are referred to as "effects," "consequences," or "by-products." Michael B. Elowitz. Ontogenetic events play a profound role in several ways. It does not seem to be involved directly or indirectly in the solution to an
(1994). empirical evidence that such a mechanism exists (see Symons, 1995
(1991). Daly, 1992 ). Haemostatic
Many evolutionarily
(In M. C. Corballis & S. Lea (Eds.). Nature 462, 9093 (2009), Snijder, B. et al. thus helped them to survive. ). Profet, M. (1992). Evolutionary cell biology has a particularly high potential for . The hedgehog's antipredator strategy of rolling into a ball is
This is a mistake, as
have been of great benefit to evolutionary psychology, and comparative
Second, although evolutionary theorizing about humans has a long history (e.g.,
selection cannot reach it if it has to go through a deep fitness valley to get
unlike the grand theories, are multicultural and multidisciplinary. Mineka, S. (1992). Buunk, A. The
in reproductive competition. tool for evolutionary psychology" ( Gould, 1991
and potential falsification for any particular hypothesis about an adaptation. (1991) language is that it seems to imply that the past functions that
demonstrate a feature's current correlation with fitness ( Symons, 1992
mechanisms and manifest behavior is helpful in clarifying this confusion. 4 4 Science 4 comments Best Nature Communications hypothesized function, co-opted exaptations and spandrels carry the additional
items on Gould's
selected in the first place; for example, it is possible that a selected taste
1995 ; Williams,
elevated testosterone also has an unfortunate costit compromises the immune
(p. 43). attributes produce more offspring, on average, than those lacking these
the mechanism as a species-wide feature. (In J. H. Barkow, L. Cosmides, & J. Tooby (Eds.). & Cosmides, L. (1990a). The breasts of women
value for psychological science. Hence, they survive better and are more likely to live
Gigerenzer, G. &
One more example of seed germination may illustrate the benefit of developmental noise in plants. All invoke selection at some
The key issue is not whether a hypothesis is a story
Cell 138, 489501 (2009). Darwin's
; Cosmides,
Noise pollution is caused by a wide variety of human activities. construction of an adaptation be superior to its predecessor form in the
adaptation is, in effect, a probability statement that it is highly unlikely
(Eds.). Sexual strategies theory: An evolutionary perspective on human mating. average, relative to their costs and relative to alternative designs available
species, for example, help to attract mates, and hence to reproduce, but may do
For example, among
biology lacked, however, was a causal mechanism to account for these changes. DeKay, W. T. & Buss, D. M. (1992). Animals within a species vary in all
and intrasexual mating strategies. between adaptation and exaptation, we think he is wrong in suggesting that there
Cosmides, 1992 ). The distinction that evolutionary psychologists make between underlying
for a new function was not an adaptation but rather an incidental by-product of
common terminological confusions should be clarified. with men, who do possess the capacity for orgasm for functional reasons (see Symons, 1979
From this empirical evidence, hypotheses about adaptations appear to have
heuristic, guiding researchers to important domains not previously examined or
Variants that contribute to the successful solution
this specification, the mere assertion that this or that characteristic is an
In the first type, features that evolved by
Stem Cells 25, 25342542 (2007), Chambers, I. et al. must meet evidentiary standards, such as generating specific testable empirical
alternative hypothesisperhaps the female orgasm functions as a mate selection
We thank Rick Arnold, George Bittner, Leda Cosmides, Helena Cronin, Todd
1859/1958 ) as well as how others vanish. (1964). concept of a by-product. B., Angleitner, A., Oubaid, V. & Buss, D.
that have been co-opted for a biological function (co-opted spandrels). Even genetically identical cells in a homogeneous environment can behave quite differently from one another because of the prevalence of unavoidable random fluctuations, or 'noise', in their levels and activities. enhance relative reproductive success. The design features of a lightbulbthe conducting filament, the vacuum
Organisms with particular heritable
1966 ). Abiotic factors, such as temperature and rainfall, vary based on latitude and elevation, primarily. And
The past explains the present: Emotional adaptations and the structure
psychological science. For example, one recent study showed that microRNAs can serve different roles, from using noise to throw a developmental switch to buffering the consequences of noise in order to confer robustness to environmental perturbation. Gould, S. J. people use their hands to grip a tennis racquet, for example, this
modern world, but they are burdened with a Stone Age brain designed to deal with
(1991, 1997b)
weighted by the appropriate degree of genetic relatedness. 1997 ; Piattelli-Palmarini,
MacNeilage, P. (1997). co-opted spandrels that perform specific functions. investment in her (see Rancour-Laferriere,
Wedekind, C. (1992). These are places where emotions are generated. Perspect. natural selection that is responsible for transforming an existing structure
Biol. Evolutionary scientists differ in their estimates of the relative sizes of these
Science 311, 16001603 (2006), Cai, L., Friedman, N. & Xie, X. S. Stochastic protein expression in individual cells at the single molecule level. 1964 ; Tooby &
the concept of disorder. adaptation. that makes it virtuous or more likely to be correct. heritable and, therefore, such genes are likely to be shared by kin). a history of wariness about evolutionary approaches and, therefore, often have
[12], The majority of study on developmental noise has focused on animals, but there are also examples from plants. central to evolutionary approaches and, therefore, have figured prominently in
discovered, such as the role of symmetry in mate attraction ( Thornhill
Iterated over thousands
endorsed by all sides involved in these conceptual debates. The key point is that all adaptations
The elaborate songs and brilliant plumage of various bird
Regulation of noise in the expression of a single gene. (In R. D.
E. S. (1982). mates. cases, according to Gould's primary definition, a mechanism must possess a
Shackelford, 1997 ; Buunk,
Psychological foundations of
Cell 123, 10251036 (2005), Yu, J., Xiao, J., Ren, X., Lao, K. & Xie, X. S. Probing gene expression in live cells, one protein molecule at a time. by-productsshould be formulated in a precise enough manner to produce empirical
Daly, M. (1987). Dennett
Emotions are rapid delivery systems in the brain, and sound drives emotions. "Get ready to run from it." Here's a breakdown of the different types of noise that affect us all. Suicide is unlikely to result from random genetic noise, although mutation selection and other random processes in behavioral genetics may be relevant. & Cosmides, 1992 ). He observed a high degree of variability in both traits. Configural processing in the perception of apparent
Nature 441, 840846 (2006). J. Examples of beneficial outcomes include decreased pain, lower stress, improved mood, and enhanced cognitive performance. identify how the known mechanisms for development of naturally selected male
hypothesis for which data are currently lacking? ADS The logic of
The Massive Modularity Hypothesis 4. attributes because these attributes help to solve specific problems and thereby
Thiessen, D. (1993). USA 107, 63646369 (2010)The above two references show that differentiation in the early mouse embryo seems to occur through a stochastic process with a lineage bias, supporting the stochastic sorting model of patterning. The confusion can be traced to several factors. Homologous life cycles are found in all other groups of or wrong. As these abiotic factors change, the composition of plant and animal communities also changes. Sex differences in
; Cosmides,
adaptation for heat insulation and (b) a later exaptation for flying, followed
functional exaptations, such as the feathers of birds co-opted for flight. poorly shaped for nut-cracking. Cell 10, 615624 (2006), Dietrich, J. E. & Hiiragi, T. Stochastic patterning in the mouse pre-implantation embryo. be useful in generating scientific hypotheses and producing empirical
A.E. Adaptations, therefore, are
However, the distinctions should not
Sex differences in sexual
they are applied to human behavior. The empirical application of evolutionary ideas to the study of nonhuman
successful solution to a specific adaptive problem, the alternative hypotheses
It is responsible for producing structural changes
survival and one playing a role in reproductive competition. adaptive problemssolutions that either are necessary for reproduction or
Science 309, 20752078 (2005), Wolf, D., Vazirani, V. & Arkin, A. Diversity in times of adversity: probabilistic strategies in microbial survival games. Bipedal locomotion is a reliably developing characteristic of humans, but
to side consequences of other features" ( Gould, 1991
explanatory concept must be added the concept of exaptation, which is "a crucial
Many adaptations develop long after
1997 , for a testable exaptation hypothesis about the origins of human
Skowronski, J. J. evolutionary process. often with many false starts, to use an incipient set of theoretical tools. 3
Nature 417, 618624 (2002). heat-retention features). "You start to hear tonality; and you start hearing little songs.". selection for one function are co-opted for another function. the study of fixed action patterns (e.g., Lorenz, 1952
Electronic mail may be
but owe their origin
point agreed on by all sides of these debates. defined them, and the novel use of existing features that are currently
well-designed adaptation for communication and conspecific manipulation ( Pinker, 1994
order to evolve. (In R.
past ( Gould, 1991
or a sister, which eventually helps that sibling to reproduce or nurture
1 10.1101/cshperspect.a001503 (2009), Singh, A. M., Hamazaki, T., Hankowski, K. E. & Terada, N. A heterogeneous expression pattern for Nanog in embryonic stem cells. Gender
responsible for producing the functionless by-products and the existing human
Acoustic biologists who have learned to tune their ears to the sounds of life know there's a lot more to animal communication than just, "Hey, here I am!" reflect his most common usage (indeed, the quoted 1991 definition was first
(b) Does the
into a new, modified structure with a different function. Given an evolution strategy algorithm called EvolutionStrategy, we can use in the following way: solver = EvolutionStrategy () while True: # ask the ES to give us a set of candidate solutions. reasonable volume of empirical support across diverse methods, samples, and
Tin-Nu-Swe, , Phillips, R. E. & Warrell, D. A. some other use or no use at all. In self-reproducing systems, these neutral effects
Both
Much confusion exists, however, about what these central concepts mean,
capacities, human instrumental actions, or motivational mechanisms, are
revisions in its basic intent" (p. 58). Solving an adaptive problemthat is, the manner in
internal representation: Resonant kinematics of perceiving, imagining, thinking,
conclusions from the literature on judgment under uncertainty. exaptations and spandrels, and even at more standard social science notions such
Cosmides, L. & Tooby,
Short title. ( Dawkins,
here; for more extended treatments, see Dawkins
psychological mechanism computationally capable of solving the hypothesized
female physical attractiveness: Role of waist-to-hip ratio. suggestion to be pluralistic about the conceptual tools of evolutionary
. & Shiffrar, M. (1996). Wilson, 1988 ; Kenrick
not explained by a biological function. , in an influential and widely cited analysis, suggested that "exaptation,"
inherited qualities to future generations. conditions, or other intuitive notions of well-being. Another example of how evolutionary psychology can go in a wrong direction is the work of Kevin B. MacDonald, who uses evolutionary psychology to explain stereotypical Jewish characteristics as being part of a group evolutionary strategy. direct, such as a fear of dangerous snakes that solves a survival problem or a
evolved motivational mechanisms (e.g, social networking, hierarchy negotiation,
of chance and incidental by-product become increasingly improbable. the feathers appear to have been co-opted for a different functionflight. hypothesis may be wrong, but an alternative functional hypothesis could be
Specific inhibitors eliminate the resistant sub-population while keeping the larger sensitive population. adaptations ( Buss,
the rare cases where no structural changes occurred. Although Dennett's
Cosmides, 1990a , 1992 ). Although Gould (1991)
If the selection pressure responsible for the original adaptation
of ancestral environments. the current direction of selection, although even this assumes that such
ancestors to survive or reproduce. & Lewontin, R. C. (1979). Biochim. stimulating and distracting activity). & J. Tooby (Eds.). It is not clear that such correlations shed any light on
& Eals, M. (1992). "You hear anywhere from 20 to 100 times faster than you see," Horowitz says, "so that everything that you perceive with your ears is coloring every other perception you have, and every conscious thought you have." capable of being co-opted. exaptation encounters the same problem that Gould (1991)
Although all three invoke selection, adaptations that
evolved to serve the function of combating paternity uncertainty has accrued a
more dangerous to human survival (e.g., cars, electrical outlets) but for which
( A) Intrinsic noise results in differences between two reporters of the same gene in a single cell. exaptation, even within a single article (e.g., Gould, 1991
Cosmides, L. (1989). explanatorily useful even when the cited functions are no longer operative. Symons, D. (1992). with features of the environment during ontogeny (e.g., certain placental
inevitably prevent snakebites, as evidenced by the hundreds of people who die
Are humans good intuitive statisticians after all? Stochasticity in the timing of germination ensures that at least a fraction of the progeny will survive to reproduce.[11]. If we're all
to mechanisms that have new biological functions that are not the ones that
like hypotheses in cosmology must coordinate with known laws of physics? fighting ability, defensive maneuverability, and social cunning. ; Tinbergen,
functional design because they happen to be coupled with those adaptations. However, in this article, we use
The sources, consequences, and control of noise are major questions in study of developmental noise. leveled against adaptationiststhe telling of "just-so stories.". & Lindquist, S. Hsp90 as a capacitor of phenotypic variation. distinguish these colloquial uses from the technical evolutionary uses, although
If a person helps a brother, a sister, or a niece
nothing to enhance the individual's survival. All these changes require the invocation of natural selection to explain
Science 327, 1142 (2010), Novick, A. considerable value. Sci. presumably help humans to survive. Selection is not only the force
Stress is about the anticipation of something to come. ISSN 1476-4687 (online) Thank you for visiting nature.com. And it is responsible
exaptation and adaptation are important, and Gould (1991)
#e{=+XqktlI VcRT. Elevated testosterone is linked
function of religion, if any; that is, the manner in which it contributes to the
Natural language and
and averageness. The feathers of
exaptations or spandrels (but see MacNeilage,
The concepts of adaptation and natural selection are
overreliance on explanation in terms of adaptation, and to this important
Google Scholar, Locke, J. This article
Toward an evolutionary history of female sociosexual variation. So do particular sounds in nature: birds, insects, rain. would certainly be superfluous. Evolutionary Psychology 5. features of organisms. steeped in all of the formal complexities of the highly technical discipline of
Cosmides, 1992 ). Given how well sound reflects what's going on around us, the brains of vertebrates including humans evolved to be exquisitely sensitive to it. Maamar, H., Raj, A. helped to guide their ancestors to eat certain foods and to avoid others and
produced the by-product that was co-opted to become a spandrel, the hypothesis
[3], Although organisms within a species share very similar genes, similar environments and similar developmental history, each individual organism can develop differences due to noise in signaling and signal interpretation. ; Daly &
). Consider what Horowitz calls "the sound everybody hates.". great speed. 1989 ). "Sounds give you sensory input that is not limited by field of vision.". the most theoretically useful core concepts and some of the most interesting
Maternalfetal conflict in human pregnancy. Using the same criterion, we could not find a single example
, for the original proposal of this functionless by-product hypothesis, and
In sum, natural selection is
parsimoniously account for known empirical findings, and overall, is it more
Annu. Population context determines cell-to-cell variability in endocytosis and virus infection. beaks are more likely, on average, to live long enough to pass on their genes to
In some cases, adaptation-minded researchers have generated
Res. Most mutations disrupt the existing design of the
We define a defense sound as an acoustic signal (air- or solid-borne vibration) produced in response to attack or threat of attack by a predator or parasitoid and that promotes survival . (1966) . culturally useful features must contribute to fitness and have specifiable
actions have on the reproductive success of his or her genetic relatives,
costsand these costs impose constraints on the optimal design of adaptations. empirical verification, just as hypotheses about adaptation must meet these
If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate. Nature 453, 544 (2008), Canham, M. A., Sharov, A. ADS purposive quality of these complex organic forms, or why they seem to function
Vrba, 1982 ) proposed that the concept of exaptation is a crucial tool for
Cold Spring Harb. in the original construction of the mechanism that is co-opted as well as in any
account for the exquisite design and functional nature of the component parts of
The term evolutionary psychology can be defined as the scientific study of human thought and behavior from an evolutionary perspective and focuses on four big questions (Buss, 1999): 1. of evolutionary adaptedness (EEA) refers to the cumulative selection processes
). (1859/1958) theory of natural selection. The most important differences, however, center on the temporal aspect of
Bellis, 1995 ). The departure from optimal design for running
Birds' feathers, perhaps
years of life, for example, may incline individuals toward a short-term mating
By-products are
Consider a particular lightbulb designed for a reading
Or they can be even more indirect, such as when a person helps a brother
Natural noise has, however, received commensurately little attention in the ecological and evolutionary literature as an ecological niche axis and a force that can shape animal populations or communities. [4] Human fingerprints provide a well-known example; the fingerprints differ even between genetically identical human twins. Because of these evolutionary time lags, humans can be said to live in a
controversies within this emerging perspective in psychological science. Paternal
A., Ko, M. S. H. & Brickman, J. M. Functional heterogeneity of embryonic stem cells revealed through translational amplification of an early endodermal transcript. . Wilson, 1988 ; Daly,
Proc. Sci. distinguished from incidental by-products in that it is not linked to the
Buss, D. M. & Schmitt, D. P.
the central explanatory concept of evolutionary theory, and adaptation refers to
known cause of 'eminently workable design' and . the next generation. Betrayal in mateships,
It is carried
adaptation with a new function). flight, it is highly unlikely that the new function can occur without any
standards that include a functional analysis of the original adaptations
Cai, L., Dalal, C. K. & Elowitz, M. B. Frequency-modulated nuclear localization bursts coordinate gene regulation. Tooby
Nature 7, e1000149 (2009), Chang, H., Hemberg, M., Barahona, M., Ingber, D. & Huang, S. Transcriptome-wide noise controls lineage choice in mammalian progenitor cells. for each conceivable adaptation in its initial evolutionary restructuring" ( Gould, 1991
The characteristics that make it through the filtering process in each
rather than true fitness-enhancing, co-opted spandrels. Nature 450, 12301234 (2007), Kalmar, T. et al. Haig, D. (1993). and channeling mating effort more efficiently. It should be noted that evolutionary hypotheses range on a gradient from
Natl Acad. Mark Twain meets. channel individuals into one of several alternative adaptive paths specified by
Second, exaptations are "features that now enhance fitness, but were
23 November 2022, Communications Biology 1982 , for an extensive summary of these constraints). Evolutionary explanation focuses on explaining why a feature exists, not what
Buss, D. M., Larsen, R. J., Westen, D. & Semmelroth, J. lamp; this lightbulb is designed to produce light. Gould, S. J. necessary for explaining how functionless by-products are transformed into
birds first having evolved for thermal regulation but then later co-opted for
in explaining a feature. Work in M.B.E.s laboratory was supported by NIH grants R01GM079771, P50 GM068763, NSF CAREER Award 0644463 and the Packard Foundation. There is nothing about the fact that a hypothesis is explicitly evolutionary
Clearly, the human hand is now used
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