcohol abuse must include understanding the body and brain responses to alcohol, prenatal or early postnatal exposure to alcohol, family histories and patterns of alcohol use, the motivational reasons for abusing alcohol, contextual analysis (including conditioned effects), and an analysis of social relationships and cultural norms which may support alcohol abuse. This focus on the causal analysis of development, how development actually happens, is fundamental to our proposed framework. As we put it a decade ago:
If evolutionary psychology’s aim is to understand the evolved psychological mechanisms that underlie behavior and the selective forces that have shaped these mechanisms, then it seems to us that uncovering the network of factors (both internal and external to the organism) that bring about or maintain (or eliminate) transgenerational similarities or differences in behavioral traits should be a prominent goal of EP. (Lickliter & Honeycutt, 2003, p. 829)
We continue to hold this view and our developmental evolutionary framework outlined here is an attempt to move forward on this significant challenge.
Implications for Psychological Science What are the implications of incorporating a developmental evolutionary framework into contemporary psychology? How would our proposal offer something different from the established version of EP, which has long argued that it provides psychology with its most comprehensive and profitable theoretical framework (e.g., Buss, 1995; Cosmides & Tooby, 1987). In the broadest sense, our metatheoretical framework identifies important areas and topics of research currently underrepresented within the field, while better situating the field of psychology within contemporary life sciences. The adoption of our approach in the psychological sciences demands greater attention be paid to analyzing behaviors under normally occurring ecological conditions. Because organism– environment systems are tuned together over developmental and evolutionary time scales, we should expect special relationships between organisms and species-typical ecological events (Johnston, 1985). In much of traditional psychological analysis, however, subjects are placed in ecologically arbitrary situations and tested for responsiveness to ecologically arbitrary events. Such artificial, simplified studies may be necessary to isolate potential mechanisms of behavior, but such studies often overstate or understate how important certain factors are in real-life situations. In the examples below, we highlight the importance of including ecologically relevant factors in psychological research. • Neurogenesis in adult zebra finches is known to be influenced by housing conditions. Rates of neurogenesis are higher in birds housed in larger social groups in large aviaries compared to birds housed alone in smaller aviaries or housed with one other bird (Lipkind, Nottebohm, Rado, & Barnea, 2002). Likewise, a number of recent studies have shown that neurogenesis in adult birds drawn from wild populations is higher compared to captive populations (Barnea, 2010). • Much of what we know about the acquisition of classically conditioned responses has been based on studies that involved conditioned stimuli that were ecologically arbitrary (e.g., pure tones or lights). However, when more ecologically realistic stimuli are used as conditioned stimuli in a variety of domains, the acquisition of conditioned responses are more rapid and resistant to extinction than when ecologically arbitrary (but still complex) stimuli are used (Domjan, 2005). • Early studies on song leaning by birds often used a tape-tutor paradigm in which young birds were isolated in chambers and exposed individually to recordings of songs. These results suggested that birds could only learn conspecific songs and only within a certain period of sensitivity. Later results using live tutors, however, indicated that not only could birds learn songs well
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beyond the proposed sensitive period for learning, but in this more ecologically realistic situation, could even learn songs of a different species (Baptista & Petrinovich, 1984). • Traditionally, in studies of human infant perception researchers often presented stimulus events in a single sensory modality (e.g., visually or acoustically). However, infant perceptual and cognitive abilities are enhanced when researchers use more realistic events in which information is simultaneously conveyed across multiple sensory modalities (Bahrick & Lickliter, 2000). By placing organisms in ecologically arbitrary physical (e.g., laboratories or operant chambers) and social (testing alone vs. in groups) situations and exposing them to ecologically arbitrary stimuli (e.g., pure tones to study associative learning, line drawings of figures to study attraction, face-like stimuli to study infant face preferences, still shot photographs to study emotional discrimination), psychologists have made great strides in understanding how and what organisms can do or learn (i.e., in understanding potential mechanisms of behavioral change), but such progress has often been at the expense of understanding what and how organisms actually do or learn in real situations. As mentioned earlier, proponents of contemporary EP would have us look to the distant (population-level) past to understand the (individual-level) present. Keeping in mind the deeply problematic issues of determining whether human behaviors were adaptive in the distant past or were influenced by natural selection (Lewontin, 1998), we believe there is value in this approach, namely in its functional orientation. Asking how humans (or any species) “solve” or adapt to various problems (like finding and securing suitable mates) has been a useful way throughout the history of psychology to frame the study of human behavior and development. Our approach, however, differs from EP by studying how organisms adapt in the course of their lifetime and across generations. Humans can certainly detect cheaters in social relationships, but how humans develop this ability, and how it can be altered, must be addressed. To be clear, we are not discounting the value in attempting to determine the evolutionary origin of traits, but we believe these questions are most strategically answered through tests of a range of different populations within and across a range of related species. That is, we favor a focus on how behavioral patterns and cognitive abilities develop, are maintained, and change in existing human and animal populations. Take, for example, a child’s theory of mind, which is believed by some to be an innate mental module (Baron-Cohen, 1995; Scholl & Leslie, 1999). In Western, middleclass (predominately White) children, the theory of mind module (as measured in false belief tasks) appears to become functional on average at about 4 years of age. However, as Boesch (2007) has noted, there are tremendous cultural differences. In some cultures, most children do not pass false belief tasks until age 7, in others age 9, and in still other cultures children continue to fail the task at 14 years of age. Any attempt to discuss the evolution (and development) of a so-called theory of mind module must take these differences into account. However, studying the evolutionary origins of any trait must also involve identifying the distribution of that trait in related species. For humans, this poses a significant challenge because there are very few closely related species living today (Lewontin, 1998). Of course, studying theory of mind in chimpanzees and other primates can help in the reconstruction of this ability, but as with humans, researchers must consider the
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social upbringing of the animals (from wild populations vs. labreared) and assess the skill in a way that is ecologically relevant to the species tested (Boesch, 2007). A major point of departure from our framework and other evolutionary approaches to behavior is our interest in treating evolution as an ongoing process. Our perspective sees behavior as both the product of evolution and also as part of the process by which evolution takes place. Note that the assumptions of the modern synthesis (and in turn, EP) limit how evolutionary concerns could be applied to the study of behavior, as the behavioral (and cognitive) characteristics of individuals are viewed as predetermined outcomes of a past, finished process of evolution (Honeycutt, 2006). The idea that behavioral changes can be an important source of evolutionary change has received increasing research attention in the last decade (see Gottlieb, 2002; Oyama, Griffiths, & Gray, 2001; West-Eberhard, 2003). Acknowledging the role of behavioral change in initiating (or inhibiting) evolutionary change involves the idea that behavioral changes can alter selection pressures by bringing organisms into different environmental relationships. In this sense, behavioral change only indirectly influences evolutionary change via its impact on selection pressures (Duckworth, 2009). For example, imagine a situation in which a group of land-dwelling mammals begin to exploit a new food resource found under water that requires some level of diving and swimming. Indirectly, this change in diet and foraging behavior could influence selection pressures for morphological traits (e.g., webbed feet, increased adipose tissue for thermal regulation, and increased lung capacity) and behavioral traits (swimming skills and underwater perceptual skills) that aid in obtaining this resource. In our developmental evolutionary approach, behavioral changes could also have more direct effects on lifecycles. Identifying these direct effects would involve investigating how developmental conditions of offspring change following their parents’ new behavior. In the hypothetical example above, we would want to know how the nutritional value of the new food might alter the physical development of offspring. Different diets could not only influence physical stature, but the rate/speed of physical growth or onset of puberty. It would most certainly be the case that the flavors and odors of the new food would be present in the amniotic fluid of offspring, which could bias the responsiveness of offspring following birth. If pregnant animals dive and swim for the food, how might the types, timing, and amount of sensory stimulation change for their embryonic offspring? Do the animals begin to nest near the water, and, if so, what might be the developmental implications of these new environments? Does the new foraging behavior (or nesting environment) somehow bring about changes in social structure or alter social interactions (e.g., levels of physical contact, grooming, patterns of parental care, or mating behavior) in the population? How might offspring learn to dive and swim in a manner different than their parents? And importantly, how do all of the aforementioned factors persist or change across each generation? Of course, addressing the questions above requires working with nonhuman species given the practical and ethical limitations of studying humans across generations. However, the questions listed above are relevant when studying how human individuals and populations adjust to environmental changes (e.g., immigrating
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populations). As a case in point, imagine all the developmental changes that might ensue in a fishing community whose primary food and water supply has been polluted by industrial waste. As with the animal studies, we would be interested in any direct effect of toxins (or new diets) on individual development, changes in foraging behavior, changes in child care and education, changes in social structure (cooperation and competition for resources, dominance hierarchies, mating strategies, and/or divisions of labor), as well as the impact that these new behaviors have on the surrounding ecology. Note that the questions and topics addressed by our developmental evolutionary approach involve unpacking developmental dynamics across numerous levels of analysis (epigenetic, behavioral, social, ecological) and time scales (developmental and transgenerational). Note too how reference to selection pressures describes only the statistical changes in population parameters (e.g., the distribution of phenotypic variants), whereas our developmental evolutionary orientation addresses how these variants actually come into being and are perpetuated across generations. A complete understanding of evolution requires both statistical (selectionist) explanations at the population level and dynamical explanations at the individual level (Ariew, 2003). Unfortunately, the former types of explanation have received the majority of attention in both evolutionary biology and evolutionary psychology. For example, changes in behavior brought about by changes in prenatal and early postnatal rearing environments have been well documented in comparative psychology (e.g., Lickliter, 2005; Michel & Moore, 1995; Renner & Rosenzweig, 1987), but this experimental work has received little empirical attention from those interested in evolutionary change. Generally speaking, our framework for psychology is fundamentally process-oriented. Because all topical areas in psychology (e.g., personality or cognition) can be framed in developmental terms, a developmental metatheory is especially well suited to unite the various disciplines in psychology under a common agenda. The developmental evolutionary framework we propose would expand this developmentally oriented metatheory to include transgenerational, cross-cultural and comparative analyses, in addition to asking functional (i.e., fitness-related) questions. In other words, should psychologists adopt our framework, a greater emphasis would be placed on the contextually contingent origins, maintenance, modifiability, and adaptiveness of psychological phenomena (e.g., personality differences, psychological disorders, problem solving, or obedience) in real-life settings across generations.
Conclusion Writing over 70 years ago, the embryologist and geneticist Conrad Waddington (1941) pointed out that A theory of evolution requires, as a fundamental part of it, some theory of development. Evolution is concerned with changes in animals, and it is impossible profitably to discuss changes in a system unless one has some picture of what the system is like. Since every aspect of an animal is a product of development, or rather is a temporary phase of a continuous process of development, a model of the nature of animal organization can only be given in developmental terms. (p. 108)
The current trends in evolutionary biology suggest we are on the verge of a new, broader synthesis of disciplines, one that shares Waddington’s insight. We believe psychology can benefit from and play a key role in its construction in the decades ahead. The metatheoretical assumptions of the developmental evolutionary framework outlined here provide a starting point for guiding how psychology can better align with contemporary life sciences to further advance our understanding of the links between the process of development and the process of evolution.
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Received April 5, 2013 Accepted April 9, 2013 䡲
