Parental investment

This article is about parental care in animals. For human parenting, see parenting.
A female calliope hummingbird feeding her chicks
Ronald Fisher
A crab-eating macaque nursing her offspring
A human mother feeding her child

Parental investment (PI), in evolutionary biology and evolutionary psychology, is any parental expenditure (time, energy etc.) that benefits one offspring at a cost to parents' ability to invest in other components of fitness,[1][2] and is thus a form of sexual selection. Components of fitness[3] include the wellbeing of existing offspring, parents' future sexual reproduction, and inclusive fitness through aid to kin.[4] Parental investment may be performed by both the male and female (biparental care), the mother alone (exclusive maternal care) or the father alone (exclusive paternal care).

Initially introduced in 1930 by the English biologist and statistician Ronald Fisher, parental care is found in a broad range of taxonomic groups, including both ectothermic (invertebrates, fish, amphibians and reptiles), and endothermic (birds and mammals) species. Care can be provided at any stage of the offspring's life: pre-natal care including behaviours such as egg guarding, preparation of nest, brood carrying, incubation, and placental nourishment in mammals; and post-natal care including food provisioning and protection of offspring.

An alternative to Fisher's theory is Trivers' parental investment theory (PIT), in which he focuses on how parental investment in humans affects their sexual behaviour.

Parental care

Parental investment theory is a branch of life history theory. The earliest consideration of parental investment is given by Ronald Fisher in his 1930 book The Genetical Theory of Natural Selection,[5] wherein Fisher argued that parental expenditure on both sexes of offspring should be equal. Clutton-Brock expanded the concept of PI to include costs to any other component of parental fitness.

A cellar spider defending spiderlings.

Reproduction is costly. Individuals are limited in the degree to which they can devote time and resources to producing and raising their young, and such expenditure may also be detrimental to their future condition, survival, and further reproductive output. However, such expenditure is typically beneficial to the offspring, enhancing their condition, survival, and reproductive success. These differences may lead to parent-offspring conflict. Parental investment can be provided by the female (female uniparental care), the male (male uniparental care), or both (biparental care). Parents are naturally selected to maximize the difference between the benefits and the costs, and parental care will tend to exist when the benefits are substantially greater than the costs.

Penguins are a prime example of a species that drastically sacrifices their own health and well-being in exchange for the survival of their offspring and the overall fitness of the population. This altruistic behavior, one that does not necessarily benefit the individual, but the population as a whole, can be seen in the King Penguin. Although some animals do exhibit altruistic behaviors towards individuals that are not of direct relation, many of these behaviors appear mostly in parent-offspring relationships. While breeding, males remain in a fasting-period at the breeding site for five weeks, waiting for the female to return for her own incubation shift. However, during this time period, males may decide to abandon their egg if the female is delayed in her return to the breeding grounds. This is an interesting case, as it shows that these penguins initially show a trade-off of their own health, in hopes of increasing the survivorship of their egg, but there comes a point where the male penguin's costs become too high in comparison to the gain of a successful breeding season. In a study, Olof Olsson investigated the correlation between how many experiences in breeding an individual has and the duration an individual will wait until abandoning his egg. He proposed that the more experienced the individual, the better that individual will be at replenishing his exhausted body reserves, allowing him to remain at the egg for a longer period of time. The males' sacrifice of their body weight and possible survivorship, in order to increase their offspring's chance of survival is a trade-off between current reproductive success and the parents' future survival.[6] This trade-off makes sense with other examples of kin-based altruism and is a clear example of the use of altruism in an attempt to increase overall fitness of a population at the expense of the individual's fitness.

A study found that male dunnocks tend to not discriminate between their own young and those of another male in polyandrous or polygynandrous systems. However, they increase their own reproductive success through feeding the offspring in relation to their own access to the female throughout the mating period, which is generally a good predictor of paternity.[7] This indiscriminative parental care by males is also observed in redlip blennies.[8]

In iteroparous species, where individuals may go through several reproductive bouts during their lifetime, a tradeoff may exist between investment in current offspring and future reproduction. Parents need to balance their offspring's demands against their own self-maintenance. This potential negative effect of parental care was explicitly formalized by Trivers in 1972, who originally defined the term parental investment to mean any investment by the parent in an individual offspring that increases the offspring's chance of surviving (and hence reproductive success) at the cost of the parent's ability to invest in other offspring.[2]

A female lizard defending her clutch against an egg-eating snake.

The benefits of parental investment to the offspring are large and are associated with the effects on condition, growth, survival, and ultimately on reproductive success of the offspring. For example, in the cichlid fish Tropheus moorii, a female has very high parental investment in her young because she mouthbroods the young and while mouthbrooding, all nourishment she takes in goes to feed the young and she effectively starves herself. In doing this, her young are larger, heavier, and faster than they would have been without it. These benefits are very advantageous since it lowers their risk of being eaten by predators and size is usually the determining factor in conflicts over resources.[9] However, such benefits can come at the cost of parent's ability to reproduce in the future e.g., through increased risk of injury when defending offspring against predators, loss of mating opportunities whilst rearing offspring, and an increase in the time interval until the next reproduction.

A special case of parental investment is when young do need nourishment and protection, but the genetic parents do not actually contribute in the effort to raise their own offspring. For example, in Bombus terrestris, oftentimes sterile female workers will not reproduce on their own, but will raise their mother's brood instead. This is common in social Hymenoptera due to haplodiploidy, whereby males are haploid and females are diploid. This ensures that sisters are more related to each other than they ever would be to their own offspring, incentivizing them to help raise their mother's young over their own.[10]

Overall, parents are selected to maximize the difference between the benefits and the costs, and parental care will be likely to evolve when the benefits exceed the costs.

In animals with small brains

Extensive parental investment is known in a wide range of small-brained animals, including some species of fish, amphibians and insects. Therefore, the bigger brains in mammals compared to most other animals cannot be due to a minimum parental care brain size. The fact that big brains consume more nutrients than smaller brains makes this especially relevant for evolutionary theories of the brain.[11][12]

Offspring and situation direction

Parental care only requires behaviors that effectively enhances offspring's chances of survival, it does not require underlying mechanisms to be potentially at a continuum with generalizable empathy appliceable to adults, nor even other situations involving young than the specific reactions, nor does it require the offspring to be altruistic back in any way.[13][14] Total ungeneralizability to adults has the evolvability advantage of not making parentally caring individuals vulnerable to being exploited by other adults.

Sexual selection

In many species, sexual selection is closely linked to parental investment. In theory, a male from such a species can produce a large number of offspring over the course of his life by minimizing parental investment in favor of investing his time instead impregnating any reproductive-age female who is fertile. In contrast, a female from said species typically can have a much smaller number of offspring during her reproductive life, partly due to an obligatory non-nil parental investment (i.e., gestation and delivery). This suggests that females will be more selective ("choosy") of mates than males will be, choosing males with good fitness (e.g., genes, high status, resources, etc.), so as to help offset any lack of direct parental investment from the male, and therefore increase reproductive success. Robert Trivers' theory of parental investment predicts that the sex making the largest investment in lactation, nurturing, and protecting offspring will be more discriminating in mating; and that the sex that invests less in offspring will compete for access to the higher-investing sex (see Bateman's principle[15]).

In species where both sexes invest highly in parental care, mutual choosiness is expected to arise. An example of this is seen in crested auklets, where parents share equal responsibility in incubating their single egg and raising the chick. In crested auklets both sexes are ornamented.[16]

The importance of parental investment can be seen especially in species in which the offspring are altricial (i.e., unable to fend for themselves from earliest ages). In many bird species and in modern humans, this leads to males spending more time caring for their offspring than do the male parents of precocial species, since reproductive success would otherwise suffer. The higher parental investment in humans is the result of an extended childhood required in order to develop the unusually large human brain. During this time, parental investment comes in the form of feeding, teaching, and protection, for example.

However, if human parental investment starts from the point when the sperm fertilizes the egg, then the minimal obligatory parental investment for the male is zero effort, while the minimal obligatory investment for the female is nine months of pregnancy, followed by delivery. This difference of minimal obligatory investment between males and females suggests that the amount of investment and effort put into mating and parenting will differ.

In some insects male parental investment is given in the form of a nuptial gift. For instance, ornate moth females receive a spermatophore containing nutrients, sperm and defensive toxins from the male during copulation. This gift, which can account for up to 10% of the male's body mass, constitutes the total parental investment the male provides.[17]

Trivers' parental investment theory

Parental investment as defined by Trivers in 1972[18] is the investment in offspring by the parent that increases the offspring's chances of surviving and hence reproductive success at the expense of the parent's ability to invest in other offspring. A large parental investment largely decreases the parents' chances of investing in other offspring. Parental investment can be split into two main categories: mating investment and rearing investment. Mating investment consist of the sexual act and the sex cells invested. The rearing investment is the time and energy expended to raise the offspring after conception. Women's parental investment in both mating and rearing efforts greatly surpasses that of the male. In terms of sex cells (egg and sperms cells), the female's investment is a lot larger, while males produce thousands of sperm cells which are supplied at a rate of twelve million per hour.[19] Women have a fixed supply of around 400 ova. Also, the acts of fertilization and gestation occur in the women, which compared to the male's investment of just one cell outweighs it. Furthermore, each intercourse could result in a nine-month commitment such as gestation (the act of breastfeeding) for the woman. From Trivers' theory of parental investment several implications follow. The first is that women are often but not always the more investing sex. The fact that they are more investing sex has meant that evolution has favoured females who are more selective of their mates to ensure that intercourse would not result in unnecessary costs. The third implication is that because women invest more and are essential for the reproductive success of their offspring they are a valuable resource for men; as a result, males often compete for sexual access to them.

Women as the more investing sex

Due to the heavy investment that women engage in after conception, evolution has favoured women who are more selective of their partners. Therefore, generally speaking women are more discriminate about their partners. Their mate preferences usually include males which are of good financial status, and could therefore invest more resources; who are more committed, and would therefore dedicate more time and energy; who are more athletic, as they would be better at protecting and defending the female and her offspring; and who are healthier, again with the prospect in mind that they would ensure healthy offspring. Women's preferences for mates, however, vary depending on their context. There are two contexts which have been identified to affect preferences. The first is women's personal resources, also referred to as the "structural powerlessness hypothesis".[20]

Women as a valuable resource for men

The second prediction that follows from Trivers' theory is that the fact that women invest more heavily in offspring makes them a valuable resource for males as it ensures the survival of their offspring which is the driving force of natural selection. Therefore, the sex that invests less in offspring with compete among themselves to breed with the more heavily investing sex. In other words, males will compete for females. It has been argued that jealousy has developed to avert the risk of potential loss of parental investment in offspring.[19] If a male redirects his resources to another female it is a costly loss of time, energy and resources for her offspring. However, the risks for males are higher because although women invest more in their offspring, they have bigger maternity certainty because they themselves have carried out the child. However, males can never have 100% paternal certainty and therefore risk investing resources and time in offspring that is genetically unrelated. Evolutionary psychology views jealousy as an adaptive response to this problem.

Very often parent-offspring conflict can find its roots in parental investment. Because parents are equally related to all offspring they try to distribute their investment equally and proportionally. However, because the offspring is more related to themselves than they are from their siblings, they often require a lot more care than the parent is able to provide. According to Trivers, this discrepancy in investment optima occurs because the parent is selected to invest in the offspring up until the point in which the cost of investing more in the current offspring and hence leading to less future reproductive success is larger than the benefit of increased survival of the current offspring. Conflict arises if the parent continues to invest in the offspring even to the critical point where the cost begins to outweigh the benefits.[21]

Sociosexuality

From an evolutionary perspective, the sexual strategy and parental investment of male and female differ considerably and females usually invest more in parental care due to a time-consuming period of pregnancy and childbirth, while male are not restricted by the sperm production but by access to more sexual partners.[18][22] There is a trade-off between mating effort and parenting effort, and males are generally more willing to have more sexual partners compared to females, as reproduction of more offspring instead of using the time and resource to invest in existing offspring would be a trade-off.[23] Therefore, the amount and duration of investment necessary to ensure the survival of offspring was unequal between the two sexes, with females investing more.[24]

On the other hand, Trivers has proposed that females prefer to mate with males who display both an ability and willingness to invest resources for the survival of the female and her offspring, thus, the costs of female parental care and investment could be balanced by benefits in terms of more paternal investment in resources for offspring,[25] This would mean that females who expect paternal investment in resources such as territory will be more conservative and careful towards having sexual intercourse with males, regardless of how attractive the potential male is.[26]

Sociosexual orientation refers to individual differences in willingness to engage in sexual relations without love or intimacy. Unrestricted sexuality refers to a kind of sexual strategy people adopts when they engage in more than one concurrent sexual relationship and have had sex with many different partners in the past, usually sex without commitment such as one-night-stands. Individuals with a restricted sexuality would often have closeness and commitment before engaging in a sexual relationship.[27][28] Using socio-sexual orientation inventory (SOI), study has demonstrated that females who expect heavy paternal investment will try to emphasise fidelity and chastity by scoring lower on the SOI, hence more restricted sexuality, while females who expect non-investing mates will imply males they are open to sexual intercourse by showing sexual attractiveness and display an unrestricted sexuality.[28][29]

The effect of father absence

Parental investment is any effort that increases quality of offspring, this could be things such as spending of time, energy and resources to help offspring to survive. Traditionally, females have greater parental effort while males have greater mating effort, and males often compete for the access to females and their investment in order to pass on their genes. In human parental investment, males are more uncertain compared to females when investing in offspring, as they don't know if the offspring belongs to them, so called paternity uncertainty. Therefore, lack of paternal investment constituted a unique and independent path to early sexual activity and adolescent pregnancy for females. Research has shown that exposure to father absence was strongly associated with risk for early sexual activity and teenage pregnancy.[30] Many studies have identified the absence of father as a major risk factor for both early sexual activity[31] and teenage pregnancy.[32] In the study, females that had no father absent since the age of 0 to 5 are more likely to engage in sexual activities earlier than females who had no father absent from 6 to 13.[33][34]

Another study looked at psychological/social stress in the developmental environment, the age of first menarche and the age of giving first birth to offspring, the data suggests that the family environment with father absence makes the family support more uncertain, which might trigger higher stress, resulting in early menarche. As menarche is the sign of puberty, females are more likely to engage in sexual activities earlier because of the facilitation of early menarche. Unrestricted female sexuality could also be explained by females who lack paternal input would seek outside for social support due to uncertain future and uncertain availability of males.[35]

Research indicates that lack of paternal investment also impacts on male development. Higher mortality, higher impulsivity, higher rape-prone activities and sexual coercion is associated with father absence, compared to males who grew up in father-presence environment.[36][37] These findings are consistent with life-course adversity models of early sexual activity and teenage pregnancy, which stated that the stress within the family environment over lifetime provokes the earlier onset of sexual activity and reproduction.[38][39]

Application of Trivers' theory in real life

Trivers' theory has been very influential as the predictions it makes correspond to differences in sexual behaviours of men and women, as demonstrated by a variety of research. Cross-cultural study from Buss (1989)[40] shows that males are tuned into physical attractiveness as it signals youth and fertility and ensures male reproductive success, which is increased by copulating with as many fertile females as possible. Women on the other hand are tuned into resources provided by potential mates, as their reproductive success is increased by ensuring their offspring will survive, and one way they do so is by getting resources for them. Alternatively, another study shows that men are more promiscuous than women, giving further support to this theory. Clark and Hatfield[41] found that 75% of men were willing to have sex with a female stranger when propositioned, compared to 0% of women. On the other hand, 50% of women agreed to a date with a male stranger. This suggests males seek short term relationships, while women show a strong preference for long-term relationships.

However, these preferences (male promiscuity and female choosiness) can be explained in other ways. In Western cultures, male promiscuity is encouraged through the availability of pornographic magazines and videos targeted to the male audience. Alternatively, both Western and Eastern cultures discourage female promiscuity through social checks such as slut-shaming.[42]

PIT also explains patterns of sexual jealousy.[19] He found males are more likely to show a stress response when imagining their partners showing sexual infidelity (having sexual relations with someone else), and women showed more stress when imagining their partner being emotionally unfaithful (being in love with another woman). PIT explains this, as woman's sexual infidelity decreases the male's paternal certainty, thus he will show more stress due to fear of cuckoldry. On the other hand, the woman fears losing the resources her partner provides. If her partner has an emotional attachment to another female it's likely that he won't invest into their offspring as much, thus a greater stress response is shown in this circumstance.

A heavy criticism of the theory comes from Thornhill and Palmer's analysis of it in A Natural History of Rape: Biological Bases of Sexual Coercion,[43] as it seems to rationalise rape and sexual coercion of females. Thornhill and Palmer claimed rape is an evolved technique for obtaining mates in an environment where women choose mates. As PIT claims males seek to copulate with as many fertile females as possible, the choice women have could result in a negative effect on the male's reproductive success. If women didn't choose their mates, Thornhill and Palmer claim there would be no rape. This ignores a variety of sociocultural factors, such as the fact that not only fertile females are raped – 34% of underage rape victims are under 12,[44] which means they are not of fertile age, thus there is no evolutionary advantage in raping them. 14% of rapes in England are committed on males,[45] who cannot increase a man's reproductive success as there will be no conception. Thus, what Thornhill and Palmer called an 'evolved machinery' might not be very advantageous.

Versus sexual strategies

Trivers' theory overlooks that women do have short-term relationships such as one-night stands, while not all men behave promiscuously. An alternative explanation to PIT and mate preferences would be Buss and Schmitt's sexual strategies theory.[23] SST argues that both sexes pursue short-term and long-term relationships, but seek different qualities in their short- and long-term partners. For a short-term relationship women will prefer an attractive partner, but in a long-term relationship they might be willing to trade-off that attractiveness for resources and commitment. On the other hand, men might be accepting of a sexually willing partner in a short-term relationships, but to ensure their paternal certainty they will seek a faithful partner instead.

See also

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References

  1. ↑ Clutton-Brock, T.H. 1991. The Evolution of Parental Care. Princeton, NJ: Princeton U. Press. pg. 9
  2. 1 2 Trivers, R.L. (1972). Parental investment and sexual selection. In B. Campbell (Ed.), Sexual selection and the descent of man, 1871-1971 (pp. 136–179). Chicago, IL: Aldine. ISBN 0-435-62157-2.
  3. ↑ Beatty, John. 1992. "Fitness: theoretical contexts," in Key Words in Evolutionary Biology. Edited by EF Keller and EA Lloyd, pp. 115–9. Cambridge, MA: Harvard U.Press
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  6. ↑ Olsson, Olof. "Clutch abandonment: a state-dependent decision in king penguins". Journal of Avian Biology. 28: 264–267. doi:10.2307/3676979.
  7. ↑ Burke, T.; Daviest, N. B.; Bruford, M. W.; Hatchwell, B. J. (1989). "Parental care and mating behaviour of polyandrous dunnocks Prunella modularis related to paternity by DNA fingerprinting". Nature. 338 (6212): 249–51. Bibcode:1989Natur.338..249B. doi:10.1038/338249a0.
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  9. ↑ Schürch, Roger, Roger & Taborsky, Barbara (2005). "The Functional Significance of Buccal Feeding in the Mouthbrooding Cichlid Tropheus moorii". Behaviour. 142 (3): 265–281. doi:10.1163/1568539053778274. JSTOR 4536244.
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  11. ↑ How the body shapes the way we think: A new view of intelligence, Rolf Pfeifer, Josh Bongard
  12. ↑ The unpredictable species: What makes humans unique, Philip Lieberman
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  14. ↑ Epstein, R. (1984). The principle of parsimony and some applications in psychology. Journal of Mind and Behavior, 5
  15. ↑ Bateman AJ (December 1948). "Intra-sexual selection in Drosophila". Heredity. 2 (3): 349–68. doi:10.1038/hdy.1948.21. PMID 18103134.
  16. ↑ Amundsen, Trond (1 April 2000). "Why are female birds ornamented". Trends in Ecology & Evolution. 15 (4): 149–55. doi:10.1016/S0169-5347(99)01800-5.
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  18. 1 2 Trivers, R. (1972). Parental investment and sexual selection. Sexual Selection & the Descent of Man, Aldine de Gruyter, New York, 136-179.
  19. 1 2 3 Buss, D. M., Larsen, R. J., Westen, D., & Semmelroth, J. (1992). Sex differences in jealousy: Evolution, physiology, and psychology. Psychological science, 3(4), 251-255.
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  23. 1 2 Buss, D. M., & Schmitt, D. P. (1993). Sexual strategies theory: an evolutionary perspective on human mating. Psychological review, 100(2), 204.
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  26. ↑ Baumeister, R. F., & Vohs, K. D. (2004). Sexual economics: Sex as female resource for social exchange in heterosexual interactions. Personality and Social Psychology Review, 8(4), 339-363.
  27. ↑ Gangestad, S. W., Simpson, J. A., DiGeronimo, K., & Biek, M. (1992). Differential accuracy in person perception across traits: examination of a functional hypothesis. Journal of personality and social psychology, 62(4), 688.
  28. 1 2 Simpson, J. A., & Gangestad, S. W. (1991). Individual differences in sociosexuality: evidence for convergent and discriminant validity. Journal of personality and social psychology, 60(6), 870.
  29. ↑ Cashdan, E. (1993). Attracting mates: Effects of paternal investment on mate attraction strategies. Ethology and Sociobiology, 14(1), 1-23.</Socio-sexuality
  30. ↑ Ellis, B. J., Bates, J. E., Dodge, K. A., Fergusson, D. M., John Horwood, L., Pettit, G. S., & Woodward, L. (2003). Does father absence place daughters at special risk for early sexual activity and teenage pregnancy?. Child development, 74(3), 801-821.
  31. ↑ Kiernan, K. E., & Hobcraft, J. (1997). Parental divorce during childhood: age at first intercourse, partnership and parenthood. Population Studies, 51(1), 41-55.
  32. ↑ McLanahan, S. S. (1999). Father absence and the welfare of children. Coping with divorce, single parenting, and remarriage: A risk and resiliency perspective, 117-145.
  33. ↑ Ellis, B. J., Bates, J. E., Dodge, K. A., Fergusson, D. M., John Horwood, L., Pettit, G. S., & Woodward, L. (2003). Does father absence place daughters at special risk for early sexual activity and teenage pregnancy?. Child development, 74(3), 801-821.
  34. ↑ Scaramella, L. V., Conger, R. D., Simons, R. L., & Whitbeck, L. B. (1998). Predicting risk for pregnancy by late adolescence: a social contextual perspective. Developmental psychology, 34(6), 1233.
  35. ↑ Chisholm, J. S., Quinlivan, J. A., Petersen, R. W., & Coall, D. A. (2005). Early stress predicts age at menarche and first birth, adult attachment, and expected lifespan. Human Nature, 16(3), 233-265.
  36. ↑ Lynn, D. B., & Sawrey, W. L. (1959). The effects of father-absence on Norwegian boys and girls. The Journal of Abnormal and Social Psychology, 59(2), 258.
  37. ↑ Malamuth, N. M. (1981). Rape proclivity among males. Journal of social issues, 37(4), 138-157.
  38. ↑ Belsky, J., Steinberg, L., & Draper, P. (1991). Childhood experience, interpersonal development, and reproductive strategy: An evolutionary theory of socialization. Child development, 62(4), 647-670.
  39. ↑ Coley, R. L., & Chase-Lansdale, P. L. (1998). Adolescent pregnancy and parenthood: recent evidence and future directions. American Psychologist, 53(2), 152.
  40. ↑ Buss, D. M. (1989). Sex differences in human mate preferences: Evolutionary hypotheses tested in 37 cultures. Behavioral and brain sciences, 12(01), 1-14.
  41. ↑ Clark, R. D., & Hatfield, E. (1989). Gender differences in receptivity to sexual offers. Journal of Psychology & Human Sexuality, 2(1), 39-55.
  42. ↑ Armstrong, E. A., Hamilton, L. T., Armstrong, E. M., & Seeley, J. L. (2014). “Good Girls” Gender, Social Class, and Slut Discourse on Campus. Social Psychology Quarterly, 77(2), 100-122
  43. ↑ Thornhill, R., & Palmer, C. T. (2001). A natural history of rape: Biological bases of sexual coercion. MIT press.
  44. ↑ "Children and Teens: Statistics - RAINN".
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Further reading

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