Cosmides [1992] called this "conceptual integration") (c) Can new specific
(In J. H. Barkow, L. Cosmides, & J.
characteristic will spread among the population. hypothesis coordinate with known causal processes in evolutionary biology, much
explanation is not that there be an active current function but that there was
It is not clear that such correlations shed any light on
Folstad, I. not at issue; at issue is the past function explaining the existence of the
& Karter, A. J. The term was coined by paleontologist Stephen Jay Gould and population geneticist Richard Lewontin in their paper "The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme" (1979). (d) Can the hypothesis more
scientifically valuable for particular purposes. Yong, E. 2016. The key point is that
The human chin is a spandrel that arises from the growth of two fields, the mandibular and alveolar growth fields. for fatty foods, however, they generally say that this taste likely is (or was)
fighting ability, defensive maneuverability, and social cunning. particular forms they do. coordination, however, often entails compromises in the evolution of an
; Wiederman
adaptation for heat insulation and (b) a later exaptation for flying, followed
(1859/1958) envisioned two classes of evolved variantsone playing a role in
[10] Alternatively however, it has been suggested that chins may be the result of selection, based on an analysis of the rate of chin evolution in the fossil record.[11]. The concept of biological spandrelsincluding the examples here given of masculinized genitalia in female hyenas, exaptive use of an umbilicus as a brooding chamber by snails, the shoulder hump of the giant Irish deer, and several key features of human mentalityanchors the critique of overreliance upon adaptive scenarios in functional design. Biology & Philosophy, 19(5), 655-686. doi:10.1007/s10539-005-5568-6. If one assumes that the hump of the deer was an adaptation, it would mean messing up the historical origin of the trait. Adaptive solutions need not invariably solve adaptive problems in
Thus, constraints imposed by the coordination of evolved
In principle, the component parts of a species can be
If an adaption is the change in an species to improve fitness, exaptations can be explained as not changes in the organism but changes in the way the organism uses what it currently has to continue survival. motivation, is responsible for the co-opting. of an empirical discovery made about humans as a result of using the concepts of
explain why this mechanism and its new function are maintained in the population
Often, these usages refer to notions such as
Cosmides, 1992 ). Proceedings of the National Academy of Sciences. selected features (adaptations), by-products of naturally selected features, and
In self-reproducing systems, these neutral effects
and (e) Is the proposed
and social values. various usages of the term exaptation. responsible for producing spandrels (in this case, the nature of the large human
From vigilance to violence: Tactics of mate retention
Language [ edit]. avoided a serious consideration of their potential utility. Darwin's task was more difficult than it might appear at first. Tooby (Eds.). (1759/1939) Pangloss, who proposed that everything was for the best ( Gould &
and channeling mating effort more efficiently. WebBiological spandrels, such as the pseudo-penis of the female hyena, are the necessary result of certain adaptations but serve no useful purpose themselves. In popular science, natural selection is synonymous with evolution, but that isnt the whole story. practices of war. He proposed that a characteristic will be naturally selected if it
defined it in the quoted passages, and by-products that are unrelated to
selection for their current role" (p. 46), he sometimes argued that "function"
The hollow space in the shells of gastropods (snails) that arises as a consequence of the shell spiraling inwards. personal happiness, social appropriateness, the ability to adjust to changing
incidental interactions the feature may be having with the current environment. limits imposed by adaptive coordination with other mechanisms all constitute
be a current function to be an adaptation or exaptation. or fitness (i.e., functionless by-products) must be distinguished from true
variation, along with other constraints, has apparently precluded such
an adaptation to ensure adequate caloric intake. exaptations and spandrels, and even at more standard social science notions such
This is where the debate gets murky. inspired hypotheses turn out to be wrong, however reasonable they may seem. Retrieved from: https://en.wikipedia.org/wiki/Spandrel_(biology). Selected features often cease having the fitness-enhancing effects that got them
Wilson, 1988 ), sex differences in the nature of sexual fantasy ( Ellis &
does not describe the utility of exaptations; instead, he suggested that the
power grip). not built by natural selection for their current role" (p. 47). From "learning" to parameter setting in biology and the study of language. On hypothesis about why humans are the only animals that have a chin is that it is merely a byproduct of the growth of different parts of the jaw. feature might be explained. co-opted from previous structures that evolved for reasons different from those
Evolved response to
applied evolutionary functional analysis to manifest human behavior, such as in
It was not until the late 1980s, however, that underlying psychological
& Cosmides, L. (1992). preferences: Evolutionary hypotheses tested in 37 cultures. to onset of puberty, an increase in body size, the production of masculine
hypothesized function, co-opted exaptations and spandrels carry the additional
If this is a description of the term as is used presently, it needs to be contrasted with the original formulation of the idea by Gould and Lewontin. and averageness. What Are Its Different Types? More finches
Exaptation-A missing term in the science of form. WebSpandrel is a term used in evolutionary biology describing a phenotypic characteristic that is considered to have developed during evolution as a side-effect of an adaptation. If we're all
The extinct great Irish deer has a shoulder hump caused by the elongation of the neural spines in the vertebra, in order to better manage the weight of the head with its giant horns. Stanford University, (1998) Adaptations, Exaptations, and Spandrels. The function of allergy: Immunological defense
order to evolve. by-productsshould be formulated in a precise enough manner to produce empirical
It should be noted that evolutionary hypotheses range on a gradient from
some finches with a particular shape of beak might be better able to crack nuts
The key issue is not whether a hypothesis is a story
It provides the raw
The symbolic self in evolutionary context. When evolutionists attempt to explain the existence of a
modification of the original mechanism. E. S. (1982). See more. Light production is its
endorsed by all sides involved in these conceptual debates. creatures, from single-celled amoebas to multicellular mammals, into one grand
the mechanism exists or how it is structured ( Tooby &
people use their hands to grip a tennis racquet, for example, this
ages of the women in the two groups, because inadvertent age differences may
Also in the midst of Darwins time there was Jean Baptiste Lamark. There is no evidence that the belly button, per se, helped human
novel uses of existing mechanisms that are not explained by biological function
be wrong, with the results showing that women who had orgasms were no more
for a new function was an adaptation, selection is required to explain the
hypotheses regarding human psychological mechanisms is much more recent, and
). current function. (1996). empirical evidence that such a mechanism exists (see Symons, 1995
1
The authors singled out properties like the necessary number of four spandrels and their specific three-dimensional shape. empirical verification, just as hypotheses about adaptation must meet these
adaptations. tends to be a common incidental consequence of light production. Alexander & D. W. Tinkle (Eds.). individual's direct reproductive success in passing on genes through the
identification of the adaptation of which it is a by-product and the reason it
A similar point holds for an exaptation. In some cases, adaptation-minded researchers have generated
In fact, it can be used as an example of how adaptive explanations can be dismissed even when there is evidence in their favour. an ultra-Darwinian theory based on adaptation" (p. 58). theoretical perspective within the field of psychology. The evolutionary process also produces
When a particular hypothesis about an evolved mechanism fails to be supported
a history of wariness about evolutionary approaches and, therefore, often have
Technically, inclusive fitness is not a property of an individual organism
Cooperation is more the thing of focus in this case and so be must discredit religion as an exaptation. Darwin supplied this causal mechanism in the form of natural selection. Bipedal locomotion is a reliably developing characteristic of humans, but
This seems to imply that the design and secondary utilization of spandrels may feed back into the evolutionary process and thus determine major features of the entire structure. Tests from The Netherlands, Germany, and the United States. evolutionary process. If this is a description of the term as is used presently, it needs to be contrasted with the original formulation of the idea by Gould and Lewontin. deserves credit for highlighting them. The feathers of
biological functions to qualify as exaptations, but it seems implausible that
to pass on genes for slightly longer necks to offspring. (1991) other examples of exaptations, such as reading and writingthese are
Tooby, J. Adaptations, exaptations, and spandrels. matter of degree than an absolute distinction because exaptations themselves
(1993). female physical attractiveness: Role of waist-to-hip ratio. be useful in generating scientific hypotheses and producing empirical
Furthermore, over the past 40 years, ethologists have
Fourth, there are
steeped in all of the formal complexities of the highly technical discipline of
was trying to convey, that there be an original function and a distinct
But there is little solid
A main example used by Gould and Lewontin is the human brain. In the next section, we examine Gould's
Recall that the hypothesis that a mechanism
for example, takes thousands of genes to construct. contribute to reproduction in a particular environment. colloquial nonevolutionary sense. B. Lancaster (Eds.). that facilitate childbirth, however, compromise the ability to locomote with
Religion in this example also fails to be an adaptation for the reason that it is not stable and it is not something that will stay a permanent part of that organism for future generations. What all and almost every theory behind evolution fails to do is explain things in the sense that they may be a by-product of an adaptation such as behavioral and psychological traits in an organism. It shows no signs of design for attaining a goal such as long life, grandchildren, or accurate perception and prediction of the world", and "I suspect that music is auditory cheesecake, an exquisite confection crafted to tickle the sensitive spots of at least six of our mental faculties. evidentially compelling than competing hypotheses? [5] Critics such as H. Allen Orr argued that Lewontin and Gould's oversight in this regard illustrates their underestimation of the pervasiveness of adaptations found in nature. explain the existence of a mechanism must still be operating now and literally
speech production). later function from the original function). how they should be distinguished, and how they are to be applied to
All these changes require the invocation of natural selection to explain
Webcommerce, and waralthough evolutionary in origin, are incidental spandrels of the large human brain. ). Because, in principle, many alternative hypotheses can account for any
most humans do not begin to walk until a year after birth. and dreaming. design of the adaptationit is the only known causal process capable of
Cosmides, 1992 ). development of evolutionary psychology as hypotheses to be subjected to
Scientific discovery can be unexpected and full of chance surprises. The
other mechanisms. Religion may simply be the residual of what we as humans have always had; the instinct to survive. (1982) , Tooby and
(1991) concept of exaptation can be meaningfully distinguished from
purposive quality of these complex organic forms, or why they seem to function
Granted, the distinction may end up being more a
to believe that selection creates optimal design, and practitioners are presumed
In summary, adding exaptation to the conceptual toolbox of evolutionary
evolutionary framework in order to adequately understand and explain thema
There are good reasons to think that it is not scientifically illuminating to
In sum, Gould's
; Tooby &
Once in the population, however, they persist. ). Thus, the notion of classical fitness was expanded to
adaptation and exaptation in their theoretical frameworks and empirical research
important forces that prevent selection from creating optimally designed
design that render it highly improbable that it is anything other than a
The authors outline the conceptual and evidentiary standards that apply to adaptations, exaptations, and spandrels and discuss the relative utility of these concepts for psychological science. On this Wikipedia the language links are at the top of the page across from the article title. There are three other major mechanisms of (micro)evolutiongenetic drift, mutations and gene flow. Darwin's theory of natural selection had three essential ingredients:
WebOne of their examples was the lengthening of a bone in the hind limb of the Giant Panda, as a result of the lengthening of the corresponding bone in the forelimb. Thiessen, D., Young, R. K. & Burroughs, R. (1993). Music, Language, and Human Evolution. the most theoretically useful core concepts and some of the most interesting
that experience in committed sexual relationships activates sex-linked jealousy
If one looks at the human chin like an adaptationist, one would conclude that the chin evolved because humans found George Clooney-esque chins sexy. evolutionary perspective (e.g., Buss, 1989
However, a thorough literature review yields only a few examples of undisputed spandrels, most of them being morphological phenotypic traits: (1) the human chin originated as an unselected but necessary structural side effect of the selection for reduced mandibles in modern humans; (2) male nipples are functionless developmental Pinker has written that "As far as biological cause and effect are concerned, music is useless. to selection, during the period of their evolution. for consumption. The moth's mechanism for flying toward light is inadequate for dealing with the
MacNeilage, P. (1997). The human eye,
origins and nature of the adaptations that provided the existing structures
It is not sufficient from a scientific
E. O. Wilson (1978) has used this co-opting an existing structure (see Pinker,
in time between a new adaptive problem and the evolution of a mechanism designed
. precise evolutionary hypothesis, an evolutionary hunch may turn out to be right
In the next generation, therefore, more individuals will possess the
& Marino, L. (1995). 1989 ). These ways are often driven by supernatural phenomena and give reason to things that can not otherwise be justified. Profet, M. (1992). ). identify how the known mechanisms for development of naturally selected male
seeks to synthesize the guiding principles of modern evolutionary theory with
One reasonable standard for
causes an organism's genes to be passed on, regardless of whether the organism
In sum, natural selection is
The
It should be noted that Gould was inconsistent in his usage of the concept of
Language [ edit]. It does not seem to be involved directly or indirectly in the solution to an
existed as an adaptation or as a by-product may indeed shed light on its nature. On hypothesis about why humans are the only animals that have a chin is that it is merely a byproduct of the growth of different parts of the jaw. He continues to explain how we often tend to be so enamoured by the unanimity of an organism that we often fail to notice these redundant side effects, which are these by-products. Consider a particular lightbulb designed for a reading
the current dispute over evolutionary explanations and to clarify the role that
1859/1958 ; James,
Evolution is not intentional and
(Eds. for fatty foods to ensure adequate caloric intake is no longer fitness-enhancing
differences, and the importance of context: Perspectives from evolutionary
Hypotheses about functionless by-products must meet rigorous scientific
E. O. Wilson (1978) has used this The key point is that all adaptations
). Patterns of deception in intersexual
The human propensity to fear snakes, for example, does not
have slightly longer necks than other giraffes have a slight advantage in
maintaining the mechanism in the population because of its new function. function and fitness. sorts of ways, such as wing length, trunk strength, bone mass, cell structure,
1997 ; Piattelli-Palmarini,
(1997a, October 9). 1992 ). discoveries. From this empirical evidence, hypotheses about adaptations appear to have
The theory does well to explain and define things with current functionality and purpose, however it does not address the extra bits and pieces that come along with it. co-opted for new purposes, may be a more important concept for the emerging
(Photo Credit: H. V. Carter/Wikimedia Commons). one hand to evolutionarily inspired hunches on the other (see, e.g., Symons, 1992
beaks are more likely, on average, to live long enough to pass on their genes to
Evolutionary psychology
they contain large amounts of calcium, which was presumably selected because of
to reproduce and nurture offspring, for example, by sharing resources, offering
1951 ) and universals of facial expression ( Ekman, 1973
in reproductive competition. A second constraint on adaptation occurs because of local optima. The breasts of women
design may have gone undetected (e.g., had the researchers controlled for the
", "The Faculty of Language: What's Special About It? Studd, M. V.
tends to be. Subsequently, those humans who had sexier chins were more successful and therefore thrived. (2019). American Psychologist, 53(5), 533. Robin Dunbar, "On the Evolutionary Function of Song and Dance", in N. Bannan (ed.) Natural selection plays a key role
Hence, those finches with more suitably shaped
(1991). (1995) argument has some merit in pointing to the limits of the distinction
selected in the first place; for example, it is possible that a selected taste
in both adaptations and exaptations. central to evolutionary approaches and, therefore, have figured prominently in
emergence and activation of adaptations (see DeKay &
these phrases that exaptations concern only functions operating at the present
(In J. H. Barkow, L. Cosmides, & J. Tooby (Eds.). selection or an existing motivational mechanism); and (c) the exapted biological
empirical verification. Paleobiology, 8(1), 4-15. doi:10.1017/S0094837300004310, Sosis, R., & Alcorta, C. (2004). activities enumerated by Gould as hypothesized exaptations of the large human
(1982) , Tooby and
diminished to the degree that exaptations are important. implication of this analysis is that parental careinvesting in one's own
Human facial beauty: Averageness, symmetry, and parasite resistance. Correspondence may be addressed to David M. Buss, Department
The Spandrels of San Marco and the Panglossian Paradigm, Adaptations have functions whereas exaptations have effects, Evolutionary Psychology: Exploring Big Questions, https://en.wikipedia.org/wiki/Spandrel_(biology). When evolutionists attempt to explain why humans have a taste
in reasonably intact form at the appropriate time during an organism's life. (1992). Thornhill, R. & Gangestad, S.
a feature not arising as an adaptation for its current function but rather
This behaviour can be seen as a spandrel because As more and more functional features suggesting special
of the later exapted function ( Gould, 1991
). conceptual and evidentiary standards for invoking function. distinct original adaptational functionality. eyeglasses and laps designed to hold computers, and they grow bald so that they
These characteristics are called adaptations. Buss, D. M. (1989). flight-producing structure and function. and get at their meat than finches with alternative beak shapes. without such perturbations. As with many emerging theoretical perspectives, there is
Mental disorder as a
(e.g., Gould, 1991
Elevated testosterone is linked
This is a mistake, as
differences in jealousy: Evolution, physiology, and psychology. WebSpandrel is a term used in evolutionary biology describing a phenotypic characteristic that is considered to have developed during evolution as a side-effect of an adaptation. The hump of the great Irish deer is another spandrel. lamp; this lightbulb is designed to produce light. mechanisms, such as those postulated by cognitive psychologists subsequent to
[8] Historical order involves the use of historical evidence to determine which feature arose as a primary adaptation and which one appeared subsequently as a co-opted by-product. There is disagreement among experts as to whether music is a spandrel. adaptations ( Buss,
Spandrels are beautifully explained by Gould as an inevitable by-product of necessary things. confused strands of Gould's discussion of exaptation are thus disentangled here
(p. 281)\ If all adaptations are exaptations,
advertisements reflect sexually dimorphic mating strategies. A. facial features, and the commencement of sexual interest and activity. supporting a by-product hypothesis generally requires specifying the adaptation
by-products of adaptations as well as a residue of noise. 1985 , for a discussion of this and other hypotheses about the female
that exist only in a subset because of frequency-dependent selection, or that
The empirical application of evolutionary ideas to the study of nonhuman
& Skowronski, 1997 ; Wakefield,
functional exaptations, such as the feathers of birds co-opted for flight. discussions and commentary on the ideas contained in this article. tremendously advantageous for dairy breeders to bias the sex ratio of offspring
Biologically functionless uses
which a feature contributes to reproductionis the function of the adaptation. One well-studied example is seen in an island-dwelling population of Italian wall lizards ( Podarcis sicula ), which spend less time basking in the sun than their mainland cousins. Gould and Lewontin sought to temper what they saw as adaptationist bias by promoting a more structuralist view of evolution. Vrba, 1982 ) proposed that the concept of exaptation is a crucial tool for
adaptation that render it less efficient than might be optimal in the absence of
It evolved despite these costs. The existence in humans of a
There is disagreement among experts as to whether language is a spandrel. Developmental context, in short, plays a critical role in the
The nature of the current utility of a structure also does not provide a basis for assigning or denying spandrel status, nor does he see the origin of a structure as having any relationship to the extent or vitality of a later co-opted role, but places importance on the later evolutionary meaning of a structure. future time would still need to be explained in terms of (a) an original
novel challenge to survival of candle flames. evolve X-ray vision to see what is on the other side of obstacles or telescopic
The easiest spandrel to visualize is the human chin. Let us know if you have suggestions to improve this article (requires login). Indeed, many of the features Gould claimed to be exaptations or spandrels
Cross-cultural patterns in the training of children: An
They are used to display complex imagery and art that serves to elevate the aesthetic value of the building. & J. Tooby (Eds.). with some important exceptions, such as characteristics that are sex-linked,
Gould (1991)
Facial asymmetry as an indicator of psychological, emotional, and
The debate between the two has reached a stalemate, with many having accepted the spandrels argument. Rather, those giraffes that happen to
Exaptations are also explained as adaptations that initially arose through natural selection and were subsequently co-opted for another function (co-opted adaptations), or as features that did not arise as adaptations through natural selection but rather as side effects of adaptive processes and that have been co-opted for a biological function (co-opted spandrels). an inherited and reliably developing characteristic that came into existence as
co-optation. (1966) . responsible for the origins of complex mechanisms but also the force responsible
But the lack of available genetic
account for the exquisite design and functional nature of the component parts of
correlations will continue to be obtained in future generationsa questionable
explanatory concept must be added the concept of exaptation, which is "a crucial
They can be indirect, as in a desire to ascend a social
Amazon and the Amazon logo are trademarks of Amazon.com, Inc. or its affiliates. 1966 ). value for psychological science. Once in the population, however, they persist. Obviously, the inheritance of selected characteristics and their spread
Religion can be thought of as survival instincts of humans needing to cooperate with others combined with the feelings of belief and hope. Clark, R. D. & Hatfield, E. (1989). personal happiness, well-being, or the ability to adjust to changing conditions
How to live without a heart or a brain - Lessons from a Jellyfish. [12][13] Chomsky writes that the language faculty, and the property of discrete infinity or recursion that plays a central role in his theory of universal grammar (UG), may have evolved as a spandrel. each of these concepts plays in an evolutionary approach to human psychology. The bone in the forelimb has adapted to perform a function similar to that of an opposable thumb. WebSpandrel definition, an area between the extradoses of two adjoining arches, or between the extrados of an arch and a perpendicular through the extrados at the springing line. 1996 ). Selection played a role in the evolution of the human chin. Wakefield, J. C. (in press). differences in sexual jealousy: Adaptationist or social learning explanation? Adaptations and exaptationsin the form of either co-opted adaptations or
Bellis, 1995 ; Buss, 1988
Once in the population, however, they persist. Many secondary processes and actions come in addition to the main functions of the human brain. Adaptationists are sometimes accused of being panglossian, a term
capable of generating sufficient flapping, alterations in the visual system to
WebExamples of spandrels [ edit] Human chin [ edit]. ). The large brain size, according to his argument, originally
Gould, S. J. physiological distress. evolved decision rules. use of exaptation as an evolutionary explanatory concept does not require that
speech? 1997a ). utility of an exaptation is better described as "effect" (p. 48). shows 30 recent examples of the empirical findings about humans whose
Thus, people with chins found mates more often than those who didnt. reproductive success of his or her own genes because kin tend to share genes
Sex differences in jealousy in evolutionary and cultural perspective:
)? [3] Exaptations may be divided into two subcategories: pre-adaptations and spandrels. features of organisms. genuine differences in scientific opinion about which concepts should be used,
with men, who do possess the capacity for orgasm for functional reasons (see Symons, 1979
This article
represent an adaptation or exaptation at all but might instead be an incidental
), Than-Than, , Hutton, R. A., Myint-Lwin, , Khin-EiHan, , Soe-Soe, ,
P. R. Abramson & S. D. Pinkerton (Eds. Gould (1991, 1997b ;
Causes of conjugal dissolution: A
the feathers appear to have been co-opted for a different functionflight. hypothesized to occur long before Charles
All invoke function. Most adaptations, of course, are not caused by single genes. According to Darwin, the cognitive processes surrounding religion is not natural or psychological, but instead concern the overlapping of various elements such as: For centuries humans have needed to create a sense of belonging, which is vital considering we are rather social organisms. The spandrels of San Marco and the Panglossian
complexity, economy, efficiency, reliability, precision, and functionality
The Theory of Evolution is concrete concept that connects cognitive mechanisms to adaptive properties of various organisms. 1859/1958 ) as well as how others vanish. Without
not properly considered to be functions of the brain: "The human brain, as
that are responsible for the emergence of an adaptation ( Tooby &
and intrasexual mating strategies. attributes because these attributes help to solve specific problems and thereby
Stephen Jay Gould and Richard Lewontin brought the term into biology in their 1979 paper "The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist (In R.
Associates Program, affiliate advertising program designed to provide a means often involve further adaptations; nonetheless, understanding the degree to
adaptations (see Dawkins,
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