Review of General Psychology 2013 American Psychological Association

2013, Vol. 17, No. 2, 184 189 1089-2680/13/$12.00 DOI: 10.1037/a0032932

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,
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epigenetics, and developmental science. The authors outline a developmental evolutionary framework for
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psychology, a relational metatheory that integrates the study of developmental and evolutionary mech-
anisms 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 pro-

cesses, dynamic systems

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 pre-
vailing theory of evolution during the 20th century. Some have
claimed that EP could serve as a metatheoretical basis for psy-
chology (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 & Mller, 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 Uni-
versity; Hunter Honeycutt, Department of Psychology, Bridgewater Col-
lege.
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: licklite@fiu.edu
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 en-
code programs or instructions that predetermine the development
of traits. Moreover, genetic information was not thought to mod-
ifiable 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 morpholog-
ical 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 individuals genes (directly or indi-
rectly via kinship relations) in subsequent generations. EP emerged
in the 1980s, focusing specifically on human behavior. EP modi-

184
 DEVELOPMENTAL AND EVOLUTIONARY PSYCHOLOGY
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 gener-
ate behavior. Specifically, EP argues that there exists a large set of
domain-specific cognitive modules in the minds of modern hu-
mans 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
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behavior and cognition, EP argues that psychologists must look to
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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 & Cos-
mides, 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 devel-
opment, and developmental influences (viewed as proximate
causes) from evolutionary processes. These divisions in turn fos-
tered the view that developmental analyses could add no explan-
atory 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 evolu-
tionary concerns with embryology and developmental biology
(Amundson, 2005; Gottlieb, 1992). This population genetics ap-
proach 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, molec-
ular, cellular, and developmental biology, neuroscience, develop-
mental psychology and psychobiology, and evolutionary biology
to reconsider the nature of the links between development and
evolution (e.g., Gottlieb, 2002; Hallgrmsson & Hall, 2011; Pigli-
ucci & Mller, 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 suf-
ficient explanation for the achievement of any phenotypic out-
185
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 cogni-
tion).
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 devel-
opment 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 stimu-
lation such as maternal behaviors (levels of licking and grooming),
abusive caregiving, and experimenter-provided handling (Cham-
pagne, 2010; Crews, 2008; Harper, 2005; Honeycutt, 2006).
These modern advances challenge the foundational dichotomies
(genetic vs. environmental; innate vs. acquired; ultimate vs. prox-
imate; 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 psy-
chology. 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 devel-
opment of our framework, below we describe several metatheo-
retical assumptions that should contribute to the course of psy-
chologys future:
 186 LICKLITER AND HONEYCUTT
Individual development takes place within a hierarchy of
embedded relational systems (gene celltissue organ system
organismproximate environments distal environments) that re-
ciprocally 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 com-
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peting alternatives. Development is situated, contingent, and
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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 inher-
itance 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 re-
sources. These are reconstructed in each organisms 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, neu-
ral, physiological, behavioral, and social mechanisms act syner-
gistically as a system from which behaviors emerge and are
maintained within and across generations. For example, explana-
tions 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 motiva-
tional 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 devel-
opment actually happens, is fundamental to our proposed frame-
work. As we put it a decade ago:
If evolutionary psychologys aim is to understand the evolved psy-
chological mechanisms that underlie behavior and the selective forces
that have shaped these mechanisms, then it seems to us that uncov-
ering the network of factors (both internal and external to the organ-
ism) 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 evolu-
tionary 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 (John-
ston, 1985). In much of traditional psychological analysis, how-
ever, 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 un-
derstate 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 pop-
ulations (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 sug-
gested 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
 DEVELOPMENTAL AND EVOLUTIONARY PSYCHOLOGY
beyond the proposed sensitive period for learning, but in this more
ecologically realistic situation, could even learn songs of a differ-
ent species (Baptista & Petrinovich, 1984).
Traditionally, in studies of human infant perception research-
ers 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 real-
istic 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 draw-
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ings of figures to study attraction, face-like stimuli to study infant
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face preferences, still shot photographs to study emotional discrim-
ination), 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 organ-
isms 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 develop-
ment. Our approach, however, differs from EP by studying how
organisms adapt in the course of their lifetime and across gener-
ations. Humans can certainly detect cheaters in social relation-
ships, 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 childs theory
of mind, which is believed by some to be an innate mental module
(Baron-Cohen, 1995; Scholl & Leslie, 1999). In Western, middle-
class (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
187
social upbringing of the animals (from wild populations vs. lab-
reared) 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 con-
cerns could be applied to the study of behavior, as the behavioral
(and cognitive) characteristics of individuals are viewed as prede-
termined outcomes of a past, finished process of evolution (Hon-
eycutt, 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 behav-
ioral change in initiating (or inhibiting) evolutionary change in-
volves the idea that behavioral changes can alter selection pres-
sures by bringing organisms into different environmental
relationships. In this sense, behavioral change only indirectly
influences evolutionary change via its impact on selection pres-
sures (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 behav-
ior 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. Identi-
fying these direct effects would involve investigating how devel-
opmental 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 gen-
eration?
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
 188 LICKLITER AND HONEYCUTT
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, dom-
inance hierarchies, mating strategies, and/or divisions of labor), as
well as the impact that these new behaviors have on the surround-
ing ecology.
Note that the questions and topics addressed by our develop-
mental evolutionary approach involve unpacking developmental
dynamics across numerous levels of analysis (epigenetic, behav-
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ioral, social, ecological) and time scales (developmental and trans-
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generational). Note too how reference to selection pressures de-
scribes only the statistical changes in population parameters (e.g.,
the distribution of phenotypic variants), whereas our developmen-
tal evolutionary orientation addresses how these variants actually
come into being and are perpetuated across generations. A com-
plete understanding of evolution requires both statistical (selec-
tionist) explanations at the population level and dynamical expla-
nations 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 docu-
mented 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 funda-
mentally 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 ad-
dition to asking functional (i.e., fitness-related) questions. In other
words, should psychologists adopt our framework, a greater em-
phasis 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 gener-
ations.
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 ani-
mals, 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
Waddingtons 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