Parental investment

Parental investment, in evolutionary biology and evolutionary psychology, is any parental expenditure (e.g. time, energy, resources) that benefits offspring. Parental investment may be performed by both males and females (biparental care), females alone (exclusive maternal care) or males alone (exclusive paternal care). Care can be provided at any stage of the offspring's life, from pre-natal (e.g. egg guarding and incubation in birds, and placental nourishment in mammals) to post-natal (e.g. food provisioning and protection of offspring). Parental investment, in evolutionary biology and evolutionary psychology, is any parental expenditure (e.g. time, energy, resources) that benefits offspring. Parental investment may be performed by both males and females (biparental care), females alone (exclusive maternal care) or males alone (exclusive paternal care). Care can be provided at any stage of the offspring's life, from pre-natal (e.g. egg guarding and incubation in birds, and placental nourishment in mammals) to post-natal (e.g. food provisioning and protection of offspring). Parental investment theory, a term coined by Robert Trivers in 1972, predicts that the sex that invests more in its offspring will be more selective when choosing a mate, and the less-investing sex will have intra-sexual competition for access to mates. This theory has been influential in explaining sex differences in sexual selection and mate preferences, throughout the animal kingdom and in humans. In 1859, Charles Darwin published On the Origin of Species. This introduced the concept of natural selection to the world, as well as related theories such as sexual selection. For the first time, evolutionary theory was used to explain why females are 'coy' and males are 'ardent' and compete with each other for females' attention. In 1930, Ronald Fisher wrote The Genetical Theory of Natural Selection, in which he introduced the modern concept of parental investment, introduced the sexy son hypothesis, and introduced Fisher's principle. In 1948, Angus John Bateman published an influential study of fruit flies in which he concluded that because female gametes are more costly to produce than male gametes, the reproductive success of females was limited by the ability to produce ovum, and the reproductive success of males was limited by access to females. In 1972, Trivers continued this line of thinking with his proposal of parental investment theory, which describes how parental investment affects sexual behavior. He concludes that the sex that has higher parental investment will be more selective when choosing a mate, and the sex with lower investment will compete intra-sexually for mating opportunities. In 1974, Trivers extended parental investment theory to explain parent-offspring conflict, the conflict between investment that is optimal from the parent's versus the offspring's perspective. 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, wherein Fisher argued that parental expenditure on both sexes of offspring should be equal. Clutton-Brock expanded the concept of parental investment to include costs to any other component of parental fitness. Male dunnocks tend to not discriminate between their own young and those of another male in polyandrous or polygynandrous systems. 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. This indiscriminative parental care by males is also observed in redlip blennies. 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. In some species, such as humans and many birds, the offspring are altricial and unable to fend for themselves for an extended period of time after birth. In these species, males invest more in their offspring than do the male parents of precocial species, since reproductive success would otherwise suffer. 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. 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.