Paternal care

From Wikipedia, the free encyclopedia

https://en.wikipedia.org/wiki/Paternal_care

In biology, paternal care is parental investment provided by a male to his own offspring. It is a complex social behaviour in vertebrates associated with animal mating systems, life history traits, and ecology. Paternal care may be provided in concert with the mother (biparental care) or, more rarely, by the male alone (so called exclusive paternal care).

The provision of care, by either males or females, is presumed to increase growth rates, quality, and/or survival of young, and hence ultimately increase the inclusive fitness of parents. In a variety of vertebrate species (e.g., about 80% of birds and about 6% of mammals), both males and females invest heavily in their offspring. Many of these biparental species are socially monogamous, so individuals remain with their mate for at least one breeding season.

Exclusive paternal care has evolved multiple times in a variety of organisms, including invertebrates, fishes, and amphibians.

Mammals

Pack of African wild dogs (Lycaon pictus) resting. A mammal in which males remain as care helpers.

Male mammals employ different behaviors to enhance their reproductive success (e.g. courtship displays, mate choice). However, the benefits of paternal care have rarely been studied in mammals, largely because only 5-10% of mammals exhibit such care (mostly present in primates, rodents and canids). In those species in which males provide extensive care for their offspring, indirect evidence suggests that its costs can be substantial. For example, mammalian fathers that care for their young may undergo changes in body mass and an increase in production of a number of costly hormones (e.g. androgens, glucocorticoids, leptin). Nonetheless, there is evidence that suggest that across all mammals, when males carry and groom their offspring their female partner fecundity increases, and if males provision the females, their litter size tend to be larger.

Humans

Main article: Father

Human cultures and societies vary widely in the expression of paternal care. Some cultures recognize paternal care via celebration of Father's Day. Human paternal care is a derived characteristic (evolved in humans or our recent ancestors) and one of the defining characteristics of Homo sapiens. Different aspects of human paternal care (direct, indirect, fostering social or moral development) may have evolved at different points in our history, and together they form a unique suite of behaviors as compared with the great apes. One study of humans has found evidence suggesting a possible evolutionary trade-off between mating success and parenting involvement; specifically, fathers with smaller testes tend to be more involved in care of their children.

Research on the effects of paternal care on human happiness have yielded conflicting results. However, one recent study concluded that fathers generally report higher levels of happiness, positive emotion, and meaning in life as compared with non-fathers.

According to the United States Census Bureau, approximately one third of children in the U.S. grow up without their biological father in their home. Numerous studies have documented negative consequences of being raised in a home that lacks a father, including increased likelihood of living in poverty, having behavioral problems, committing crimes, spending time in prison, abusing drugs or alcohol, becoming obese, and dropping out of school.

Non-human primates

In non-human primates, paternal investment is often dependent on the type of mating system exhibited by each species. Mating systems influence paternity certainty and the likelihood that a male is providing care towards his own biological offspring. Paternal certainty is high in monogamous pair-bonded species and males are less likely to be at risk for caring for unrelated offspring and not contributing to their own fitness. In contrast, polygamous primate societies create paternity uncertainty and males are more at risk of providing care for unrelated offspring and compromising their own fitness. Paternal care by male non-human primates motivated by biological paternity utilize past mating history and phenotypic matching in order to recognize their own offspring. Comparing male care efforts exhibited by the same species can provide insight on the significant relationship between paternity certainty and the amount of paternal care exhibited by a male. For example, Siamangs (Symphalangus syndactylus) utilize both polyandrous and monogamous mating systems but, it was found that monogamous males are more likely to carry infants and contribute to parental duties compared to those in promiscuous mating systems. Studies in Primatology have used primate mating systems and social organization to help theorize the evolutionary significance of paternal care in Primates.

Strepsirrhines

Ring tailed Lemur

Strepsirrhini is a suborder of the order Primates and includes lemurs, lorises, and bush babies. In this sub-order, males exhibit the lowest levels of paternal care for infants among primates. Examples of observed male care in this group include playing, grooming, and occasionally transporting infants. Males have also been observed interacting with infants while mothers park them and temporarily leave in order to feed. When female strepsirrhines park or nest their infants in nearby trees, males frequently use this as an opportunity to play with the unattended infants. In this suborder, male care and affection is directed toward multiple infants including non-biological offspring, and young strepsirrhines can be found interacting with various males. Paternal care does not influence infant growth rates or shorten inter-birth intervals of mothers as it can in haplorrhines. Strepsirrhini males exhibit the lowest intensity of care towards infants in non-human primates.

Strepsirrhines are constrained by their life history traits and reproductive rates are not flexible within this group of primates. This group of primates are programmed to give birth when food is abundant resulting in strict seasonal breeding periods. Shortening inter-birth intervals, which is theorized to be a possible outcome of increased male care, is not beneficial for Strepsirrhine mothers and can decrease infant survival. Studies also show that paternity can be highly skewed in Strepsirrhines, with only one or few male members being the only biological father within a single group. Instead of relying on a singular paternal figure, female mothers in this group rely on alloparenting from other group members. Infant parking and strict reproductive schedules are more beneficial for successful infant development in Strepsirrhines.

Haplorrhines

Chimpanzee infant

Haplorhini, a sub-order of the order Primate, includes tarsiers, New World Monkeys, Old World monkeys, apes, and humans. Haplorrhini is broken into two sister groups which are commonly distinguished by the characteristic of the primate nose: Catarrhini (narrow turned down nose) and Platyrrhini (flat nose). Paternal care is highly variable between the two sister groups and the species within them.

Catarrhines

Catarrhini is composed of Old World Monkeys (Cercopithecidae) and apes (Hylobatidae and Hominoidea). These primates are geographically located in Africa, Asia, and Madagascar.

Cercopithecines, the largest primate family, include primates species such as baboons, macaques, colobus, and vervet monkeys.

Apes consist of species of gibbons, siamangs, bonobos, chimpanzees, gorillas, orangutans and humans.

Catarrhines (non-human) are often organized into a multimale-multifemale social systems and utilize polygamous mating systems which results in paternity uncertainty. It is predicted that males in promiscuous mating systems do not engage in infant care due to the high costs of caring for an infant and missing opportunities to mate with receptive females. Male care in this group of primates is often portrayed through actions such as grooming, carrying, tolerance of the infant, as well as protection against agonistic interactions and infanticide. High ranking males can also provide access to food for developing infants. Direct care such as grooming and playing is not as common compared to male intervention on behalf of the infant when it is being harassed by conspecifics.

Baboon and her infant

In Cercopiths, male involvement in the infant's interactions with others is common in many species of baboons but between species paternal care is not always biased towards biological offspring. Male Savannah baboons (Papio cynocephalus) direct care towards their own biological offspring. Males in this species are more likely to intervene and protect infants from harassment against other group members when the infant is predicted to be their own. Studies have shown that male Savannah baboons selectively choose to remain in closer proximity to their own offspring and engage in long-term investment beyond early infancy, when the infant is at greatest risk for infanticide. Infants receiving paternal investment in Savannah baboons have shown enhanced fitness and accelerated maturation through males creating a safe zone for infants to exist in. Similarly to Savannah Baboons, Yellow baboon (Papio cynocephalus) males provide elevated care for their own offspring. Long-term care and investment beyond early infancy is better linked to paternity in this species and affecting infant growth and development. Male baboons also direct care towards unrelated offspring based on male affiliations with female mothers. Baboon males and females within a social group often exhibit “friendships” with females which begin during birth of her infant and has been observed to end abruptly if the infant dies. Males establish associations with females in which they have previously mated resulting in affiliative behaviour and protection towards her offspring. Relationships created by male and female members are significant for infant survival in Chacma baboons (Papio ursinus) because the risk of infanticide in early infancy is higher in this species. Paternal care in the form of protection for the infant is therefore more beneficial than long term investment in Chacma baboons and is believed to be directed towards both biological and non-biological infants in the group.

Rhesus Macaque

Similarly to baboons, paternal roles and the underlying mechanisms as to why paternal care evolved vary within macaque species. In Sulawesi crested macaques (Macaca nigra) both male rank and the relationship to the mother predicted male care towards an infant instead of true biological paternity. In both Sulawesi and Barbary macaques (Macaca sylvanus) males adopted a “care-then-mate” strategy, in which care is provided to infants regardless of paternity in order for the male to increase future mating opportunities with the mother. In both species, it was observed that male macaques are more likely to initiate care towards and positively interact with the infant in the presence of the mother. In Assamese macaques (Macaca assamensis) biological paternity was the most significant predictor of male affiliations with infants and therefore males biased care towards infants presumed to be their own. Observers found that Assamese males were more likely to engage and provide care for infants in the absence of their mothers reducing the likelihood that care provided to infants will impress the mother and secure access to mating possibilities. In Rhesus macaques, male's providing protection and greater access to food resulted in higher weight gain for both male and female infants. This had a positive effect on infant survival and was significant in the first year of infancy when the risk of infanticide is the highest.

Chimpanzees (Pan troglodytes) are organized into fission-fusion social groups and provide an example of a polygamous mating society. Male chimpanzees often engage with infants in the form of grooming, playing, and providing protection towards other group members. In both Western and Eastern chimpanzees it was found that males were more likely to engage with their own biological offspring meaning that male care is directed by paternity in this species. In both chimpanzee and bonobo social groups, high ranking alpha males sire approximately half of the offspring within their social group. More research needs to be done addressing how reproductive skew affects paternal care and infant-male relationships in non-human primates including chimpanzees and bonobos.

Platyrrhines

Titi Monkey

Platyrrhini is a sub-order of the order Primate and are commonly referred to as the New World Monkeys. These primates occupy Central and South America, and Mexico. This group is broken into five families, range in body size, and include species such as spider monkeys, capuchins, and howler monkeys.

Among primate species, the highest levels of male care found in New World monkeys are observed in Owl monkeys (Aotus azarai ) and Titi monkeys (Callicebus caligatus). In both of these species, males and females are monogamous, pair-bonded, and exhibit bi-parental care for their offspring. The social group in both these species consists of female and male parents along with their offspring. Males in these species serve as the primary caregivers and play a major role in infant survival.

Male Titi monkeys are more involved than the mother in all aspects of male care except nursing, and engage in more social activities such as grooming, food sharing, play, and transportation of the infant. The bond between an infant and its father is established right after birth and maintained into adolescence making the father the infant's predominant attachment figure. Similarly, the male Owl monkey acts as the main caregiver and is crucial to the survival of his offspring. If a female gives birth to twins, the male is still responsible for transporting both the infants. In the absence of a father, infant mortality increases in both these species and it is unlikely that the infant will survive. One study found that the replacement of a male enacting as the role of the father resulted in higher mortality during infancy emphasizing the importance of the social bond created between father and offspring at birth.

White-faced Capuchin

In White‐faced Capuchins (Cebus capucinus) one study found that parental care was exhibited in the form of playful behaviour, proximity to, inspection of, and collecting discarded food items from infants as determined by male rank and dominance status rather than biological relatedness to the infant. Scientists believe that future research on kin recognition needs to be done on capuchins to determine if males choose to bias their care as well as in other non-human primates relying on phenotypic matching to distinguish biological offspring.

Evolutionary Perspectives on Paternal Care in Primates

Squirrel monkeys

The Theory of Paternal Investment: Differences in infant care between sexes stems from females investing more time and energy in their offspring than males, while males compete with one another for access to females. Although paternal care is rare among mammalians, males across many primate species still play a paternal role in infant care.

Hypotheses addressing the rise of paternal care in several primate species

The Paternal Care hypothesis: Paternal care and investment will be designated to biological offspring, increasing the infant's chance of survival, and therefore increasing the male's own fitness. This hypothesis requires the on male to use recognition and behavioural cues to distinguish their own offspring from other infants. Paternal uncertainty is high in multimale-multifemale primate groups so males must use these cues to recognize and bias care towards their own offspring. This allows males to provide both short and long-term investment for infants. Primates living in monogamous pairs or single-male groups exhibit high paternity certainty and assist with the Paternal Care hypothesis.

The Mating Effort hypothesis: Males provide care for infants in order to increase mating opportunities with females. This means that males are more likely to engage in affiliative behaviours with the infant in the presence of the mother as a form of male mating effort in order to enhance future reproductive success. This theory is independent of genetics and evolved independent of paternity.

The Maternal Relief hypothesis: Males provide care infants to help reduce reproductive burdens of the female, ultimately resulting shorter inter-birth intervals and more successful offspring. This stems from the male alleviating the female from her parental duties in order to keep her resources from becoming depleted and subsequently allowing her to produce high quality milk for the infant. Similarly to the mating effort hypothesis, the maternal relief hypothesis is independent of genetics and does not require the male to be the biological father to take part in infant care.

Rodents

California mice (Peromyscus californicus) are well known for have intensive and sustained paternal behavior.

Several species of rodents have been studied as models of paternal care, including prairie voles (Microtus ochrogaster), Campbell's dwarf hamster, the Mongolian gerbil, and the African striped mouse. The California mouse (Peromyscus californicus) is a monogamous rodent that exhibits extensive and essential paternal care, and hence has been studied as a model organism for this phenomenon. One study of this species found that fathers had larger hindlimb muscles than did non-breeding males. Quantitative genetic analysis has identified several genomic regions that affect paternal care.

Birds

Main article: Parental care in birds

Fathers contribute equally with mothers to the care of offspring in as many as 90% of bird species, sometimes including incubating the eggs. Most paternal care is associated with biparental care in socially monogamous mating systems (about 81% of species), but in approximately 1% of species, fathers provide all care after eggs are laid. The unusually high incidence of paternal care in birds compared to other vertebrate taxa is often assumed to stem from the extensive resource requirements for production of flight-capable offspring. By contrast, in bats (the other extant flying vertebrate lineage), care of offspring is provided by females (although males may help guard pups in some species). In contrast to the large clutch sizes found in many bird species with biparental care, bats typically produce single offspring, which may be a limitation related to lack of male help. It has been suggested, though not without controversy, that paternal care is the ancestral form of parental care in birds.

Amphibians

Paternal care occurs in a number of species of anuran amphibians, including glass frogs.

Fish

According to the Encyclopedia of Fish Physiology: From Genome to Environment:

About 30% of the 500 known fish families show some form of parental care, and most often (78% of the time) care is provided by only one parent (usually the male). Male care (50%) is much more common than female care (30%) with biparental care accounting for about 20%, although a more recent comparative analysis suggests that male care may be more common (84%).

There are three common theoretical explanations for the high levels of paternal care in fish, with the third one currently favoured. First, external fertilization protects against paternity loss; however, sneaker tactics and strong sperm competition have evolved many times. Second, the earlier release of eggs than sperm gives females an opportunity to flee; however, in many paternal care species, eggs and sperm are released simultaneously. Third, if a male is already protecting a valuable spawning territory in order to attract females, defending young adds minimal parental investment, giving males a lower relative cost of parental care.

One well-known example of paternal care is in seahorses, where males brood the eggs in a brood pouch until they are ready to hatch.

Males from the Centrarchidae (sunfish) family exhibit paternal parental care of their eggs and fry through a variety of behaviors such as nest guarding and nest fanning (aerating eggs).

In jawfish, the female lays the eggs and the male then takes them in his mouth. A male can have up to 400 eggs in his mouth at one time. The male can't feed while he hosts the young, but as the young get older, they spend more time out of the mouth. This is sometimes termed mouthbrooding.

During the breeding season, male three-spined sticklebacks defend nesting territories. Males attract females to spawn in their nests and defend their breeding territory from intruders and predators. After spawning, the female leaves the male's territory and the male is solely responsible for the care of the eggs. During the ~6-day incubation period, the male 'fans' (oxygenates) the eggs, removes rotten eggs and debris, and defends the territory. Even after embryos hatch, father sticklebacks continue to tend their newly hatched offspring for ~7 days, chasing and retrieving fry that stray from the nest and spitting them back into the nest.

Arthropods

Paternal care is rare in arthropods, but occurs in some species, including the giant water bug and the arachnid Iporangaia pustulosa, a harvestman. In several species of crustaceans, males provide care of offspring by building and defending burrows or other nest sites. Exclusive paternal care, where males provide the sole investment after egg-laying, is the rarest form, and is known in only 13 taxa: giant water bugs, sea spiders, two genera of leaf-footed bugs, two genera of assassin bugs, three genera of phlaeothripid thrips, three genera of harvestmen, and in millipedes of the family Andrognathidae.

Theoretical models of the evolution of paternal care

Mathematical models related to the prisoner's dilemma suggest that when female reproductive costs are higher than male reproductive costs, males cooperate with females even when they do not reciprocate. In this view, paternal care is an evolutionary achievement that compensates for the higher energy demands that reproduction typically involves for mothers.

Other models suggest that basic life-history differences between males and females are adequate to explain the evolutionary origins of maternal, paternal, and bi-parental care. Specifically, paternal care is more likely if male adult mortality is high, and maternal care is more likely to evolve if female adult mortality is high. Basic life-history differences between the sexes can also cause evolutionary transitions among different sex-specific patterns of parental care.

Consequences for offspring survival and development

Care by fathers can have important consequences for survival and development of offspring in both humans and other species. Mechanisms underlying such effects may include protecting offspring from predators or environmental extremes (e.g., heat or cold), feeding them or, in some species, direct teaching of skills. Moreover, some studies indicate a potential epigenetic germline inheritance of paternal effects.

The effects of paternal care on offspring can be studied in various ways. One way is to compare species that vary in the degree of paternal care. For example, an extended duration of paternal care occurs in the gentoo penguin, as compared with other Pygoscelis species. It was found that their fledging period, the time between a chick's first trip to sea and its absolute independence from the group, was longer than other penguins of the same genus. The authors hypothesized that this was because it allowed chicks to better develop their foraging skills before becoming completely independent from their parents. By doing so, a chick may have a higher chance of survival and increase the population's overall fitness.

Proximate mechanisms

The proximate mechanisms of paternal care are not well understood for any organism. In vertebrates, at the level of hormonal control, vasopressin apparently underlies the neurochemical basis of paternal care; prolactin and testosterone may also be involved. As with other behaviors that affect Darwinian fitness, reward pathways in the brain may reinforce the expression of paternal care and may be involved in the formation of attachment bonds.

The mechanisms that underlie the onset of parental behaviors in female mammals have been characterized in a variety of species. In mammals, females undergo endocrine changes during gestation and lactation that "prime" mothers to respond maternally towards their offspring.

Paternal males do not undergo these same hormonal changes and so the proximate causes of the onset of parental behaviors must differ from those in females. There is little consensus regarding the processes by which mammalian males begin to express parental behaviors. In humans, evidence ties oxytocin to sensitive care-giving in both women and men, and with affectionate infant contact in women and stimulatory infant contact in men. In contrast, testosterone decreases in men who become involved fathers and testosterone may interfere with aspects of paternal care.

Placentophagia (the behavior of ingesting the afterbirth after parturition) has been proposed to have physiological consequences that could facilitate a male's responsiveness to offspring. Non-genomic transmission of paternal behavior from fathers to their sons has been reported to occur in laboratory studies of the biparental California mouse, but whether this involves (epigenetic) modifications or other mechanisms is not yet known.

Challenge hypothesis

From Wikipedia, the free encyclopedia
https://en.wikipedia.org/wiki/Challenge_hypothesis

The challenge hypothesis outlines the dynamic relationship between testosterone and aggression in mating contexts. It proposes that testosterone promotes aggression when it would be beneficial for reproduction, such as mate guarding, or strategies designed to prevent the encroachment of intrasexual rivals. The positive correlation between reproductive aggression and testosterone levels is seen to be strongest during times of social instability. The challenge hypothesis predicts that seasonal patterns in testosterone levels are a function of mating system (monogamy versus polygyny), paternal care, and male-male aggression in seasonal breeders.

The pattern between testosterone and aggression was first observed in seasonally breeding birds, where testosterone levels rise modestly with the onset of the breeding season to support basic reproductive functions. However, during periods of heightened male aggression, testosterone levels increase further to a maximum physiological level. This additional boost in testosterone appears to facilitate male-male aggression, particularly during territory formation and mate guarding, and is also characterized by a lack of paternal care. The challenge hypothesis has come to explain patterns of testosterone production as predictive of aggression across more than 60 species.

Patterns of testosterone

The challenge hypothesis presents a three-level model at which testosterone may be present in circulation. The first level (Level A) represents the baseline level of testosterone during the non-breeding season. Level A is presumed to maintain feedback regulation of both GnRH and gonadotropin release, which are key factors in testosterone production. The next level (Level B) is a regulated, seasonal breeding baseline. This level is sufficient for the expression of reproductive behaviors in seasonal breeders and the development of some secondary sex characteristics. Level B is induced by environmental cues, such as length of day. The highest level (Level C) represents the physiological testosterone maximum and is reached through social stimulation, such as male-male aggression. The challenge hypothesis proposes that social stimulation which leads to this rise in testosterone above breeding baseline serves to increase the frequency and intensity of aggression in males, particularly for competing with other males or interacting with sexually receptive females.

In birds

It is thought that testosterone plays an integral part of the territorial behavior within bird species, in particular the fluctuation of testosterone mitigated by luteinizing hormone (LH) during different seasons. Generally, mating behavior is demonstrated in the spring and accordingly, male birds show a sharp increase in LH as well as testosterone during this time. This acute rise in LH and testosterone can be attributed to the increased need for aggressive behaviors. The first need for aggressive behavior comes from the drive to establish territory. This typically occurs within the first few weeks of mating season. The second need for aggression occurs after the first clutch of eggs have been laid. The male not only needs to guard the eggs, but also to guard his sexually receptive mate from other potential suitors. Thus, the male adopts an “alpha male status” when acquiring territory as well as during the egg laying period. This alpha male status, as mentioned before, comes from the significant increase of testosterone that occurs during the mating season. Further evidence of LH and testosterone mitigating aggression in bird species comes from studies on bird species such as the song sparrow and the European blackbird who build highly accessible refuges, known as open cup nests.

Song sparrow

Because open cup nests can essentially be built anywhere, there is little competition when it comes to nest building sites. Accordingly, both the song sparrow and the European blackbird do not show an increase in luteinizing hormone or testosterone during territory acquisition . However, not all species of birds show increased levels of testosterone and LH during aggressive behavior. In a landmark study, it was found that male western screech owls, when exposed to another male during the non-mating season showed aggressive behavior without the increase in LH and testosterone. However, when the owls were put in a situation that warranted aggressive behavior during the mating season, there was a large spike in LH and testosterone during the aggressive act. This suggests that the mechanisms of aggressive behavior during the mating and non-mating seasons are independent of each other or perhaps the increase in testosterone somehow increases the aggressive response during the mating season. Estradiol (E2), a type of non gonadal estrogen, seems to play a key role in regulating aggressive behavior during the non-mating season in several species of birds. As previously noted, many bird species during the non-mating season have low testosterone levels yet still manage to display aggression. As a primary example, when the Washington State song sparrow, a bird which shows fairly high levels of aggression during non-mating season despite low testosterone, is exposed to fadrozole, an aromatase inhibitor, the levels of aggression are greatly decreased. When the E2 was replaced, the aggressive behaviors reestablished themselves thus confirming that E2 governs aggressive behavior during the non-mating season. It is unknown however if this is just specific to birds, or if this extends to other animal species.

These examples all culminate in a challenge hypothesis model which focuses on the role of testosterone on aggression in the breeding season. The challenge hypothesis most likely cannot be applied to the non-breeding season since, as mentioned above, there is most likely a mechanism independent of testosterone governing aggression in the non-mating season. A sigmoidal relationship between testosterone plasma levels and male-male aggression is observed under the challenge hypothesis when the birds’ testosterone levels were above seasonal breeding testosterone baseline levels. If birds remained at the seasonal breeding baseline levels during the breeding season, then there is not a significant difference observed in male-male aggression. In addition, there is a negative, sigmoidal relationship between testosterone levels in the birds and the amount of parental care provided when parents are above the seasonal breeding testosterone baseline levels. As such, the relationship between testosterone plasma levels and male-male aggression is context-specific to the species. Figure 2 and 3 describe the relationships observed of many single- or double-brooded bird species, from male western gulls to male turkeys.

In other animals

The challenge hypothesis has been used to describe the testosterone levels in other species to certain social stimuli. The challenge hypothesis predicts the testosterone influence on aggressive male-male interactions between male northern fence lizards. This reinforces the challenge hypothesis by showing rapid changes in aggressive behaviors of the lizards do not correlate with testosterone concentrations. Yet, over the mating season, the intensity of the behavior and the levels of testosterone levels yielded a positive correlation. Research has also shown the challenge hypothesis applies to specific monogamous fish species, with a greater correlation in species with stronger pair bonding.

In addition, the challenge hypothesis has been adapted to primate species. In 2004, Martin N. Muller and Richard W. Wrangham applied a modified challenge hypothesis to chimpanzees. Similar to the original hypothesis, they predicted that there would be increased male-male aggressive interaction when a receptive and fertile female chimpanzee was present. Muller and Wrangham also correctly predicted the testosterone levels of more dominant chimpanzees to be higher as compared to lower status chimpanzees. Therefore, chimpanzees significantly increased both testosterone levels and aggressive male-male interactions when receptive and fertile females presented sexual swellings. This study also highlighted how male testosterone and aggression levels rise only when males are in the presence of parous females. This is because nulliparous females are less attractive to males, and they are not guarded, meaning there is little competition. This evidence suggests that the increase in testosterone is related to only aggression – not sexual activity – as male chimpanzees mate equally with both parous and nulliparous females. Currently, no research has specified a relationship between the modified challenge hypothesis and human behavior, yet, many testosterone/human behavior studies support the modified hypothesis applying to human primates.

Cornerstones

Mating effort versus parenting effort

A fundamental feature of male life history is the tradeoff between the energy devoted to male-male competition and mate attraction (mating effort) versus that allocated to raising offspring (parenting effort). There is a trade off because decreased paternal effort caused by increased testosterone dramatically decreases reproductive success, due to decreased parental care and protection for the offspring. Therefore, to maximise reproductive success, the optimal balance between the two must be found. The challenge hypothesis proposes testosterone as the key physiological mechanism underlying this tradeoff. When the opportunity to reproduce arises—namely, the species enters the breeding season or females enter estrus—males should exhibit a rise in testosterone levels to facilitate sexual behavior. This will be characterized by increased mating effort and decreased parenting effort, as investment in the former may be incompatible with parental care due to insufficient time and energy to engage in all of these facets of reproductive effort.

Research on nonhuman species has found that testosterone levels are positively associated with mating effort and negatively related to parenting effort. Moreover, experimental manipulations have revealed a causal role of testosterone, such that elevations in testosterone result in increased mating effort and decreased parenting effort.

Paternal care

The challenge hypothesis makes different predictions regarding testosterone secretion for species in which males exhibit paternal care versus those in which males do not. When aggressive interactions among males arise in species that exhibit paternal care, testosterone levels are expected to be elevated. Males are predicted to exhibit an increase in testosterone to Level C (physiological maximum), but only during periods of territory establishment, male-male challenges, or when females are fertile so that paternal care is not compromised. When aggression is minimal, specifically during parenting, testosterone levels should decrease to Level B (breeding baseline). Level B represents the minimal levels of testosterone required for the expression of reproductive behaviors, and is not expected to drastically interfere with parenting behavior.

In species where males exhibit minimal to no paternal care, testosterone levels are hypothesized to be at Level C throughout the breeding season because of intense and continued interactions between males and the availability of receptive females. In polygynous species, where a single male tends to breed with more than one female, males generally do not exhibit a heightened endocrine response to challenges, because their testosterone levels are already close to physiological maximum throughout the breeding season. Experimental support for the relationship between heightened testosterone and polygyny was found, such that if testosterone was implanted into normally monogamous male birds (i.e., testosterone levels were manipulated to reach Level C) then these males became polygynous.

Mating Effort versus Maintenance

There is a broader trade-off to consider when it comes to the challenge hypothesis: maintenance vs reproductive effort. Reproductive effort includes both mating and parental effort. In order to gain the benefits of reproductive effort, individuals have to suffer the costs of testosterone, which can hinder their physiological maintenance. This is a form of life history tradeoff, due to the fact that natural selection favors reproductive success rather than maintenance. Therefore, the ability to find the correct balance between reproductive effort and maintenance would have been positively selected for by natural selection, leading to the physiological and social behaviour we now know as the challenge hypothesis.

One such cost is that increased aggressive activity due to high levels of testosterone is hypothesised to expose males to increased predation, which not only endangers them, but also their offspring. A study on the lizard Sceloporus jarrovi, supported this prediction, as those with induced high levels of testosterone for extended periods of time had a higher mortality rate than those with lower levels of testosterone. Prolonged high levels of testosterone have also been seen to suppress the immune system, with evidence ranging from human natural experiments to male-ring-tailed lemurs. Maintaining high levels of testosterone is energetically expensive, which can hinder reproductive success when a male frequently finds himself in aggressive and physically demanding situations. Due to increased aggression as a result of high testosterone levels, individuals expose themselves to higher injury risk than usual.

Therefore, the costs of maintaining a high testosterone level may outweigh increased reproductive success. A study on male ring-tailed lemurs (Lemur catta) supports the idea of a compromise between costs and benefits of increased testosterone levels, as increased levels were tightly timed around days of female estrus. This shows that there is an optimum length of time to have high testosterone levels when considering the costs and benefits.

Male-male aggression

It has long been known that testosterone increases aggressive behavior. While castration tends to decrease the frequency of aggression in birds and replacement therapy with testosterone increases aggression, aggression and testosterone are not always directly related. The challenge hypothesis proposes that testosterone is most immediately related to aggression when associated with reproduction, such as mate-guarding. An increase in male-male aggression in the reproductive context as related to testosterone is strongest in situations of social instability, or challenges from another male for a territory or access to mates.

The relationship between aggression and testosterone can be understood in light of the three-level model of testosterone as proposed by the challenge hypothesis. As testosterone reaches Level B, or breeding baseline, there is minimal increase in aggression. As testosterone increases above Level B and approaches Level C, male-male aggression rapidly increases.

Continuous breeders

The challenge hypothesis was established based upon data examining seasonal breeders. There are many species, however, who are continuous breeders—namely, species that breed year-round and whose mating periods are distributed throughout the year (e.g., humans). In continuous breeders, females are sexually receptive during estrus, at which time ovarian follicles are maturing and ovulation can occur. Evidence of ovulation, the phase during which conception is most probable, is advertised to males among many non-human primates via swelling and redness of the genitalia.

Support for the challenge hypothesis has been found in continuous breeders. For example, research on chimpanzees demonstrated that males became more aggressive during periods when females displayed signs of ovulation. Moreover, male chimpanzees engaged in chases and attacks almost 2.5 times more frequently when in groups containing sexually receptive females.

Implications for humans

The predictions of the challenge hypothesis as applied to continuous breeders partially rests upon males' ability to detect when females are sexually receptive. In contrast to females of many animal species who advertise when they are sexually receptive, human females do not exhibit cues but are said to conceal ovulation. While the challenge hypothesis has not been examined in humans, some have proposed that the predictions of the challenge hypothesis may apply.

Several lines of converging evidence in the human literature suggest that this proposition is plausible. For example, testosterone is lower in fathers as compared to non-fathers, and preliminary evidence suggests that men may be able to discern cues of fertility in women. The support for the challenge hypothesis in non-human animals provides a foundation for which to explore the relationship between testosterone and aggression in humans.

The Dual Hormone Hypothesis as an Extension of the Challenge Hypothesis

The challenge hypothesis claims that there is an association between testosterone and aggression in mating contexts, and more broadly status-seeking behaviours. However, findings linking testosterone to status-seeking behaviours, especially in humans, are often inconsistent and leave room for critique. In some cases, testosterone has been seen to positively correlate with status-seeking behaviours such as aggression and competitiveness, however, testosterone has also been found to have weak or even null correlations with the same behaviours. Some scholars blame these inconsistencies on limitations in study methods, but the dual-hormone hypothesis has emerged as a theoretical explanation to some of these inconsistencies.

Dual-Hormone Hypothesis

Stress plays a fundamental role in competition and mating, and therefore, the hormones released in response to stress should be considered as well as testosterone when looking at the challenge hypothesis. Cortisol is produced in the hypothalamic-pituitary-adrenal gland and is released when one is under physical or psychological stress; this is relevant to the challenge hypothesis as testosterone-linked status-seeking interactions are often stressful situations.

According to the dual-hormone hypothesis, the correlation between testosterone levels and aggression/status-seeking behaviour is reliant on corresponding cortisol levels; there is a strong correlation between the two when cortisol levels are low, and a weaker or sometimes reversed correlation when cortisol levels are high. There is supporting evidence for this relationship from a study done on humans, which looked at social status, leadership, and aggression. Cortisol is seen as a moderator of the relationship between testosterone and status-seeking/reproductive aggression in this hypothesis.

However, the dual-hormone hypothesis also has its own flaws, and current evidence appears to only partially support the hypothesis, according to a meta-analytical evaluation in 2019 by Dekkers et al. A proposed reasoning for the occasional weak evidence is that cortisol and testosterone, further interact with social context and individual psychology to regulate status-seeking behaviours. One such context is ‘victory-defeat’, where testosterone and cortisol will interact to increase desire to compete again more after losing than winning, as a method of regaining social status. Individual personality also has an effect on the interaction between cortisol and testosterone, and studies have shown that the cortisol x testosterone interaction was statistically significant for only those with high disagreeableness and high emotional instability.

Male warrior hypothesis

From Wikipedia, the free encyclopedia

https://en.wikipedia.org/wiki/Male_warrior_hypothesis

Ancient Hungarian warriors

The male warrior hypothesis (MWH) is an evolutionary psychology hypothesis by Professor Mark van Vugt which argues that human psychology has been shaped by between-group competition and conflict. Specifically, the evolutionary history of coalitional aggression between groups of men may have resulted in sex-specific differences in the way outgroups are perceived, creating ingroup vs. outgroup tendencies that are still observable today.

Overview

Violence and warfare

Violence and aggression are universal across human societies, and have likely been features of human behavior since prehistory. Archaeologists have found mass graves dating to the late Pleistocene and early Holocene that contain primarily male skeletons showing signs of blunt force trauma, indicating the cause of death was by weapons used in combat.

Violence among humans occurs in distinct patterns, differing most obviously by sex. Ethnographic findings and modern crime data indicate that the majority of violence is both perpetrated by and targeted at males, and males are the most likely to be victims of violence. This male-male pattern of violence has been observed so repeatedly and in so many cultures that it may qualify as a human universal.

Tribal behavior

Men preparing for a raid.

Humans are a social species with a long history of living in tribal groups. The psychological mechanisms that evolved to handle the complexities of group living have also created heuristics for quickly categorizing others as ingroup or outgroup members, with different behavioral strategies for each: treat ingroup members (those in one’s own group) favorably, and react to outgroup members (those who belong to a different group) with fear and aggression. These tendencies arise with little motivation, and have been provoked over superficial groups in lab studies—for example, by showing paintings to participants and creating groups based on which painting participants prefer.

The male warrior hypothesis suggests that the ease with which individuals discriminate against others is an adaptation resulting from a long history of being threatened by outgroup males, who are in competition for resources.

Sex differences in parental investment

The MWH argues that the sex differences in attitudes towards outgroup members may be a result of the different reproductive strategies used by males and females—specifically, the greater competition among males for mates. In mammals, males and females have distinct reproductive strategies based on the physiology of reproduction. Because females gestate, birth, feed, and invest more overall resources in each of their offspring, they are more selective with their mates but have greater certainty of being able to reproduce.

Males, in contrast, can mate at a very low energetic cost once they have found a partner, but are only able to attract a female if they have physical or social characteristics that can be converted into resources—e.g., territory, food resources, status, power, or influence—or the strength and alliances to coerce females to mate. As a result, there is typically much greater variability in the reproductive success of males within a species and higher competition among males for mates. The strongest, best adapted, and most powerful males may have a harem, while less fit males never reproduce.

For more details on this topic, see Trivers' theory of parental investment.

Male attitudes towards groups

The male warrior hypothesis predicts that because males may have historically remained in the groups in which they were born rather than moving away at adulthood (see patrilocality), they have a higher overall relatedness to their group than the female members, who would have moved to their new husbands’ group upon marriage. Males may have a stronger interest in defending their group, and will be more likely to act aggressively towards outgroup males they encounter who may be attempting to steal resources or weaken the group with violence.

For men at risk of never finding a mate, the fitness benefit to engaging in aggressive, violent behavior could outweigh the potential costs of fightings, especially if fighting alongside a coalition. Furthermore, the groups with more individuals who formed coalitions and acted altruistically to in-group members but aggressively to outgroup members would prosper.

Observational evidence/studies

Sex differences

Consistent with the expectations of the male warrior hypothesis, several studies have shown more ethnocentric and xenophobic beliefs and behaviors among men (compared to women), including the more frequent use of dehumanizing speech to describe outgroup members; stronger identification with their groups; greater cooperation when faced with competition from another group; a greater desire to engage in war when presented with images of attractive (but not unattractive) members of the opposite sex; greater overall rates of male-male competition and violence (as shown in violent crime and homicide statistics); and larger body size correlating with quicker anger responses.

Studies have also tested the responses of women to outgroups, and have shown that women are most likely to fear outgroup males during the periovulatory phase of the menstrual cycle, when fertility is at its peak. Women also have more negative responses around peak fertility when the males belong to an outgroup that the woman associates with physical formidability, even if the group was constructed in the lab. Overall, women who feel most at risk of sexual coercion are more likely to fear outgroup males, which aligns with the predictions of the MWH.

Prepared learning studies

In studies of prepared learning, conditioned fear responses to images of outgroup males were far more difficult to extinguish than conditioned fear responses to outgroup females or ingroup members of either sex, as measured by conductivity tests of perspiration on the skin. These results held true whether the participant was male or female. Because the neural circuitry for fear responses are more developed towards stimuli that have posed a larger threat for most of human history (snakes and spiders, for example, which were dangers frequently encountered by foragers), these findings suggest that outgroup males may have been more of a threat to physical safety than outgroup women or ingroup members, supporting the male warrior hypothesis.

Sport matches

It is hypothesized that sport began as a way for men to develop the skills needed in primitive hunting and warfare, and later developed to act primarily as a lek where male athletes display and male spectators evaluate the qualities of potential allies and rivals. This hypothesis is supported by the observation that the most popular modern male sports require the skills needed for success in male-male physical competition and primitive hunting and warfare, and that champion male athletes obtain high status and thereby reproductive opportunities in ways that parallel those gained by successful primitive hunters and warriors. There is evidence that male and female athletes generally differ in their motivation in sports, specifically their competitiveness and risk taking, in accordance with the spectator lek hypothesis.

The male warrior hypothesis proposes that men must engage in maximally effective intra-group cooperation. Post-conflict affiliation between opponents is proposed to facilitate future cooperation. Regarding sports matches as a proxy for intra-group conflict, a study found that unrelated human males are more predisposed than females to invest in post-conflict affiliation that is expected to facilitate future intra-group cooperation.

Non-human evidence

Coalitionary violence has also been observed in social species besides humans, including other primates. Chimpanzee (Pan troglodytes) males demonstrate similar violent behavior: groups of males form coalitions that patrol the borders of their territory and attack neighboring bands. Chimpanzees also have patrilocal living patterns, which aid with forming close coalitions, as all males are likely kin.

A study of 72 species of group-living mammals found that males are more involved than females in inter-group conflict where male fitness is limited by access to mates whereas female fitness is limited by access to food and safety.

Sex differences in emotional intelligence

From Wikipedia, the free encyclopedia

https://en.wikipedia.org/wiki/Sex_differences_in_emotional_intelligence
 

Emotional intelligence (EI) involves using cognitive and emotional abilities to function in interpersonal relationships, social groups as well as manage one's emotional states. It consists of abilities such as social cognition, empathy and also reasoning about the emotions of others.

The literature finds women have higher emotional intelligence ability than men based on common ability tests such as the MSCEIT. Physiological measures and behavioral tests also support this finding.

Emotional intelligence

Emotional intelligence (EI) involves using cognitive and emotional abilities to function in interpersonal relationships, social groups as well as manage one's emotional states. A person with high EI ability can perceive, comprehend and express emotion accurately, and also has the ability to access and generate feelings when needed to improve one's self and relationships with others.

Women tend to score higher than men on measures of emotional intelligence, but gender stereotypes of men and women can affect how they express emotions. The sex difference is small to moderate, somewhat inconsistent, and is often influenced by the person's motivations or social environment. Bosson et al. say "physiological measures of emotion and studies that track people in their daily lives find no consistent sex differences in the experience of emotion", which "suggests that women may amplify certain emotional expressions, or men may suppress them".

Tests

Mayer-Salovey-Caruso Emotional Intelligence Test (MSCEIT)

The Mayer-Salovey-Caruso Emotional Intelligence Test (MSCEIT) is used to get emotional intelligence IQs (EIQ). It is the most widely used test for the ability of emotional intelligence (AEI), and is well-validated. Much of the evidence for ability EI is based on the MSCEIT, partly because it was the only test available to measure EI ability. It is also the only omnibus test to measure all four branches of the EI ability model in one standardized assessment. The area scores include experiential EIQ and strategic EIQ. Experiential EIQ includes being able to recognize emotions to compare them to other sensations and their connection to the thought process. Strategic EIQ focuses on the meaning behind emotions, how emotions affect relationships, and how to manage emotions. After area scores, branch scores include four different sections: perceiving emotions, using emotions, understanding emotions, and managing emotions. Using these categories, the test analyzes people's ability to perform tasks and solve emotional problems or situations. No self-perceived assessments are used in the test; it is an objective assessment of a subject's ability to solve emotional problems.

A 2010 meta-analysis published in the Journal of Applied Psychology by researchers Dana L. Joseph and Daniel A. Newman found that women scored higher than men by around half a deviation, which amounts to 6–7 points difference.

Test of Emotional Intelligence (TIE)

The Test of Emotional Intelligence (TIE) focuses on measuring perception and comprehending emotions and the ability to use emotions and manage them. It is considered to be the Polish equivalent of the MSCEIT.

Sex differences

Social cognition

Every day, people use social cognition subconsciously, as it is part of most of modern society. Social cognition is an important part of emotional Intelligence and incorporates social skills such as processing facial expressions, body language and other social stimulus.

A 2012 review published in the journal Neuropsychologia found that men were more responsive to threatening cues while women could express themselves more easily and were better at recognizing others emotional states. A 2014 meta-analysis of 215 study samples by researchers Ashley E. Thompson and Daniel Voyer in the journal Cognition and Emotion found that there was "a small overall advantage in favour of females on emotion recognition tasks". Two 2015 reviews published in the journal Emotion review also found that adult women are more emotionally expressive, but that the size of this gender difference varies with the social and emotional context. Researchers distinguish three factors that predict the size of gender differences in emotional expressiveness: gender-specific norms, social role and situational constraints, and emotional intensity.

Empathy

See also: Sex differences in psychology § Empathy, and Empathy § Sex differences

A 2014 meta-analysis, in Cognition and Emotion, found overall female advantage in non-verbal emotional recognition.

A 2014 analysis from the journal Neuroscience & Biobehavioral Reviews also found that there are sex differences in empathy from birth, growing larger with age and which remains consistent and stable across lifespan. Females, on average, were found to have higher empathy than males at all ages, and children with higher empathy regardless of gender continue to possess high empathy throughout development in life. Further analysis of brain tools such as event related potentials found that females who viewed human suffering had higher ERP waveforms than males, an indication of greater empathetic response. Another investigation with similar brain tools such as N400 amplitudes found higher N400 in females in response to social situations which then positively correlated with self-reported empathy. Structural fMRI studies have also found females to have larger grey matter volumes in posterior inferior frontal and anterior inferior parietal cortex areas which have been correlated with mirror neurons indicated by the fMRI literature. Mirror neurons are crucial for many if not most aspects of empathy. Females were also found to have a stronger link between emotional and cognitive empathy. The researchers use The Primary Caretaker Hypothesis to explain the stability of these sex differences in development. According to the hypothesis, prehistoric males did not have the same selective pressure as women and this led to sex differences in emotion recognition and empathy.

Heritability of IQ

From Wikipedia, the free encyclopedia

https://en.wikipedia.org/wiki/Heritability_of_IQ

Research on the heritability of IQ inquires into the degree of variation in IQ within a population that is due to genetic variation between individuals in that population. There has been significant controversy in the academic community about the heritability of IQ since research on the issue began in the late nineteenth century. Intelligence in the normal range is a polygenic trait, meaning that it is influenced by more than one gene, and in the case of intelligence at least 500 genes. Further, explaining the similarity in IQ of closely related persons requires careful study because environmental factors may be correlated with genetic factors. Outside the normal range, certain single gene genetic disorders, such as phenylketonuria, can negatively affect intelligence.

Early twin studies of adult individuals have found a heritability of IQ between 57% and 73%, with some recent studies showing heritability for IQ as high as 80%. IQ goes from being weakly correlated with genetics for children, to being strongly correlated with genetics for late teens and adults. The heritability of IQ increases with the child's age and reaches a plateau at 14–16 years old, continuing at that level well into adulthood. However, poor prenatal environment, malnutrition and disease are known to have lifelong deleterious effects. Estimates in the academic research of the heritability of IQ have varied from below 0.5 to a high of 0.8 (where 1.0 indicates that monozygotic twins have no variance in IQ and 0 indicates that their IQs are completely uncorrelated). Eric Turkheimer and colleagues (2003) found that for children of low socioeconomic status heritability of IQ falls almost to zero. These results have been challenged by other researchers. IQ heritability increases during early childhood, but it is unclear whether it stabilizes thereafter. A 1996 statement by the American Psychological Association gave about 0.45 for children and about .75 during and after adolescence. A 2004 meta-analysis of reports in Current Directions in Psychological Science gave an overall estimate of around 0.85 for 18-year-olds and older. The general figure for heritability of IQ is about 0.5 across multiple studies in varying populations.

Although IQ differences between individuals have been shown to have a large hereditary component, it does not follow that disparities in IQ between groups have a genetic basis. The scientific consensus is that genetics does not explain average differences in IQ test performance between racial groups.

Heritability and caveats

Main article: Heritability

Heritability is a statistic used in the fields of breeding and genetics that estimates the degree of variation in a phenotypic trait in a population that is due to genetic variation between individuals in that population. The concept of heritability can be expressed in the form of the following question: "What is the proportion of the variation in a given trait within a population that is not explained by the environment or random chance?"

Estimates of heritability take values ranging from 0 to 1; a heritability estimate of 1 indicates that all variation in the trait in question is genetic in origin and a heritability estimate of 0 indicates that none of the variation is genetic. The determination of many traits can be considered primarily genetic under similar environmental backgrounds. For example, a 2006 study found that adult height has a heritability estimated at 0.80 when looking only at the height variation within families where the environment should be very similar. Other traits have lower heritability estimates, which indicate a relatively larger environmental influence. For example, a twin study on the heritability of depression in men estimated it as 0.29, while it was 0.42 for women in the same study.

Caveats

There are a number of points to consider when interpreting heritability:

• Heritability measures the proportion of variation in a trait that can be attributed to genes, and not the proportion of a trait caused by genes. Thus, if the environment relevant to a given trait changes in a way that affects all members of the population equally, the mean value of the trait will change without any change in its heritability (because the variation or differences among individuals in the population will stay the same). This has evidently happened for height: the heritability of stature is high, but average heights continue to increase. Thus, even in developed nations, a high heritability of a trait does not necessarily mean that average group differences are due to genes. Some have gone further, and used height as an example in order to argue that "even highly heritable traits can be strongly manipulated by the environment, so heritability has little if anything to do with controllability."
• A common error is to assume that a heritability figure is necessarily unchangeable. The value of heritability can change if the impact of environment (or of genes) in the population is substantially altered. If the environmental variation encountered by different individuals increases, then the heritability figure would decrease. On the other hand, if everyone had the same environment, then heritability would be 100%. The population in developing nations often has more diverse environments than in developed nations. This would mean that heritability figures would be lower in developing nations. Another example is phenylketonuria which previously caused intellectual disabilities in everyone who had this genetic disorder and thus had a heritability of 100%. Today, this can be prevented by following a modified diet, resulting in a lowered heritability.
• A high heritability of a trait does not mean that environmental effects such as learning are not involved. Vocabulary size, for example, is very substantially heritable (and highly correlated with general intelligence) although every word in an individual's vocabulary is learned. In a society in which plenty of words are available in everyone's environment, especially for individuals who are motivated to seek them out, the number of words that individuals actually learn depends to a considerable extent on their genetic predispositions and thus heritability is high.
• Since heritability increases during childhood and adolescence, and even increases greatly between 16 and 20 years of age and adulthood, one should be cautious drawing conclusions regarding the role of genetics and environment from studies where the participants are not followed until they are adults. Furthermore, there may be differences regarding the effects on the g-factor and on non-g factors, with g possibly being harder to affect and environmental interventions disproportionately affecting non-g factors.
• Contrary to popular belief, two parents of higher IQ will not necessarily produce offspring of equal or higher intelligence. Polygenic traits often appear less heritable at the extremes. A heritable trait is definitionally more likely to appear in the offspring of two parents high in that trait than in the offspring of two randomly selected parents. However, the more extreme the expression of the trait in the parents, the less likely the child is to display the same extreme as the parents. In fact, parents whose IQ is at either extreme are more likely to produce offspring with IQ closer to the mean (or average) than they are to produce offspring with high IQ. At the same time, the more extreme the expression of the trait in the parents, the more likely the child is to express the trait at all. For example, the child of two extremely tall parents is likely to be taller than the average person (displaying the trait), but unlikely to be taller than the two parents (displaying the trait at the same extreme). See also regression toward the mean.

Estimates

Various studies have estimated the heritability of IQ to be between 0.7 and 0.8 in adults and 0.45 in childhood in the United States. It has been found that estimates of heritability increase as individuals age. Heritability estimates in infancy are as low as 0.2, around 0.4 in middle childhood, and as high as 0.8 in adulthood. The brain undergoes morphological changes in development which suggests that age-related physical changes could contribute to this effect.

A 1994 article in Behavior Genetics based on a study of Swedish monozygotic and dizygotic twins found the heritability of the sample to be as high as 0.80 in general cognitive ability; however, it also varies by trait, with 0.60 for verbal tests, 0.50 for spatial and speed-of-processing tests, and 0.40 for memory tests. In contrast, studies of other populations estimate an average heritability of 0.50 for general cognitive ability.

In 2006, David Kirp, writing in The New York Times Magazine, summarized a century's worth of research as follows, "about three-quarters of I.Q. differences between individuals are attributable to heredity" while also highlighting that "much of what is labeled 'hereditary' becomes meaningful only in the context of experience." 

Shared family environment

See also: Environment and intelligence

There are some family effects on the IQ of children, accounting for up to a quarter of the variance. However, adoption studies show that by adulthood adoptive siblings aren't more similar in IQ than strangers, while adult full siblings show an IQ correlation of 0.24. However, some studies of twins reared apart (e.g. Bouchard, 1990) find a significant shared environmental influence, of at least 10% going into late adulthood. Judith Rich Harris suggests that this might be due to biasing assumptions in the methodology of the classical twin and adoption studies.

There are aspects of environments that family members have in common (for example, characteristics of the home). This shared family environment accounts for 0.25-0.35 of the variation in IQ in childhood. By late adolescence it is quite low (zero in some studies). There is a similar effect for several other psychological traits. These studies have not looked into the effects of extreme environments such as in abusive families.

The American Psychological Association's report "Intelligence: Knowns and Unknowns" (1996) asserts the necessity of a certain minimum level of responsible care for normal child development. Environments that are severely deprived, neglectful, or abusive negatively affect various developmental aspects, including intellectual growth. Beyond this minimum threshold, the influence of family experience on child development is contentious. Variables such as home resources and parents' use of language are correlated with children's IQ scores; however, these correlations may be influenced by genetic as well as environmental factors. The extent to which variance in IQ results from differences between families, compared to the varying experiences of different children within the same family, is a subject of debate. Recent twin and adoption studies indicate that the effect of the shared family environment is significant in early childhood but diminishes substantially by late adolescence. These findings suggest that differences in family lifestyles, while potentially important for many aspects of children's lives, have little long-term impact on the skills measured by intelligence tests.

Non-shared family environment and environment outside the family

Although parents treat their children differently, such differential treatment explains only a small amount of non-shared environmental influence. One suggestion is that children react differently to the same environment due to different genes. More likely influences may be the impact of peers and other experiences outside the family. For example, siblings grown up in the same household may have different friends and teachers and even contract different illnesses. This factor may be one of the reasons why IQ score correlations between siblings decreases as they get older.

Malnutrition and diseases

Certain single-gene metabolic disorders can severely affect intelligence. Phenylketonuria is an example, with publications documenting the capacity of treated phenylketonuria to produce a reduction of 10 IQ points on average. Meta-analyses have found that environmental factors, such as iodine deficiency, can result in large reductions in average IQ; iodine deficiency has been shown to produce a reduction of 12.5 IQ points on average.

Heritability and socioeconomic status

The APA report "Intelligence: Knowns and Unknowns" (1996) also stated that:

"We should note, however, that low-income and non-white families are poorly represented in existing adoption studies as well as in most twin samples. Thus it is not yet clear whether these studies apply to the population as a whole. It remains possible that, across the full range of income and ethnicity, between-family differences have more lasting consequences for psychometric intelligence."

A study (1999) by Capron and Duyme of French children adopted between the ages of four and six examined the influence of socioeconomic status (SES). The children's IQs initially averaged 77, putting them near retardation. Most were abused or neglected as infants, then shunted from one foster home or institution to the next. Nine years later after adoption, when they were on average 14 years old, they retook the IQ tests, and all of them did better. The amount they improved was directly related to the adopting family's socioeconomic status. "Children adopted by farmers and laborers had average IQ scores of 85.5; those placed with middle-class families had average scores of 92. The average IQ scores of youngsters placed in well-to-do homes climbed more than 20 points, to 98."

Stoolmiller (1999) argued that the range of environments in previous adoption studies was restricted. Adopting families tend to be more similar on, for example, socio-economic status than the general population, which suggests a possible underestimation of the role of the shared family environment in previous studies. Corrections for range restriction to adoption studies indicated that socio-economic status could account for as much as 50% of the variance in IQ.

On the other hand, the effect of this was examined by Matt McGue and colleagues (2007), who wrote that "restriction in range in parent disinhibitory psychopathology and family socio-economic status had no effect on adoptive-sibling correlations [in] IQ"

Turkheimer and colleagues (2003) argued that the proportions of IQ variance attributable to genes and environment vary with socioeconomic status. They found that in a study on seven-year-old twins, in impoverished families, 60% of the variance in early childhood IQ was accounted for by the shared family environment, and the contribution of genes is close to zero; in affluent families, the result is almost exactly the reverse.

In contrast to Turkheimer (2003), a study by Nagoshi and Johnson (2005) concluded that the heritability of IQ did not vary as a function of parental socioeconomic status in the 949 families of Caucasian and 400 families of Japanese ancestry who took part in the Hawaii Family Study of Cognition.

Asbury and colleagues (2005) studied the effect of environmental risk factors on verbal and non-verbal ability in a nationally representative sample of 4-year-old British twins. There was not any statistically significant interaction for non-verbal ability, but the heritability of verbal ability was found to be higher in low-SES and high-risk environments.

Harden, Turkheimer, and Loehlin (2007) investigated adolescents, most 17 years old, and found that, among higher income families, genetic influences accounted for approximately 55% of the variance in cognitive aptitude and shared environmental influences about 35%. Among lower income families, the proportions were in the reverse direction, 39% genetic and 45% shared environment."

In the course of a substantial review, Rushton and Jensen (2010) criticized the study of Capron and Duyme, arguing their choice of IQ test and selection of child and adolescent subjects were a poor choice because this gives a relatively less hereditable measure. The argument here rests on a strong form of Spearman's hypothesis, that the hereditability of different kinds of IQ test can vary according to how closely they correlate to the general intelligence factor (g); both the empirical data and statistical methodology bearing on this question are matters of active controversy.

A 2011 study by Tucker-Drob and colleagues reported that at age 2, genes accounted for approximately 50% of the variation in mental ability for children being raised in high socioeconomic status families, but genes accounted for negligible variation in mental ability for children being raised in low socioeconomic status families. This gene–environment interaction was not apparent at age 10 months, suggesting that the effect emerges over the course of early development.

A 2012 study based on a representative sample of twins from the United Kingdom, with longitudinal data on IQ from age two to age fourteen, did not find evidence for lower heritability in low-SES families. However, the study indicated that the effects of shared family environment on IQ were generally greater in low-SES families than in high-SES families, resulting in greater variance in IQ in low-SES families. The authors noted that previous research had produced inconsistent results on whether or not SES moderates the heritability of IQ. They suggested three explanations for the inconsistency. First, some studies may have lacked statistical power to detect interactions. Second, the age range investigated has varied between studies. Third, the effect of SES may vary in different demographics and different countries.

Maternal (fetal) environment

A meta-analysis by Devlin and colleagues (1997) of 212 previous studies evaluated an alternative model for environmental influence and found that it fits the data better than the 'family-environments' model commonly used. The shared maternal (fetal) environment effects, often assumed to be negligible, account for 20% of covariance between twins and 5% between siblings, and the effects of genes are correspondingly reduced, with two measures of heritability being less than 50%. They argue that the shared maternal environment may explain the striking correlation between the IQs of twins, especially those of adult twins that were reared apart. IQ heritability increases during early childhood, but whether it stabilizes thereafter remains unclear. These results have two implications: a new model may be required regarding the influence of genes and environment on cognitive function; and interventions aimed at improving the prenatal environment could lead to a significant boost in the population's IQ.

Bouchard and McGue reviewed the literature in 2003, arguing that Devlin's conclusions about the magnitude of heritability is not substantially different from previous reports and that their conclusions regarding prenatal effects stands in contradiction to many previous reports. They write that:

Chipuer et al. and Loehlin conclude that the postnatal rather than the prenatal environment is most important. The Devlin et al. (1997a) conclusion that the prenatal environment contributes to twin IQ similarity is especially remarkable given the existence of an extensive empirical literature on prenatal effects. Price (1950), in a comprehensive review published over 50 years ago, argued that almost all MZ twin prenatal effects produced differences rather than similarities. As of 1950 the literature on the topic was so large that the entire bibliography was not published. It was finally published in 1978 with an additional 260 references. At that time Price reiterated his earlier conclusion (Price, 1978). Research subsequent to the 1978 review largely reinforces Price's hypothesis (Bryan, 1993; Macdonald et al., 1993; Hall and Lopez-Rangel, 1996; see also Martin et al., 1997, box 2; Machin, 1996).

Dickens and Flynn model

Dickens and Flynn (2001) argued that the "heritability" figure includes both a direct effect of the genotype on IQ and also indirect effects where the genotype changes the environment, in turn affecting IQ. That is, those with a higher IQ tend to seek out stimulating environments that further increase IQ. The direct effect can initially have been very small but feedback loops can create large differences in IQ. In their model an environmental stimulus can have a very large effect on IQ, even in adults, but this effect also decays over time unless the stimulus continues. This model could be adapted to include possible factors, like nutrition in early childhood, that may cause permanent effects.

The Flynn effect is the increase in average intelligence test scores by about 0.3% annually, resulting in the average person today scoring 15 points higher in IQ compared to the generation 50 years ago. This effect can be explained by a generally more stimulating environment for all people. Some scientists have suggested that such enhancements are due to better nutrition, better parenting and schooling, as well as exclusion of the least intelligent people from reproduction. However, Flynn and a group of other scientists share the viewpoint that modern life implies solving many abstract problems which leads to a rise in their IQ scores.

Influence of genes on IQ stability

Recent research has illuminated genetic factors underlying IQ stability and change. Genome-wide association studies have demonstrated that the genes involved in intelligence remain fairly stable over time. Specifically, in terms of IQ stability, "genetic factors mediated phenotypic stability throughout this entire period [age 0 to 16], whereas most age-to-age instability appeared to be due to non-shared environmental influences". These findings have been replicated extensively and observed in the United Kingdom, the United States, and the Netherlands. Additionally, researchers have shown that naturalistic changes in IQ occur in individuals at variable times.

Influence of parents genes that are not inherited

Kong reports that, "Nurture has a genetic component, i.e. alleles in the parents affect the parents' phenotypes and through that influence the outcomes of the child." These results were obtained through a meta-analysis of educational attainment and polygenic scores of non-transmitted alleles. Although the study deals with educational attainment and not IQ, these two are strongly linked.

Spatial ability component of IQ

Spatial ability has been shown to be unifactorial (a single score accounts well for all spatial abilities), and is 69% heritable in a sample of 1,367 pairs of twins from the ages 19 through 21. Further only 8% of spatial ability can be accounted for by shared environmental factors like school and family. Of the genetically determined portion of spatial ability, 24% is shared with verbal ability (general intelligence) and 43% was specific to spatial ability alone.

Molecular genetic investigations

A 2009 review article identified over 50 genetic polymorphisms that have been reported to be associated with cognitive ability in various studies, but noted that the discovery of small effect sizes and lack of replication have characterized this research so far. Another study attempted to replicate 12 reported associations between specific genetic variants and general cognitive ability in three large datasets, but found that only one of the genotypes was significantly associated with general intelligence in one of the samples, a result expected by chance alone. The authors concluded that most reported genetic associations with general intelligence are probably false positives brought about by inadequate sample sizes. Arguing that common genetic variants explain much of the variation in general intelligence, they suggested that the effects of individual variants are so small that very large samples are required to reliably detect them. Genetic diversity within individuals is heavily correlated with IQ.

A novel molecular genetic method for estimating heritability calculates the overall genetic similarity (as indexed by the cumulative effects of all genotyped single nucleotide polymorphisms) between all pairs of individuals in a sample of unrelated individuals and then correlates this genetic similarity with phenotypic similarity across all the pairs. A study using this method estimated that the lower bounds for the narrow-sense heritability of crystallized and fluid intelligence are 40% and 51%, respectively. A replication study in an independent sample confirmed these results, reporting a heritability estimate of 47%. These findings are compatible with the view that a large number of genes, each with only a small effect, contribute to differences in intelligence.

Correlations between IQ and degree of genetic relatedness

The relative influence of genetics and environment for a trait can be calculated by measuring how strongly traits covary in people of a given genetic (unrelated, siblings, fraternal twins, or identical twins) and environmental (reared in the same family or not) relationship. One method is to consider identical twins reared apart, with any similarities that exist between such twin pairs attributed to genotype. In terms of correlation statistics, this means that theoretically the correlation of tests scores between monozygotic twins would be 1.00 if genetics alone accounted for variation in IQ scores; likewise, siblings and dizygotic twins share on average half alleles and the correlation of their scores would be 0.50 if IQ were affected by genes alone (or greater if there is a positive correlation between the IQs of spouses in the parental generation). Practically, however, the upper bound of these correlations are given by the reliability of the test, which is 0.90 to 0.95 for typical IQ tests.

If there is biological inheritance of IQ, then the relatives of a person with a high IQ should exhibit a comparably high IQ with a much higher probability than the general population. In 1982, Bouchard and McGue reviewed such correlations reported in 111 original studies in the United States. The mean correlation of IQ scores between monozygotic twins was 0.86, between siblings 0.47, between half-siblings 0.31, and between cousins 0.15.

The 2006 edition of Assessing adolescent and adult intelligence by Alan S. Kaufman and Elizabeth O. Lichtenberger reports correlations of 0.86 for identical twins raised together compared to 0.76 for those raised apart and 0.47 for siblings. These numbers are not necessarily static. When comparing pre-1963 to late 1970s data, researchers DeFries and Plomin found that the IQ correlation between parent and child living together fell significantly, from 0.50 to 0.35. The opposite occurred for fraternal twins.

Every one of these studies presented next contains estimates of only two of the three factors which are relevant. The three factors are G, E, and GxE. Since there is no possibility of studying equal environments in a manner comparable to using identical twins for equal genetics, the GxE factor can not be isolated. Thus the estimates are actually of G+GxE and E. Although this may seem like nonsense, it is justified by the unstated assumption that GxE=0. It is also the case that the values shown below are r correlations and not r(squared), proportions of variance. Numbers less than one are smaller when squared. The next to last number in the list below refers to less than 5% shared variance between a parent and child living apart.

Another summary:

• Same person (tested twice over time) .85 or above
• Identical twins—Reared together .86
• Identical twins—Reared apart .76
• Fraternal twins—Reared together .55
• Fraternal twins—Reared apart .35
• Biological siblings—Reared together .47
• Biological siblings—Reared apart .24
• Biological siblings—Reared together—Adults .24
• Unrelated children—Reared together—Children .28
• Unrelated children—Reared together—Adults .04
• Cousins .15
• Parent-child—Living together .42
• Parent-child—Living apart .22
• Adoptive parent–child—Living together .19

Between-group heritability

Main article: Race and intelligence

In the US, individuals identifying themselves as Asian generally tend to score higher on IQ tests than Caucasians, who tend to score higher than Hispanics, who tend to score higher than African Americans. Yet, although IQ differences between individuals have been shown to have a large hereditary component, it does not follow that between-group differences in average IQ have a genetic basis. In fact, greater variation in IQ scores exists within each ethnic group than between them. The scientific consensus is that genetics does not explain average differences in IQ test performance between racial groups. Growing evidence indicates that environmental factors, not genetic ones, explain the racial IQ gap.

Arguments in support of a genetic explanation of racial differences in average IQ are sometimes fallacious. For instance, some hereditarians have cited as evidence the failure of known environmental factors to account for such differences, or the high heritability of intelligence within races. Jensen and Rushton, in their formulation of Spearman's Hypothesis, argued that cognitive tasks that have the highest g-load are the tasks in which the gap between black and white test takers is greatest, and that this supports their view that racial IQ gaps are in large part genetic. However, in separate reviews, Mackintosh, Nisbett et al. and Flynn have all concluded that the slight correlation between g-loading and the test score gap offers no clue to the cause of the gap. Further reviews of both adoption studies and racial admixture studies have also found no evidence for a genetic component behind group-level IQ differences. Hereditarian arguments for racial differences in IQ have been criticized from a theoretical point of view as well. For example, the geneticist and neuroscientist Kevin Mitchell has argued that "systematic genetic differences in intelligence between large, ancient populations" are "inherently and deeply implausible" because the "constant churn of genetic variation works against any long-term rise or fall in intelligence." As he argues, "To end up with systematic genetic differences in intelligence between large, ancient populations, the selective forces driving those differences would need to have been enormous. What's more, those forces would have to have acted across entire continents, with wildly different environments, and have been persistent over tens of thousands of years of tremendous cultural change."

In favor of an environmental explanation, on the other hand, numerous studies and reviews have shown promising results. Among these, some focus on the gradual closing of the black–white IQ gap over the last decades of the 20th century, as black test-takers increased their average scores relative to white test-takers. For instance, Vincent reported in 1991 that the black–white IQ gap was decreasing among children, but that it was remaining constant among adults. Similarly, a 2006 study by Dickens and Flynn estimated that the difference between mean scores of black people and white people closed by about 5 or 6 IQ points between 1972 and 2002, a reduction of about one-third. In the same period, the educational achievement disparity also diminished. Reviews by Flynn and Dickens, Mackintosh, and Nisbett et al. all accept the gradual closing of the gap as a fact. Other recent studies have focused on disparities in nutrition and prenatal care, as well as other health-related environmental disparities, and have found that these disparities may account for significant IQ gaps between population groups. Still other studies have focused on educational disparities, and have found that intensive early childhood education and test preparation can diminish or eliminate the black–white IQ test gap. In light of these and similar findings, a consensus has formed that genetics does not explain differences in average IQ test performance between racial groups.