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oth the process and products of evolution the process that creates complex traits and the products functions anatomical structures biochemicals etc that may be imperfectly developed 1 2 the term exaptation was proposed by stephen jay gould and elisabeth vrba as a replacement for pre adaptation which they considered to be a teleologically loaded term 3 i e it falsely implies that adaptation or evolution generally acts in pursuit of some goal history and definitions edit charles darwin the idea that the function of a trait might shift during its evolutionary history originated with charles darwin darwin 1859 for many years the phenomenon was labeled preadaptation but since this term suggests teleology in biology appearing to conflict with natural selection it has been replaced by the term exaptation the idea had been explored by several scholars a when in 1982 stephen jay gould and elisabeth vrba introduced the term exaptation however this definition had two categories with different implications for the role of adaptation 1 a character previously shaped by natural selection for a particular function an adaptation is coopted for a new use cooptation 2 a character whose origin cannot be ascribed to the direct action of natural selection a nonaptation is coopted for a current use cooptation gould and vrba 1982 table 1 the definitions are silent as to whether exaptations had been shaped by natural selection after cooption although gould and vrba cite examples e g feathers of traits shaped after cooption note that the selection pressure upon a trait is likely to change if it is especially primarily or solely used for a new purpose potentially initiating a different evolutionary trajectory to avoid these ambiguities david buss et al suggested the term co opted adaptation which is limited to traits that evolved after cooption 6 however the commonly used terms of exaptation and cooption are ambiguous in this regard preadaptation edit in some circumstances the pre in preadaptation can be interpreted as applying for non teleological reasons prior to the adaptation itself creating a meaning for the term that is distinct from exaptation 7 8 for example future environments say hotter or drier ones may resemble those already encountered by a population at one of its current spatial or temporal margins 7 this is not actual foresight but rather the luck of having adapted to a climate which later becomes more prominent cryptic genetic variation may have the most strongly deleterious mutations purged from it leaving an increased chance of useful adaptations 8 9 but this represents selection acting on current genomes with consequences for the future rather than foresight function may not always come before form developed structures could change or alter the primary functions they were intended for due to some structural or historical cause 10 examples edit bird feathers of various colors exaptations include the co option of feathers which initially evolved for heat regulation for display and later for use in bird flight another example is the lungs of many basal fish which evolved into the lungs of terrestrial vertebrates but also underwent exaptation to become the gas bladder a buoyancy control organ in derived fish 11 a third is the repurposing of two of the three bones in the amniote jaw to become the malleus and incus of the mammalian ear leaving the mammalian jaw with just one hinge 12 arthropods provide the earliest identifiable fossils of land animals from about 419 million years ago in the late silurian and terrestrial tracks from about 450 million years ago appear to have been made by arthropods 13 arthropods were well pre adapted to colonize land because their existing jointed exoskeletons provided support against gravity and mechanical components that could interact to provide levers columns and other means of locomotion that did not depend on submergence in water 14 metabolism can be considered an important part of exaptation as one of the oldest biological systems and being central to life on the earth studies have shown that metabolism may be able to use exaptation in order to increase fitness given some new set of conditions or environment 15 studies have shown that up to 44 carbon sources are viable for metabolism to successfully take place and that any one adaptation in these specific metabolic systems is due to multiple exaptations 16 taking this perspective exaptations are important in the origination of adaptations in general a recent example comes from richard lenski s e coli long term evolution experiment in which aerobic growth on citrate arose in one of twelve populations after 31 000 generations of evolution 17 genomic analysis by blount and colleagues showed that this novel trait was due to a gene duplication that caused a citrate transporter that is normally expressed only under anoxic conditions to be expressed under oxic conditions thus exapting it for aerobic use 18 gould and brosius took the concept of exaptation to the genetic level it is possible to look at a retroposon originally thought to be simply junk dna and deduce that it may have gained a new function to be termed as an exaptation 19 20 21 given an emergency situation in the past a species may have co opted junk dna for a useful purpose this may have occurred with mammalian ancestors when confronted with the permian triassic extinction event about 250 million years ago and a substantial increase in the level of oxygen in earth s atmosphere more than 100 loci have been found to be conserved only among mammalian genomes and are thought to have essential roles in the generation of features such as the placenta diaphragm mammary glands neocortex and auditory ossicles it is believed that as a result of exaptation or making previously useless dna into dna that could be used in order to increase survival chance mammals were able to generate new brain structures as well as behavior to better survive the mass extinction and adapt to new environments similarly viruses and their components have been repeatedly exapted for host functions the functions of exapted viruses typically involve either defense from other viruses or cellular competitors or transfer of nucleic acids between cells or storage functions koonin and krupovic suggested that virus exaptation can reach different depths from recruitment of a fully functional virus to exploitation of defective partially degraded viruses to utilization of individual virus proteins 22 adaptation and exaptation cycle edit it was speculated by gould and vrba 23 in one of the first papers written about exaptation that when an exaptation arises it may not be perfectly suited for its new role and may therefore evolve new adaptations to promote its use in a better manner in other words the beginning of evolving a particular trait starts out with a primary adaptation toward a fit or specific role followed by a primary exaptation a new role is derived using the existing feature but may not be perfect for it which in turn leads to the evolution of a secondary adaptation the feature is improved by natural selection for better performance promoting further evolution of an exaptation and so forth once again feathers are an important example in that they may have first been adapted for thermoregulation and with time became useful for catching insects and therefore served as a new feature for another benefit for instance large contour feathers with specific arrangements arose as an adaptation for catching insects more successfully which eventually led to flight since the larger feathers served better for that purpose implications edit evolution of complex traits edit one of the challenges to darwin s theory of evolution was explaining how complex structures could evolve gradually 24 given that their incipient forms may have been inadequate to serve any function as george jackson mivart a critic of darwin pointed out 5 percent of a bird wing would not be functional the incipient form of complex traits would not have survived long enough to evolve to a useful form as darwin elaborated in the last edition of the origin of species 25 many complex traits evolved from earlier traits that had served different functions by trapping air primitive wings would have enabled birds to efficiently regulate their temperature in part by lifting up their feathers when too warm individual animals with more of this functionality would more successfully survive and reproduce resulting in the proliferation and intensification of the trait eventually feathers became sufficiently large to enable some individuals to glide these individuals would in turn more successfully survive and reproduce resulting in the spread of this trait because it served a second and still more beneficial function that of locomotion hence the evolution of bird wings can be explained by a shifting in function from the regulation of temperature to flight jury rigged design edit darwin explained how the traits of living organisms are well designed for their environment but he also recognized that many traits are imperfectly designed they appear to have been made from available material that is jury rigged b understanding exaptations may suggest hypotheses regarding subtleties in the adaptation for instance that feathers evolved initially for thermal regulation may help to explain some of their features unrelated to flight 26 however this is readily explained by the fact that they serve a dual purpose some of the chemical pathways for physical pain and pain from social exclusion overlap 26 the physical pain system may have been co opted to motivate social animals to respond to threats to their inclusion in the group evolution of technology edit exaptation has received increasing attention in innovation and management studies inspired by evolutionary dynamics where it has been proposed as a mechanism that drives the serendipitous expansion of technologies and products in new domains 27 clarification needed in resilience and regional studies edit in resilience and regional studies exaptation has been used to describe the capacity of complex social economic and territorial systems to repurpose existing resources traits or capabilities for new functions in response to external shocks dávid kollár and józsef kollár introduced the term exaptive resilience in their 2020 article the art of shipwrecking the information society and the rise of exaptive resilience contrasting it with adaptive resilience as a response based on the refunctionalization of existing capacities rather than the preservation of predefined functions 28 the concept has also been applied in territorial research including the espon project terra res territorial exaptive resilience along eu eastern borders which examined how border regions mobilize and repurpose existing assets and capabilities in response to crises and structural change 29 in the cognitive science of religion edit in the cognitive science of religion exaptation is used to explain religious behavior and belief as such religion is seen as the byproduct of mental evolution 30 see also edit architectural exaptation practice of repurposing architectural elements atavism reappearance of a genetic trait once thought extinct ecological fitting biological process evolutionary anachronism attributes of living species that arose due to coevolution with other now extinct species neofunctionalization genetic process repurposing using object intended for one purpose in alternative way spandrel biology evolutionary byproduct of some other characteristic vestigiality evolutionary retention of no longer needed structures in living organisms notes edit see jacob 1977 4 and mayr 1982 5 for references jacob 1977 4 sees much of evolution as tinkering that is working with available traits tinkering includes but is not limited to shifts in function references edit bock w j 1959 preadaptation and multiple evolutionary pathways evolution 13 2 194 211 doi 10 2307 2405873 jstor 2405873 hayden eric j ferrada evandro wagner andreas 2 june 2011 cryptic genetic variation promotes rapid evolutionary adaptation in an rna enzyme pdf nature 474 7349 92 95 doi 10 1038 nature10083 pmid 21637259 s2cid 4390213 gould vrba 1982 1 2 jacob f 1977 evolution and tinkering science 196 4295 1161 6 bibcode 1977sci 196 1161j doi 10 1126 science 860134 pmid 860134 mayr ernst 1982 the growth of biological thought diversity evolution and inheritance harvard university press isbn 978 0 674 36445 5 buss david m martie g haselton todd k shackelford et al 1998 adaptations exaptations and spandrels american psychologist 53 may 533 548 http www sscnet ucla edu comm haselton webdocs spandrels html 1 2 eshel i matessi c 1998 canalization genetic assimilation and preadaptation a quantitative genetic model genetics 149 4 2119 2133 doi 10 1093 genetics 149 4 2119 pmc 1460279 pmid 9691063 cite journal cs1 maint 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