Neoteny In Humans
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Neoteny in humans is the retention of juvenile traits well into adulthood. This trend is greatly amplified in humans especially when compared to non-human
primates Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
. Adult humans more closely resemble the infants of
gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or fi ...
s and
chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative th ...
s than the adults of those animals.
Neotenic Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compared ...
features of the head include the globular skull; (page 134), cited by:

"In humans, neoteny is manifested in the resemblance of many physiological features of a human to a late-stage foetal chimpanzee. These foetal characteristics include hair on the head, a globular skull, ear shape, vertical plane face, absence of penal bone (baculum) in foetal male chimpanzees, the vagina pointing forward in foetal ape, the presence of hymen in neonate ape, and the structure of the foot. 'These and many other features', Bednarik says, 'define the anatomical relationship between ape and man as the latter's neoteny'".
thinness of skull bones; the reduction of the brow ridge; the large brain; the flattened and broadened face; the hairless face; hair on (top of) the head; larger eyes; ear shape; small nose; small teeth; and the small
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
(upper jaw) and
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
(lower jaw). Neoteny of the human body is indicated by
glabrousness Glabrousness (from the Latin ''glaber'' meaning "bald", "hairless", "shaved", "smooth") is the technical term for a lack of hair, down, setae, trichomes or other such covering. A glabrous surface may be a natural characteristic of all or part of ...
(hairless body). Neoteny of the genitals is marked by the absence of a
baculum The baculum (also penis bone, penile bone, or ''os penis'', ''os genitale'' or ''os priapi'') is a bone found in the penis of many placental mammals. It is absent from the human penis, but present in the penises of some primates, such as the ...
(penis bone); the presence of a
hymen The hymen is a thin piece of mucosal tissue that surrounds or partially covers the external vaginal opening. It forms part of the vulva, or external genitalia, and is similar in structure to the vagina. In children, a common appearance of the h ...
; and the forward-facing
vagina In mammals, the vagina is the elastic, muscular part of the female genital tract. In humans, it extends from the vestibule to the cervix. The outer vaginal opening is normally partly covered by a thin layer of mucosal tissue called the hymen ...
. Neoteny in humans is further indicated by the limbs and body posture, with the limbs proportionately short compared to torso length; longer leg than arm length; the structure of the foot; and the upright stance. Humans also retain a plasticity of
behavior Behavior (American English) or behaviour (British English) is the range of actions and mannerisms made by individuals, organisms, systems or artificial entities in some environment. These systems can include other systems or organisms as wel ...
that is generally found among animals only in the young. The emphasis on learned, rather than inherited, behavior requires the human brain to remain receptive much longer. These
neotenic Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compared ...
changes may have disparate roots. Some may have been brought about by
sexual selection in human evolution Sexual selection in humans concerns the concept of sexual selection, introduced by Charles Darwin as an element of his theory of natural selection, as it affects humans. Sexual selection is a biological way one sex chooses a mate for the best repr ...
. In turn, they may have permitted the development of human capacities such as emotional communication. However, humans also have relatively large noses and long legs, both
peramorphic In evolutionary developmental biology, heterochrony is any genetically controlled difference in the timing, rate, or duration of a developmental process in an organism compared to its ancestors or other organisms. This leads to changes in the si ...
(not neotenic) traits, though these peramorphic traits separating modern humans from extant chimpanzees were present in ''
Homo erectus ''Homo erectus'' (; meaning "upright man") is an extinct species of archaic human from the Pleistocene, with its earliest occurrence about 2 million years ago. Several human species, such as '' H. heidelbergensis'' and '' H. antecessor' ...
'' to an even higher degree than in ''
Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
'', which means general neoteny is valid for the ''H.erectus'' to ''H.sapiens'' transition (although there were perimorphic changes separating ''H.erectus'' from even earlier hominins such as most ''
Australopithecus ''Australopithecus'' (, ; ) is a genus of early hominins that existed in Africa during the Late Pliocene and Early Pleistocene. The genus ''Homo'' (which includes modern humans) emerged within ''Australopithecus'', as sister to e.g. ''Australopi ...
''). Later research shows that some species of ''Australopithecus'', including ''
Australopithecus sediba ''Australopithecus sediba'' is an extinct species of australopithecine recovered from Malapa Cave, Cradle of Humankind, South Africa. It is known from a partial juvenile skeleton, the holotype MH1, and a partial adult female skeleton, the para ...
'', had the non-neotenic traits of ''H.erectus'' to at least the same extent which separate them from other ''Australopithecus'', making it possible that general neoteny applies throughout the evolution of the genus ''Homo'' depending on what species of ''Australopithecus'' that ''Homo'' descended from. The
type specimen In biology, a type is a particular wiktionary:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to a ...
of ''A.sediba'' had these non-neotenic traits, despite being a juvenile, suggesting that the adults may have been less neotenic in these regards than any ''H.erectus'' or other ''Homo''.


Neoteny and heterochrony

Heterochrony is defined as “a genetic shift in timing of the development of a tissue or anatomical part, or in the onset of a physiological process, relative to an ancestor”. Heterochrony can lead to a modification in shape, size and/or behavior of an organism through a variety of different ways. With heterochrony being more of an umbrella term, there are two different types of heterochrony where development timing is altered: paedomorphosis and peramorphosis. These terms refer to deceleration and acceleration of development, respectively. With neoteny (as described above) being defined as retention of juvenile features into adulthood, neoteny falls under paedomorphosis, as physical development of features is slowed.


Human evolution

Many prominent evolutionary theorists propose that neoteny has been a key feature in
human evolution Human evolution is the evolutionary process within the history of primates that led to the emergence of ''Homo sapiens'' as a distinct species of the hominid family, which includes the great apes. This process involved the gradual development of ...
.
Stephen Jay Gould Stephen Jay Gould (; September 10, 1941 – May 20, 2002) was an American paleontologist, evolutionary biologist, and historian of science. He was one of the most influential and widely read authors of popular science of his generation. Gould sp ...
believed that the "evolutionary story" of humans is one where we have been "retaining to adulthood the originally juvenile features of our ancestors".
J. B. S. Haldane John Burdon Sanderson Haldane (; 5 November 18921 December 1964), nicknamed "Jack" or "JBS", was a British-Indian scientist who worked in physiology, genetics, evolutionary biology, and mathematics. With innovative use of statistics in biolog ...
mirrors Gould's hypothesis by stating a "major evolutionary trend in human beings" is "greater prolongation of childhood and retardation of maturity." Delbert D. Thiessen said that "neoteny becomes more apparent as early primates evolved into later forms" and that primates have been "evolving toward flat face." Doug Jones, a
visiting scholar In academia, a visiting scholar, visiting researcher, visiting fellow, visiting lecturer, or visiting professor is a scholar from an institution who visits a host university to teach, lecture, or perform research on a topic for which the visitor ...
in anthropology at
Cornell University Cornell University is a private statutory land-grant research university based in Ithaca, New York. It is a member of the Ivy League. Founded in 1865 by Ezra Cornell and Andrew Dickson White, Cornell was founded with the intention to teach an ...
, said that human evolution's trend toward neoteny may have been caused by
sexual selection in human evolution Sexual selection in humans concerns the concept of sexual selection, introduced by Charles Darwin as an element of his theory of natural selection, as it affects humans. Sexual selection is a biological way one sex chooses a mate for the best repr ...
for neotenous facial traits in women by men with the resulting neoteny in male faces being a "by-product" of
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of t ...
for neotenous female faces. Jones said that this type of sexual selection "likely" had a major role in human evolution once a larger proportion of women lived past the age of
menopause Menopause, also known as the climacteric, is the time in women's lives when menstrual periods stop permanently, and they are no longer able to bear children. Menopause usually occurs between the age of 47 and 54. Medical professionals often d ...
. This increasing proportion of women who were too old to reproduce resulted in a greater variance in fecundity in the population of women, and it resulted in a greater sexual selection for indicators of youthful fecundity in women by men. The anthropologist
Ashley Montagu Montague Francis Ashley-Montagu (June 28, 1905November 26, 1999) — born Israel Ehrenberg — was a British-American anthropologist who popularized the study of topics such as Race (classification of human beings), race and gender and their rel ...
said that the fetalized ''Homo erectus'' represented by the juvenile
Mojokerto Mojokerto ( jv, ꦩꦗꦏꦼꦂꦠ (''Måjåkěrtå'')) is a city in East Java Province, Indonesia. It is located 40 km southwest of Surabaya, and constitutes one of the component units of the Surabaya metropolitan area (known as Gerbangkert ...
skull and the fetalized australopithecine represented by the juvenile ''
Australopithecus africanus ''Australopithecus africanus'' is an extinct species of australopithecine which lived between about 3.3 and 2.1 million years ago in the Late Pliocene to Early Pleistocene of South Africa. The species has been recovered from Taung, Sterkfonte ...
'' skull would have had skulls with a closer resemblance to those of modern humans than to those of the adult forms of their own species. Montagu further listed the roundness of the skull, thinness of the skull bones, lack of
brow ridge The brow ridge, or supraorbital ridge known as superciliary arch in medicine, is a bony ridge located above the eye sockets of all primates. In humans, the eyebrows are located on their lower margin. Structure The brow ridge is a nodule or crest ...
s, lack of
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
s, form of the teeth, relative size of the brain and form of the brain as ways in which the juvenile skulls of these human ancestors resemble the skulls of adult modern humans. Montagu said that the retention of these juvenile characteristics of the skull into adulthood by australopithecine or ''H.erectus'' could have been a way that a modern type of human could have evolved earlier than what actually happened in human evolution. The psychiatrist
Stanley Greenspan Stanley Greenspan (June 1, 1941 – April 27, 2010) was a clinical professor of Psychiatry, Behavioral Science, and Pediatrics at George Washington University Medical School and a practicing child psychiatrist. He was best known for developing the ...
and Stuart G. Shanker proposed a theory in ''The First Idea'' of psychological development in which neoteny is seen as crucial for the "development of species-typical capacities" that depend upon a long period of attachment to caregivers for the opportunities to engage in and develop their capacity for emotional communication. Because of the importance of facial expression in the process of interactive signaling, neotenous features, such as hair loss, allow for more efficient and rapid communication of socially important messages that are based on facially expressive emotional signaling. Other theorists have argued that neoteny has not been the main cause of human evolution, because humans only retain some juvenile traits, while relinquishing others. For example, the high leg-to-body ratio (long legs) of adult humans as opposed to human infants shows that there is not a holistic trend in humans towards neoteny when compared to the other
great apes The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the east ...
.
Andrew Arthur Abbie Andrew Arthur Abbie, also known as A. A. Abbie (8 February 1905 – 22 July 1976) was an Australian anatomist and anthropologist. Biography Born in Gillingham, Kent, Abbie was educated at a mathematical school in Rochester where he obtained a s ...
agrees, citing the gerontomorphic fleshy human nose and long human legs as contradicting the neoteny hominid evolution hypothesis, although he does believe humans are generally neotenous. Brian K. Hall also cites the long legs of humans as a
peramorphic In evolutionary developmental biology, heterochrony is any genetically controlled difference in the timing, rate, or duration of a developmental process in an organism compared to its ancestors or other organisms. This leads to changes in the si ...
trait, which is in sharp contrast to neoteny. On the balance, an all or nothing approach could be regarded as pointless, with a combination of heterochronic processes being more likely and more reasonable (Vrba, 1996).


Cooked food and protective genome simplification

Based on calculations that show that more complex gene networks are more vulnerable to mutations as more conditions that are necessary but not sufficient increases the risk of one of them being hit, there is a theory that
mutagen In genetics, a mutagen is a physical or chemical agent that permanently changes nucleic acid, genetic material, usually DNA, in an organism and thus increases the frequency of mutations above the natural background level. As many mutations can ca ...
s in food cooked by human ancestors short of modern human intelligence that was more likely to be burnt than in modern cooking selected against complex gene networks. This theory successfully predicts that the human genome is shorter than other Great Ape genomes and that there are significantly more defunct
pseudogene Pseudogenes are nonfunctional segments of DNA that resemble functional genes. Most arise as superfluous copies of functional genes, either directly by DNA duplication or indirectly by Reverse transcriptase, reverse transcription of an mRNA trans ...
s with functional
homolog In biology, homology is similarity due to shared ancestry between a pair of structures or genes in different taxa. A common example of homologous structures is the forelimbs of vertebrates, where the wings of bats and birds, the arms of prima ...
s in the chimpanzee genome than vice versa. While the protein coding portion of the
FOXP2 Forkhead box protein P2 (FOXP2) is a protein that, in humans, is encoded by the ''FOXP2'' gene. FOXP2 is a member of the forkhead box family of transcription factors, proteins that regulate gene expression by binding to DNA. It is expressed in ...
gene is identical to that in
Neanderthal Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ago. While th ...
s, there is one point mutation in the regulatory part thereof (modern humans having a T where Neanderthals and all nonhuman vertebrates have an A). The observation that the effect of that difference is that the modern human FOXP2 gene does not interact with
RNA Ribonucleic acid (RNA) is a polymeric molecule essential in various biological roles in coding, decoding, regulation and expression of genes. RNA and deoxyribonucleic acid ( DNA) are nucleic acids. Along with lipids, proteins, and carbohydra ...
from other genes while all other vertebrate including Neanderthal varieties did agrees with the idea that modern human origin was marked by the elimination (not formation) of complex gene networks, as predicted by this model. The researchers behind the theory argue that neoteny is a side effect of the destruction of gene networks preventing the firing of genetic activity patterns that marked adulthood in prehuman ancestors.


Growth pattern of children

In 1943
Konrad Lorenz Konrad Zacharias Lorenz (; 7 November 1903 – 27 February 1989) was an Austrian zoologist, ethologist, and ornithologist. He shared the 1973 Nobel Prize in Physiology or Medicine with Nikolaas Tinbergen and Karl von Frisch. He is often regarded ...
noted that a newborn infant's rounded facial features might encourage guardians to show greater care for them, due to their perceived cuteness. He labeled this the
Kewpie doll effect The Kewpie doll effect is a term used in developmental psychology derived from research in ethology to help explain how a child's physical features, such as lengthened forehead and rounded face, motivate the infant's caregiver to take care of them ...
, because of their similarity to the eponymous doll. Desmond Collins who was an Extension Lecturer of Archaeology at
London University The University of London (UoL; abbreviated as Lond or more rarely Londin in post-nominals) is a federal public research university located in London, England, United Kingdom. The university was established by royal charter in 1836 as a degree- ...
said that the lengthened youth period of humans is part of neoteny. Physical anthropologist
Barry Bogin Barry Bogin (born May 20, 1950) is an American physical anthropologist trained at Temple University who researches physical growth in Guatemalan Maya children, and is a theorist upon the evolutionary origins of human childhood. He is a professo ...
said that the pattern of children's growth may intentionally increase the duration of their cuteness. Bogin said that the human brain reaches adult size when the body is only 40 percent complete, when "dental maturation is only 58 percent complete" and when "reproductive maturation is only 10 percent complete". Bogin said that this
allometry Allometry is the study of the relationship of body size to shape, anatomy, physiology and finally behaviour, first outlined by Otto Snell in 1892, by D'Arcy Thompson in 1917 in ''On Growth and Form'' and by Julian Huxley in 1932. Overview Allom ...
of human growth allows children to have a "superficially infantile" appearance (large
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
, small face, small body and sexual underdevelopment) longer than in other "
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
ian species". Bogin said that this cute appearance causes a "nurturing" and "care-giving" response in "older individuals".


Genetic diversity, relaxed sexual selection and immunity

While upper body strength is on average more
sexually dimorphic Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
in humans than in most other primates, with the exception of
gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or fi ...
s, some fossil evidence suggests that male upper-body strength and muscular sexual dimorphism during human evolution peaked in ''Homo erectus'' and decreased, along with overall
robustness Robustness is the property of being strong and healthy in constitution. When it is transposed into a system, it refers to the ability of tolerating perturbations that might affect the system’s functional body. In the same line ''robustness'' ca ...
, during the evolution of ''H.sapiens'' with its neotenic traits. The reduction in sexual dimorphism would suggest that
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
with high sexual dimorphism do not necessarily have an increased evolutionary advantage. This could be explained by the theory that sexual dimorphism could reduce
genetic diversity Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species, it ranges widely from the number of species to differences within species and can be attributed to the span of survival for a species. It is dis ...
in a population, i.e., if individuals are attracted to only highly masculine or highly feminine mates, then those without distinctly gendered features are excluded as potential partners, thus creating
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
. Neoteny in ''H.sapiens'' is explained by this theory as a result of relaxed sexual selection shifting human evolution into a less speciation-prone but more intraspecies adaptable strategy, decreasing sexual dimorphism and making adults assume a more juvenile form. As a possible trigger of such a change, it has been cited that while the Neanderthal version of the
FOXP2 Forkhead box protein P2 (FOXP2) is a protein that, in humans, is encoded by the ''FOXP2'' gene. FOXP2 is a member of the forkhead box family of transcription factors, proteins that regulate gene expression by binding to DNA. It is expressed in ...
gene differed on only one point from the modern human version (not two points as the difference between chimpanzees and modern humans) interacted strongly with other genes and was part of a
gene regulatory network A gene (or genetic) regulatory network (GRN) is a collection of molecular regulators that interact with each other and with other substances in the cell to govern the gene expression levels of mRNA and proteins which, in turn, determine the fun ...
, the derived mutation that is unique to the modern human version of the gene knocked out the attachment to which
RNA Ribonucleic acid (RNA) is a polymeric molecule essential in various biological roles in coding, decoding, regulation and expression of genes. RNA and deoxyribonucleic acid ( DNA) are nucleic acids. Along with lipids, proteins, and carbohydra ...
strains from other genes connected to it so that the gene was disconnected from its former genetic network. It is suggested that since the FOXP2 gene controls
synapse In the nervous system, a synapse is a structure that permits a neuron (or nerve cell) to pass an electrical or chemical signal to another neuron or to the target effector cell. Synapses are essential to the transmission of nervous impulses from ...
s, its disconnection from a formerly complex network of genes instantly removed many instincts including ones that drove sexual selection. It is also suggested that it allowed more genetic variants that affect the
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological proper ...
to accumulate in humans, which in combination with increased
synaptic plasticity In neuroscience, synaptic plasticity is the ability of synapses to strengthen or weaken over time, in response to increases or decreases in their activity. Since memories are postulated to be represented by vastly interconnected neural circuit ...
made modern humans more able to survive environmental change and to colonize new environments and innovate. The theory that the origin of complex language was the most recent step in human evolution is considered unlikely as storytelling about past environments would be of little use in droughts with novel distributions of water while individual ability to make correct predictions would be useful and allow for differential survival that could eliminate the archaic version altogether, as opposed to selection for language in which some primitives could use imitation as long as there were enough storytellers in the group to keep the knowledge alive for long times which predicts that some individuals would have retained the archaic version if the modern version was for language. ''H.sapiens'' is known from fossils to have had a mix of modern neotenic traits and older non-neotenic traits from its origin some 300000 years ago to the transition to early agriculture when the non-neotenic traits disappeared, which is theorized to be due to selection for the
immune system The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as cancer cells and objects such as wood splinte ...
adapting to survive a higher pathogen load caused by agriculture and men who retained more childlike traits being less burdened by weakening of the immune system from upper body musculature competing with the immune system over nutrients. It is argued that the genetic evidence of only a small part of the male population of the time of early agriculture passing on their
Y chromosome The Y chromosome is one of two sex chromosomes (allosomes) in therian mammals, including humans, and many other animals. The other is the X chromosome. Y is normally the sex-determining chromosome in many species, since it is the presence or abse ...
s can be explained by the heredity of non-neotenic traits causing the male descendants of the non-neotenic men who were not killed by diseases in one generation to die from them in subsequent generations, leaving no Y chromosome evidence of their short term continuation of paternal bloodlines in present humans. Sexual selection for stereotypic masculinity causing most men to fail to breed is ruled out as it would have selected against neoteny, not for as the archaeological evidence shows.


Milder punishment as a survival advantage

One hypothesis of the premise that Stone Age humans did not record birth date but instead assumed age based on appearance holds that if milder
punishment Punishment, commonly, is the imposition of an undesirable or unpleasant outcome upon a group or individual, meted out by an authority—in contexts ranging from child discipline to criminal law—as a response and deterrent to a particular acti ...
to juvenile delinquents existed in
Paleolithic The Paleolithic or Palaeolithic (), also called the Old Stone Age (from Greek: παλαιός ''palaios'', "old" and λίθος ''lithos'', "stone"), is a period in human prehistory that is distinguished by the original development of stone too ...
times, it would have imparted milder punishment for longer on those retaining a more youthful appearance into adulthood. This hypothesis posits that those who got milder punishment for the same breach of rules had the evolutionary advantage, passing their genes on while those who got more severe punishment had more limited reproductive success due to either limiting their survival by following all rules or by being severely punished.


Neotenous features elicit help

The Multiple Fitness Model proposes that the qualities that make babies appear
cute Cuteness is a subjective term describing a type of attractiveness commonly associated with youth and appearance, as well as a scientific concept and analytical model in ethology, first introduced by Konrad Lorenz. Lorenz proposed the concept ...
to adults additionally look "desirable" to adults when they see other adults. Neotenous features in adult females may help elicit more resource investment and nurturing from adult males. Likewise, neotenous features in adult males may similarly help elicit more resource investment and nurturing from adult females in addition to possibly making neotenous adult males appear less threatening and possibly making neotenous adult males more able to elicit resources from "other resource-rich people". Therefore, it could be adaptive for adult females to be attracted to adult males that have "some" neotenous traits. Neotenous features elicits fitness benefits for mimickers. From the point of view of the mimicker, the neoteny expression signals appeasement or submissiveness. Thus, extra parental or alloparental care will be most likely be administered because the mimicker appears to be more childlike and maybe ill-equipped to survive on its own. On the other hand, the recipient often faces aggression because of this signaled vulnerability. Caroline F. Keating et al. tested the hypothesis that adult male and female faces with more neotenous features would elicit more help than adult male and female faces with less neotenous features. Keating et al. digitally modified photographs of faces of
African-American African Americans (also referred to as Black Americans and Afro-Americans) are an Race and ethnicity in the United States, ethnic group consisting of Americans with partial or total ancestry from sub-Saharan Africa. The term "African American ...
s and
European American European Americans (also referred to as Euro-Americans) are Americans of European ancestry. This term includes people who are descended from the first European settlers in the United States as well as people who are descended from more recent Eu ...
s to make them appear more or less neotenous by either enlarging or decreasing the size of their eyes and lips. Keating et al. said that the more neotenous white male, white female and black female faces elicited more help from people in the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
and
Kenya ) , national_anthem = "Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi , ...
, but the difference in help from people in the United States and Kenya for more neotenous black male faces was not significantly different from less neotenous black male faces. A 1987 study using 20 Caucasian subjects found that "babyfaced" individuals are assumed by both Korean and U.S. participants to possess more childlike psychological attributes than their mature-faced counterparts. In her dissertation from the
University of Michigan , mottoeng = "Arts, Knowledge, Truth" , former_names = Catholepistemiad, or University of Michigania (1817–1821) , budget = $10.3 billion (2021) , endowment = $17 billion (2021)As o ...
,
Sookyung Choi SooKyung Choi is a South Korean particle physicist at Gyeongsang National University. She is part of the Belle experiment and was the first to observe the X(3872) meson in 2003. She won the 2017 Ho-Am Prize in Science. Early life and education ...
explained how perception of cuteness can contribute to perception of value. Different physical cues were shown to trigger protective feelings from their adult caregivers or other adults from which they engaged in interaction. Participants in the study were asked to design their own version of a cute rectangle. They were allowed to edit the rectangle in terms of shape roundedness, color, size, orientation, etc. Associational coefficients showed that shapes with a smaller area and rounder features were found to be cuter, and that lighter coloring and contrast playing a lesser but important role in predicting cuteness. As an additional part of the study, the asymmetric dominance paradigm was introduced, where a decoy option is presented to observe how it affects a person's decision on a certain matter. In the United States this asymmetric dominance paradigm induced a person to be more prone to a cuter item, whereas in Korea the opposite effect occurred. Cho concluded that this may be due to a different attitude toward cuteness, and so the advantages related to neoteny may be different in different countries.


Brain

The developmental psychologist
Helmuth Nyborg Helmuth Sørensen Nyborg (born 5 January 1937) is a Danish psychologist and athlete. He is a former professor of developmental psychology at Aarhus University, Denmark and Olympic canoeist. His main research topic is the connection between horm ...
said that a testable
hypothesis A hypothesis (plural hypotheses) is a proposed explanation for a phenomenon. For a hypothesis to be a scientific hypothesis, the scientific method requires that one can test it. Scientists generally base scientific hypotheses on previous obse ...
can be made using his General Trait Covariance-Androgen/Estrogen (GTC-A/E) model with regards to "neoteny". Nyborg said that the hypothesis is that "feminized", slower maturing, "neotenic" " androtypes" will differ from "masculinized", faster maturing "androtypes" by having bigger brains, more fragile skulls, bigger hips, narrower shoulders, less physical strength, live in cities (as opposed to living in the countryside) and by receiving higher performance scores on ability tests. Nyborg said that if the predictions made by this hypothesis are true, then the "material basis" of the differences would be "explained". Nyborg said that some
ecological Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps wi ...
situations would favor the survival and reproduction of the "masculinized," faster maturing "androtypes" due to their "sheer brutal force" while other ecological situations would favor the survival and reproduction of the "feminized," slower maturing, "neotenic" "androtypes" due to their "subtle tactics." Aldo Poiani, an evolutionary ecologist at
Monash University Monash University () is a public research university based in Melbourne, Victoria, Australia. Named for prominent World War I general Sir John Monash, it was founded in 1958 and is the second oldest university in the state. The university has a ...
,
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
, said that he agrees that neoteny in humans may have become "accelerated" through "two-way
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of t ...
" whereby females have been choosing smart males as mates and males have been choosing smart females as mates. Somel et al. said that 48% of the genes that affect the development of the
prefrontal cortex In mammalian brain anatomy, the prefrontal cortex (PFC) covers the front part of the frontal lobe of the cerebral cortex. The PFC contains the Brodmann areas BA8, BA9, BA10, BA11, BA12, BA13, BA14, BA24, BA25, BA32, BA44, BA45, BA46, ...
change with age differently between humans and chimpanzees. Somel et al. said that there is a "significant excess of genes" related to the development of the prefrontal cortex that show "neotenic expression in humans" relative to
chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative th ...
s and
rhesus macaque The rhesus macaque (''Macaca mulatta''), colloquially rhesus monkey, is a species of Old World monkey. There are between six and nine recognised subspecies that are split between two groups, the Chinese-derived and the Indian-derived. Generally b ...
s. Somel et al. said that this difference was in accordance with the neoteny hypothesis of human evolution. In terms of brain size differences, it has been noted that given the larger skull in neoteny humans, brain volume may be larger than an average human brain. It has been hypothesized that this is one mode of which the brains of ''Homo sapiens'' grew as a species, as the prolonged development of neurons may have led to hypermorphosis, or excessive neuronal growth. Especially in the prefrontal cortex, brain pruning from childhood may be slower than usual, allowing for more time for neuronal maturation. This prolongs the transformation of otherwise very juvenile features.
Bruce Charlton Bruce Graham Charlton is a retired British medical doctor who was Visiting Professor of Theoretical Medicine at the University of Buckingham. Until April 2019, he was Reader (academic rank), Reader in Evolutionary Psychiatry at Newcastle University ...
, a
Newcastle University Newcastle University (legally the University of Newcastle upon Tyne) is a UK public university, public research university based in Newcastle upon Tyne, North East England. It has overseas campuses in Singapore and Malaysia. The university is ...
psychology professor, said what looks like immaturity — or in his terms, the "retention of youthful attitudes and behaviors into later adulthood" — is actually a valuable developmental characteristic, which he calls psychological neoteny. The ability of an adult human to learn is considered a neotenous trait. However, some studies may suggest the opposite of this idea of neoteny being beneficial. In general, the process of learning and developing new skills can be attributed to plasticity of neurons in the brain, especially in the prefrontal cortex for higher order decisions and activity. As neurons go through
ontogeny Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the stu ...
and maturity, it becomes more difficult to make new neuronal connections and change already present pathways and connections. However, during juvenile periods, cortical neurons are described to have higher plasticity and metabolic activity. In cases with neoteny, neurons are lingering in their more juvenile states since development is decelerated. On the surface this seems beneficial for the increased potential of younger cells. However, this may not be the case, as the consequences of the increased cellular activity must be taken into account. In general, oxidative phosphorylation is the process used to supply energy for neuronal processes in the brain. When resources for oxidative phosphorylation are exhausted, neurons turn to aerobic glycolysis in the place of oxygen. However, this can be taxing on a cell. Given that the neurons in question retain juvenile characteristics, they may not be entirely myelinated. Bufill, Agusti, Blesa et al. note how “The increase of the aerobic metabolism in these neurons may lead, however, to higher levels of oxidative stress, therefore, favoring the development of neurodegenerative diseases which are exclusive, or almost exclusive, to humans, such as Alzheimer's disease.” Specifically through various studies of the brain, aerobic glycolysis activity has been detected at high levels in the dorsolateral prefrontal cortex, which has functionality regarding the working memory. Stress on these working memory cells may support conditions related to neurodegenerative diseases such as Alzheimer's Disease.


Physical attractiveness


Women

Montagu said that the following neotenous traits are in women when compared to men: more delicate skeleton, smoother ligament attachments, smaller mastoid processes, reduced
brow ridge The brow ridge, or supraorbital ridge known as superciliary arch in medicine, is a bony ridge located above the eye sockets of all primates. In humans, the eyebrows are located on their lower margin. Structure The brow ridge is a nodule or crest ...
s, more forward tilt of the head, narrower joints, less hairy, retention of fetal body hair, smaller body size, more backward tilt of pelvis, greater longevity, lower basal metabolism, faster heartbeat, greater extension of development periods, higher pitched voice and larger tear ducts. In a cross-cultural study, more neotenized female faces were the most attractive to men while less neotenized female faces were the least attractive to men, regardless of the females' actual age. Using a panel of
Asian Asian may refer to: * Items from or related to the continent of Asia: ** Asian people, people in or descending from Asia ** Asian culture, the culture of the people from Asia ** Asian cuisine, food based on the style of food of the people from Asi ...
,
Hispanic The term ''Hispanic'' ( es, hispano) refers to people, Spanish culture, cultures, or countries related to Spain, the Spanish language, or Hispanidad. The term commonly applies to countries with a cultural and historical link to Spain and to Vic ...
, and
White White is the lightest color and is achromatic (having no hue). It is the color of objects such as snow, chalk, and milk, and is the opposite of black. White objects fully reflect and scatter all the visible wavelengths of light. White on ...
judges, Michael R. Cunningham found that the Asian, Hispanic and White female faces found most attractive were those that had "neonate large eyes, greater distance between eyes, and small noses" and his study led him to conclude that "large eyes" were the most "effective" of the "neonate cues". Cunningham also said that "shiny" hair may be indicative of "neonate vitality". Cunningham said that there was a "difference" in the preferences of Asian and White judges. Cunningham said that Asian judges preferred women with "less mature faces" and smaller mouths than the White judges. Cunningham hypothesized that this difference in preference may stem from "ethnocentrism" since "Asian faces possess those qualities", so Cunningham re-analyzed the data with "11 Asian targets excluded" and he concluded that "ethnocentrism was not a primary determinant of Asian preferences." Using a panel of
Black Black is a color which results from the absence or complete absorption of visible light. It is an achromatic color, without hue, like white and grey. It is often used symbolically or figuratively to represent darkness. Black and white have o ...
and White as judges, Cunningham said that more neotenous faces were perceived as having both higher "femininity" and "sociability". In contrast, Cunningham said that faces that were "low in neoteny" were judged as "intimidating". Upon analyzing the results of his study Cunningham concluded that preference for "neonate features may display the least cross-cultural variability" in terms of "attractiveness ratings". In a study of Italian women who have won beauty competitions, the study said that the women had faces characterized by more "babyness" traits compared to the "normal" women used as a reference. In a study of sixty Caucasian female faces, the average facial composite of the fifteen faces considered most attractive differed from the facial composite of the whole by having a reduced lower facial region, a thinner jaw, and a higher forehead. In a solely Westernized study, it was recorded that the high ratio of neurocranial to lower facial features, signified by a small nose and ears, and full lips, is viewed interchangeably as both youthful and or neotenous. This interchangeability between neotenous features and youth leads to the idea that male attraction to youth may also apply to females that display exaggerated age-related cues. For example, if a female was much older but retained these “youthful” features, males may find her more favorable over other females who look their biological age. Beyond the face value of what males find physically attractive, secondary sexual characteristics related to body shape are factored in so adults may be able to recognize other adults from juveniles. A major part of the cosmetic world is built around capitalizing on enhancing these neonate features. Making eyes and lips appear larger as well as reducing the appearance of any age-related blemishes such as wrinkles or skin discoloration are some of the key target areas of this industry. Doug Jones, a
visiting scholar In academia, a visiting scholar, visiting researcher, visiting fellow, visiting lecturer, or visiting professor is a scholar from an institution who visits a host university to teach, lecture, or perform research on a topic for which the visitor ...
in anthropology at
Cornell University Cornell University is a private statutory land-grant research university based in Ithaca, New York. It is a member of the Ivy League. Founded in 1865 by Ezra Cornell and Andrew Dickson White, Cornell was founded with the intention to teach an ...
, said that there is cross-cultural evidence for preference for facial neoteny in women, because of
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of t ...
for the appearance of youthful fecundity in women by men. Jones said that men are more concerned about women's sexual attractiveness than women are concerned about men's sexual attractiveness. Jones said that this greater concern over female attractiveness is unusual among animals, because it is usually the females that are more concerned with the male's sexual attractiveness in other species. Jones said that this anomalous case in humans is due to women living past their reproductive years and due to women having their reproductive capacity diminish with age, resulting in the adaption in men to be selective against physical traits of age that indicate lessening female fecundity. Jones said that the neoteny in men's faces may be a "by-product" of men's attraction to indicators of "youthful fecundity" in "adult females". Likewise, neotenous features have also been loosely linked to providing information about levels of ovarian function, which is another integral part of
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of t ...
. Both of these factors, seeming like extra help is needed as well as neotenous features expression, being tied to optimal ovarian function, lead to a fitness advantage because males respond positively. However, it was noted that neotenous face structures are not the only thing to be taken into consideration when thinking about attractiveness and mate selection. Once again,
secondary sex characteristic Secondary sex characteristics are features that appear during puberty in humans, and at sexual maturity in other animals. These characteristics are particularly evident in the sexually dimorphic phenotypic traits that distinguish the sexes of a sp ...
s come into play because they are dominated by the
endocrine system The endocrine system is a messenger system comprising feedback loops of the hormones released by internal glands of an organism directly into the circulatory system, regulating distant target organs. In vertebrates, the hypothalamus is the neu ...
and appear only when sexual maturity is reached. The facial features are ever present and may not be the strongest case for sexual selection. Other scientists, noting that other primates have not evolved neoteny to the same extent as humans despite fertility being as reproductively significant for them, argue that if human children need more
parental investment Parental investment, in evolutionary biology and evolutionary psychology, is any parental expenditure (e.g. time, energy, resources) that benefits offspring.Clutton-Brock, T.H. 1991. ''The Evolution of Parental Care''. Princeton, NJ: Princeton ...
than nonhuman primate young, that would have selected for a preference for more experienced females more capable of providing parental care. As this would make experience more relevant for effective reproductive success (producing offspring that survive to reproductive age, as opposed to simply the number of births) and therefore more able to compensate for a slight to moderate decrease in biological fertility from recent sexual maturity to late pre-menopausal life, these scientists argue that the sexual selection model of neoteny makes the false prediction that primates that need less parental investment than humans should display more neoteny than humans.


Men

A study was conducted on the attractiveness of males with the subject of the skull and its application in human morphology, using psychology and evolutionary biology to understand selection on facial features. It found that averageness was the result of
stabilizing selection Stabilizing selection (not to be confused with negative or purifying selection) is a type of natural selection in which the population mean stabilizes on a particular non-extreme trait value. This is thought to be the most common mechanism of a ...
, whereas facial paedomorphosis or juvenile traits had been caused by
directional selection In population genetics, directional selection, is a mode of negative natural selection in which an extreme phenotype is favored over other phenotypes, causing the allele frequency to shift over time in the direction of that phenotype. Under dir ...
. In directional selection, a single
phenotypic trait A phenotypic trait, simply trait, or character state is a distinct variant of a phenotypic characteristic of an organism; it may be either inherited or determined environmentally, but typically occurs as a combination of the two.Lawrence, Eleano ...
is driven by selection toward fixation in a population. In contrast, in stabilizing selection both alleles are driven toward fixation (or polymorphism) in a population. To compare the effects of directional and stabilizing selection on facial paedomorphosis, Wehr used graphic morphing to alter appearances to make faces appear more or less juvenile. The results concluded that the effect of averageness was preferred nearly twice over juvenile trait characteristics which indicates that stabilizing selection influences facial preference, and averageness was found more attractive than the retention of juvenile facial characteristics. It was perplexing to find that women tend to prefer the average facial features over the juvenile, because in animals the females tend to drive sexual selection by female choice and the
Red Queen hypothesis The Red Queen hypothesis is a hypothesis in evolutionary biology proposed in 1973, that species must constantly adapt, evolve, and proliferate in order to survive while pitted against ever-evolving opposing species. The hypothesis was intended t ...
. Because men generally exhibit uniform preference for neotenous women's faces, Elia (2013) questioned if women's varying preferences for neotenous men's faces could "help determine" the range of facial neoteny in humans.


Neoteny and its connection with human specialization features

Neoteny is not a ubiquitous trait of the human phenotype. Human expression timing, compared to chimpanzee, has a completely different trajectory uncovering that there is no uniform shift in developmental timing. Humans undergo this neotenous shift once sexual maturity is reached. A question prompted by the Mehmet Somel et al. study, is whether or not human-specific neotenic changes are indicative of human- specific cognitive traits. The tracking of where developmental landmarks occur in humans and other primates is a step towards a better understanding of how neoteny manifests specifically in our species and how it may contribute to our specialized features, such as smaller jaws. In humans, the neotenic shift is concentrated around a group of gray matter genes. This shift in neotenic genes also coincides with cortical reorganization that is related to synaptic elimination and is at a much more rapid pace over others during adolescence. It is also linked to the development of linguistic skills and the development of certain neurological disorders like ADHD.


Among primates and early humans

Delbert D. Thiessen said that ''Homo sapiens'' are more neotenized than ''Homo erectus'', ''Homo erectus'' were more neotenized than ''Australopithecus'',
Great Ape The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the east ...
s are more neotenized than
Old World monkey Old World monkey is the common English name for a family of primates known taxonomically as the Cercopithecidae (). Twenty-four genera and 138 species are recognized, making it the largest primate family. Old World monkey genera include baboons ...
s, and Old World monkeys are more neotenized than
New World monkey New World monkeys are the five families of primates that are found in the tropical regions of Mexico, Central and South America: Callitrichidae, Cebidae, Aotidae, Pitheciidae, and Atelidae. The five families are ranked together as the Ceboidea ( ...
s. Nancy Lynn Barrickman said that Brian T. Shea concluded by multivariate analysis that
Bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee and less often the dwarf chimpanzee or gracile chimpanzee, is an endangered great ape and one of the two species making up the genus '' Pan,'' the other being the comm ...
s are more neotenized than the
common chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of Hominidae, great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close r ...
, taking into account such features as the proportionately long torso length of the Bonobo. Montagu said that part of the differences seen in the morphology of "modernlike types of man" can be attributed to different rates of "neotenous mutations" in their early populations. Regarding behavioral neoteny, Mathieu Alemany Oliver says that neoteny partly (and theoretically) explains stimulus seeking, reality conflict, escapism, and control of aggression in consumer behavior. However, if these characteristics are more or less visible among people, Alemany Oliver argues, it is more the fact of cultural variables than the result of different levels of neoteny. Such a view makes behavioral neoteny play a non-significant role in gender and race differences, and puts an emphasis on culture.


Specific neotenies

Populations with a history of dairy farming have evolved to be lactose tolerant in adulthood, whereas other populations generally lose the ability to break down lactose as they grow into adults.
Down syndrome Down syndrome or Down's syndrome, also known as trisomy 21, is a genetic disorder caused by the presence of all or part of a third copy of chromosome 21. It is usually associated with physical growth delays, mild to moderate intellectual dis ...
neotenizes the brain and body. The syndrome is characterized by decelerated maturation (neoteny), incomplete morphogenesis (vestigia) and
atavism In biology, an atavism is a modification of a biological structure whereby an ancestral genetic trait reappears after having been lost through evolutionary change in previous generations. Atavisms can occur in several ways; one of which is when ...
s.
Dwarfism Dwarfism is a condition wherein an organism is exceptionally small, and mostly occurs in the animal kingdom. In humans, it is sometimes defined as an adult height of less than , regardless of sex; the average adult height among people with dw ...
and
achondroplasia Achondroplasia is a genetic disorder with an autosomal dominant pattern of inheritance whose primary feature is dwarfism. In those with the condition, the arms and legs are short, while the torso is typically of normal length. Those affected ha ...
also neotenize the size of the human height as well as the limbs. This is due to
dwarfing Dwarfing is a process in which a breed of animals or cultivar of plants is changed to become significantly smaller than standard members of their species. The effect can be induced through human intervention or non-human processes, and can include ...
in the
growth hormone deficiency Growth hormone deficiency (GHD), or human growth hormone deficiency, is a medical condition resulting from not enough growth hormone (GH). Generally the most noticeable symptom is that an individual attains a short height. Newborns may also presen ...
.


See also

*
Ageing Ageing ( BE) or aging ( AE) is the process of becoming older. The term refers mainly to humans, many other animals, and fungi, whereas for example, bacteria, perennial plants and some simple animals are potentially biologically immortal. In ...
*
Cuteness Cuteness is a subjective term describing a type of attractiveness commonly associated with youth and appearance, as well as a scientific concept and analytical model in ethology, first introduced by Konrad Lorenz. Lorenz proposed the concept o ...
*
Kawaii ''Kawaii'' is the culture of cuteness in Japan. It can refer to items, humans and non-humans that are charming, vulnerable, shy and childlike.Kerr, Hui-Ying (23 November 2016)"What is kawaii – and why did the world fall for the ‘cult of cu ...
*
Sexual selection Sexual selection is a mode of natural selection in which members of one biological sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of t ...


References

{{reflist, 30em Vertebrate developmental biology Evolutionary biology Taxonomy (biology) Human biology