Parent–offspring Conflict
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Parent–offspring conflict (POC) is an expression coined in 1974 by
Robert Trivers Robert Ludlow "Bob" Trivers (; born February 19, 1943) is an American evolutionary biologist and sociobiologist. Trivers proposed the theories of reciprocal altruism (1971), parental investment (1972), facultative sex ratio determination (197 ...
. It is used to describe the evolutionary conflict arising from differences in optimal
parental investment Parental investment, in evolutionary biology and evolutionary psychology, is any parental expenditure (e.g. time, energy, resources) that benefits offspring.Clutton-Brock, T.H. 1991. ''The Evolution of Parental Care''. Princeton, NJ: Princeton ...
(PI) in an offspring from the standpoint of the
parent A parent is either the progenitor of a child or, in humans, it can refer to a caregiver or legal guardian, generally called an adoptive parent or step-parent. Parents who are progenitors are First-degree relative, first-degree relatives and have ...
and the
offspring In biology, offspring are the young creation of living organisms, produced either by sexual reproduction, sexual or asexual reproduction. Collective offspring may be known as a brood or progeny. This can refer to a set of simultaneous offspring ...
. PI is any investment by the parent in an individual offspring that decreases the parent's ability to invest in other offspring, while the selected offspring's chance of surviving increases. POC occurs in sexually reproducing species and is based on a genetic conflict: Parents are equally related to each of their offspring and are therefore expected to equalize their investment among them. Offspring are only half or less related to their
sibling A sibling is a relative that shares at least one parent with the other person. A male sibling is a brother, and a female sibling is a sister. A person with no siblings is an only child. While some circumstances can cause siblings to be raised ...
s (and fully related to themselves), so they try to get more PI than the parents intended to provide even at their siblings' disadvantage. However, POC is limited by the close genetic relationship between parent and offspring: If an offspring obtains additional PI at the expense of its siblings, it decreases the number of its surviving siblings. Therefore, any
gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...
in an offspring that leads to additional PI decreases (to some extent) the number of surviving copies of itself that may be located in siblings. Thus, if the costs in siblings are too high, such a gene might be selected against despite the benefit to the offspring. The problem of specifying how an individual is expected to weigh a relative against itself has been examined by
W. D. Hamilton William Donald Hamilton (1 August 1936 – 7 March 2000) was a British evolutionary biologist, recognised as one of the most significant evolutionary theorists of the 20th century. Hamilton became known for his theoretical work expounding a ...
in 1964 in the context of
kin selection Kin selection is a process whereby natural selection favours a trait due to its positive effects on the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin selection can lead ...
.
Hamilton's rule Kin selection is a process whereby natural selection favours a trait due to its positive effects on the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin selection can lead ...
says that altruistic behavior will be positively selected if the benefit to the recipient multiplied by the genetic relatedness of the recipient to the performer is greater than the cost to the performer of a social act. Conversely, selfish behavior can only be favoured when Hamilton's inequality is not satisfied. This leads to the prediction that, other things being equal, POC will be stronger under half siblings (e.g., unrelated males father a female's successive offspring) than under full siblings.


Occurrence


In plants

In
plants Plants are the eukaryotes that form the kingdom Plantae; they are predominantly photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with cyanobacteria to produce sugars f ...
, POC over the allocation of resources to the brood members may affect both brood size (number of
seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s matured within a single fruit) and seed size. Concerning brood size, the most economic use of maternal resources is achieved by packing as many seeds as possible in one fruit, i.e., minimizing the cost of packing per seed. In contrast, offspring benefits from low numbers of seeds per fruit, which reduces sibling competition before and after dispersal. Conflict over seed size arises because there usually exists an inverse exponential relationship between seed size and fitness, that is, the fitness of a seed increases at a diminishing rate with resource investment but the fitness of the maternal parent has an optimum, as demonstrated by Smith and Fretwell (see also
marginal value theorem The marginal value theorem (MVT) is an optimality model that usually describes the behavior of an optimally foraging individual in a system where resources (often food) are located in discrete patches separated by areas with no resources. Due to t ...
). However, the optimum resource investment from the offspring's point of view would be the amount that optimizes its
inclusive fitness Inclusive fitness is a conceptual framework in evolutionary biology first defined by W. D. Hamilton in 1964. It is primarily used to aid the understanding of how social traits are expected to evolve in structured populations. It involves partit ...
(direct and indirect fitness), which is higher than the maternal parent's optimum. This conflict about resource allocation is most obviously manifested in the reduction of brood size (i.e. a decrease in the proportion of
ovule In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the sporangium, megasporangium), ...
s matured into seeds). Such reduction can be assumed to be caused by the offspring: If the maternal parent's interest were to produce as few seeds as observed,
selection Selection may refer to: Science * Selection (biology), also called natural selection, selection in evolution ** Sex selection, in genetics ** Mate selection, in mating ** Sexual selection in humans, in human sexuality ** Human mating strat ...
would not favour the production of extra ovules that do not mature into seeds. (Although other explanations for this phenomenon exist, such as genetic load, resource depletion or maternal regulation of offspring quality, they could not be supported by experiments.) There are several possibilities how the offspring can affect paternal resource allocation to brood members. Evidence exists for
siblicide Siblicide (attributed by behavioural ecologist Doug Mock to Barbara M. Braun) is the killing of an infant individual by its close relatives (full or half siblings). It may occur directly between siblings or be mediated by the parents, and is dr ...
by dominant embryos: Embryos formed early kill the remaining embryos through an aborting chemical. In oaks, early fertilized ovules prevent the
fertilization Fertilisation or fertilization (see American and British English spelling differences#-ise, -ize (-isation, -ization), spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give ...
of other ovules by inhibiting the
pollen tube A pollen tube is a tubular structure produced by the male gametophyte of seed plants when it germinates. Pollen tube elongation is an integral stage in the plant life cycle. The pollen tube acts as a conduit to transport the male gamete cells fr ...
entry into the
embryo sac In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the fe ...
. In some species, the maternal parent has evolved postfertilization
abortion Abortion is the early termination of a pregnancy by removal or expulsion of an embryo or fetus. Abortions that occur without intervention are known as miscarriages or "spontaneous abortions", and occur in roughly 30–40% of all pregnan ...
of few seeded pods. Nevertheless, cheating by the offspring is also possible here, namely by late siblicide, when the postfertilization abortion has ceased. According to the general POC model, reduction of brood size – if caused by POC – should depend on genetic relatedness between offspring in a fruit. Indeed, abortion of embryos is more common in
out-crossing Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity. Outcrossing in animals Out ...
than in
self-pollinating Self-pollination is a form of pollination in which pollen arrives at the Stigma (botany), stigma of a flower (in flowering plants) or at the ovule (in gymnosperms) of the same plant. The term cross-pollination is used for the opposite case, where ...
plants (seeds in cross-pollinating plants are less related than in self-pollinating plants). Moreover, the level of solicitation of resources by the offspring is also increased in cross-pollinating plants: There are several reports that the average weight of crossed seeds is greater than of seeds produced by self-fertilization.


In birds

Some of the earliest examples of parent-offspring conflict were seen in
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
broods and especially in raptor species. While parent birds often lay two eggs and attempt to raise two or more young, the strongest fledgling takes a greater share of the food brought by parents and will often kill the weaker sibling (
siblicide Siblicide (attributed by behavioural ecologist Doug Mock to Barbara M. Braun) is the killing of an infant individual by its close relatives (full or half siblings). It may occur directly between siblings or be mediated by the parents, and is dr ...
). Such conflicts have been suggested as a driving force in the evolution of optimal clutch size in birds. In the
blue-footed booby The blue-footed booby (''Sula nebouxii'') is a marine bird native to subtropical and tropical regions of the eastern Pacific Ocean. It is one of six species of the genus ''booby, Sula'' – known as boobies. It is easily recognizable by its dis ...
, parent-offspring conflict results in times of food scarcity. When there is less food available in a given year, the older, dominant chick will often kill the younger chick by either attacking directly, or by driving it from the nest. Parents try to prevent siblicide by building nests with steeper sides and by laying heavier second eggs.


In mammals

Even before POC theory arose, debates took place over whether
infant In common terminology, a baby is the very young offspring of adult human beings, while infant (from the Latin word ''infans'', meaning 'baby' or 'child') is a formal or specialised synonym. The terms may also be used to refer to juveniles of ...
s
wean Weaning is the process of gradually introducing an infant human or other mammal to what will be its adult diet while withdrawing the supply of its mother's milk. In the UK, weaning primarily refers to the introduction of solid foods at 6 mont ...
themselves or mothers actively wean their infants. Furthermore, it was discussed whether maternal rejections increase infant independence. It turned out that both mother and infant contribute to infant independence. Maternal rejections can be followed by a short-term increase in infant contact but they eventually result in a long-term decrease of contact as has been shown for several primates: In wild
baboon Baboons are primates comprising the biology, genus ''Papio'', one of the 23 genera of Old World monkeys, in the family Cercopithecidae. There are six species of baboon: the hamadryas baboon, the Guinea baboon, the olive baboon, the yellow ba ...
s infants that are rejected early and frequently spend less time in contact whereas those that are not rejected stay much longer in the proximity of their mother and suckle or ride even in advanced ages. In wild
chimpanzee The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of Hominidae, great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close rel ...
s an abrupt increase in maternal rejections and a decrease in mother-offspring contact is found when mothers resume
estrus The estrous cycle (, originally ) is a set of recurring physiological changes induced by reproductive hormones in females of mammalian subclass Theria. Estrous cycles start after sexual maturity in females and are interrupted by anestrous phas ...
and consort with males. In
rhesus macaque The rhesus macaque (''Macaca mulatta''), colloquially rhesus monkey, is a species of Old World monkey. There are between six and nine recognised subspecies split between two groups, the Chinese-derived and the Indian-derived. Generally brown or g ...
s a high probability of conception in the following mating season is associated with a high rate of maternal rejection. Rejection and behavioral conflicts can occur during the first months of an infant's life and when the mother resumes estrus. These findings suggest that the
reproduction Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. There are two forms of reproduction: Asexual reproduction, asexual and Sexual ...
of the mother is influenced by the interaction with their offspring. So there is a potential for conflicts over PI. It was also observed in rhesus macaques that the number of contacts made by offspring is significantly higher than the number of contacts made by mother during a mating season, whereas the opposite holds for the number of broken contacts. This fact suggests that the mother resists offspring's demands for contact, whereas offspring is apparently more interested in spending time in contact. At three months of infant age a shift from mother to infant in responsibility for maintaining contact takes place. So when the infant becomes more independent, its effort to maintain proximity to its mother increases. This might sound paradoxical but becomes clear when one takes into account that POC increases during the period of PI. In summary, all these findings are consistent with POC-theory. One might object that time in contact is not a reasonable measure for PI and that, for example, time for milk transfer (
lactation Lactation describes the secretion of milk from the mammary glands and the period of time that a mother lactates to feed her young. The process naturally occurs with all sexually mature female mammals, although it may predate mammals. The process ...
) would be a better one. Here one can argue that mother and infant have different thermoregulatory needs due to the fact that they have different surface-to-volume ratios resulting in more rapid loss of heat in infants compared to adults. So infants may be more sensitive to low temperatures than their mothers. An infant might try to compensate by increased contact time with their mother, which could initiate a behavioral conflict over time. Consistency of this hypothesis has been shown for
Japanese macaque The Japanese macaque (''Macaca fuscata''), also known as the snow monkey, is a terrestrial Old World monkey species that is native to Japan. Colloquially, they are referred to as "snow monkeys" because some live in areas where snow covers the g ...
s where decreasing temperatures result in higher maternal rejections and increased number of contacts made by infants.


In social insects

In eusocial species, the parent-offspring conflict takes on a unique role because of
haplodiploidy Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky. Haplodiploidy determines the s ...
and the prevalence of sterile workers. Sisters are more related to each other (0.75) than to their mothers (0.5) or brothers (0.25). In most cases, this drives female workers to try and obtain a sex ratio of 3:1 (females to males) in the colony. However, queens are equally related to both sons and daughters, so they prefer a sex ratio of 1:1. The conflict in social insects is about the level investment the queen should provide for each sex for current and future offspring. It is generally thought that workers will win this conflict and the sex ratio will be closer to 3:1, however there are examples, like in '' Bombus terrestris'', where the queen has considerable control in forcing a 1:1 ratio.


In amphibians

Many species of frogs and salamanders display complex social behavior with highly involved parental care that includes egg attendance, tadpole transport, and tadpole feeding.


Energy expenditure

Both males and females of the strawberry poison-dart frog care for their offspring, however, females invest in more costly ways. Females of certain poison frog species produce unfertilized, non-developing
trophic Trophic, from Ancient Greek τροφικός (''trophikos'') "pertaining to food or nourishment", may refer to: * Trophic cascade * Trophic coherence * Trophic egg * Trophic function * Trophic hormone * Trophic level index * Trophic level * ...
eggs which provide nutrition to her tadpoles. The tadpoles vibrate vigorously against mother frogs to solicit nutritious eggs. These maternal trophic eggs are beneficial for offspring, positively influencing larval survival, size at metamorphosis, and post metamorphic survival. In the neotropical, foam-nesting pointedbelly frog ('' Leptodactylus podicipinus''), females providing parental care to tadpoles have reduced body condition and food ingestion. Females that are attending to her offspring have significantly lower body mass, ovary mass, and stomach volume. This indicates that the cost of parental care in the pointedbelly frog has the potential to affect future reproduction of females due to the reaction in body condition and food intake. In the Puerto Rican common coqui, parental care is performed exclusively by males and consists of attending to the eggs and tadpoles at an oviposition site. When brooding, males have a higher frequency of empty stomachs and lose a significant portion of their initial body mass during parental care. Abdominal fat bodies of brooding males during the middle of parental care were significantly smaller than those of non-brooding males. Another major behavioral component of parental care is nest defense against conspecific egg cannibals. This defense behavior includes aggressive calling, sustained biting, wrestling, and blocking directed against the nest intruder. Females of the Allegheny Mountain dusky salamander exhibit less activity and become associated with the nest site well in advance of oviposition in preparation for the reproductive season. This results in a reduced food intake and a decrease in body weight over the
brooding Broodiness is the action or behavioral tendency to sit on a clutch of eggs to Egg incubation, incubate them, often requiring the non-expression of many other behaviors including feeding and drinking.Homedes Ranquini, J. y Haro-García, F. Zoogen ...
period. Females either stop or greatly reduce their foraging activities and instead will eat opportunistically following oviposition. Since nutritional intake is reduced, there is a decrease in body weight in females. Females of the red-backed salamander make a substantial parental investment in terms of clutch size and brooding behavior. When brooding, females usually do not leave their eggs to forage but rather rely upon their fat reserves and any resources they encounter at their oviposition site. In addition, females could experience metabolic costs while safeguarding their offspring from desiccation, intruders, and predators.


Time investment

The plasticity of tadpoles may play a role in the weaning conflict in egg-feeding frogs, in which the offspring prefer to devote resources to growth, while the mother prefers nutrients to help her young become independent. A similar conflict happens in direct-developing frogs that care for clutches, with protected tadpoles having the advantage of a slower, safer development, but they need to be ready to reach independence rapidly due to the risks of predation or
desiccation Desiccation is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. The ...
. In the neotropical Zimmerman’s poison frog, the males provide a specific parental care in the form of transportation. The tadpoles are cannibalistic, hence why the males typically separate them from their siblings after hatching by transporting them to small bodies of water. However, in some cases parents do not transport their tadpoles but let them all hatch into the same pool. In order to escape their cannibalistic siblings, the tadpoles will actively seek transportative parental care. When a male frog approaches the water body in which the tadpoles had been deposited in, tadpoles will almost “jump” on the back of the adult, mimicking an attack, while adults would not assist with this movement. While this is an obvious example of sibling conflict, the one-sided interaction between tadpoles and frogs could be seen as a form of parent-offspring conflict, in which the offspring attempts to extract more from the interaction than the parent is willing to provide. In this scenario, a tadpole climbing onto an unwilling frog— who enters the pool for reasons other than tadpole transportation, such as egg deposition, cooling off, or sleeping— might be analogous to mammalian offspring seeking to nurse after weaning. In times of danger, the tadpoles of Zimmerman’s poison frog don't passively await parental assistance but instead exhibit an almost aggressive approach in mounting the adult frogs.


Trade-offs with mating

Reproductive attempts in strawberry poison-dart frog such as courtship activity, significantly decreases or will entirely cease in tadpole-rearing females compared to non-rearing females. Most brooding males of the common coqui cease calling during parental care while gravid females are still available and known to mate, hence why non-calling males miss potential opportunities to reproduce. Caring for tadpoles comes at the cost of other current reproductive opportunities for females, leading to the hypothesis that frequent reproduction is associated with reduced survival in frogs.


In humans

An important illustration of POC within humans is provided by David Haig’s (1993) work on genetic conflicts in pregnancy. Haig argued that
fetal A fetus or foetus (; : fetuses, foetuses, rarely feti or foeti) is the unborn offspring of a viviparous animal that develops from an embryo. Following the embryonic stage, the fetal stage of development takes place. Prenatal development is a ...
genes would be selected to draw more resources from the mother than would be optimal for the mother to give. The
placenta The placenta (: placentas or placentae) is a temporary embryonic and later fetal organ that begins developing from the blastocyst shortly after implantation. It plays critical roles in facilitating nutrient, gas, and waste exchange between ...
, for example, secretes allocrine
hormone A hormone (from the Ancient Greek, Greek participle , "setting in motion") is a class of cell signaling, signaling molecules in multicellular organisms that are sent to distant organs or tissues by complex biological processes to regulate physio ...
s that decrease the sensitivity of the mother to
insulin Insulin (, from Latin ''insula'', 'island') is a peptide hormone produced by beta cells of the pancreatic islets encoded in humans by the insulin (''INS)'' gene. It is the main Anabolism, anabolic hormone of the body. It regulates the metabol ...
and thus make a larger supply of blood sugar available to the fetus. The mother responds by increasing the level of insulin in her bloodstream and to counteract this effect the placenta has insulin receptors that stimulate the production of insulin-degrading
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
s. About 30 percent of human conceptions do not progress to full term (22 percent before becoming clinical pregnancies) creating a second arena for conflict between the mother and the fetus. The fetus will have a lower quality cut off point for spontaneous abortion than the mother. The mother's quality cut-off point also declines as she nears the end of her reproductive life, which becomes significant for older mothers. Older mothers have a higher incidence of offspring with genetic defects. Indeed, with parental age on both sides, the mutational load increases as well. Initially, the maintenance of pregnancy is controlled by the maternal hormone
progesterone Progesterone (; P4) is an endogenous steroid and progestogen sex hormone involved in the menstrual cycle, pregnancy, and embryogenesis of humans and other species. It belongs to a group of steroid hormones called the progestogens and is the ma ...
, but in later stages it is controlled by the fetal human chorionic gonadotrophin released into the maternal bloodstream. The release of fetal human chorionic gonadotrophin causes the release of maternal progesterone. There is also conflict over blood supply to the placenta, with the fetus being prepared to demand a larger blood supply than is optimal for the mother (or even for itself, since high birth weight is a risk factor). This results in
hypertension Hypertension, also known as high blood pressure, is a Chronic condition, long-term Disease, medical condition in which the blood pressure in the artery, arteries is persistently elevated. High blood pressure usually does not cause symptoms i ...
and, significantly, high
birth weight Birth weight is the body weight of a baby at their birth. The average birth weight in babies of European and African descent is , with the normative range between . 15% of babies born in 2012 had a low birth weight and 14.7% in 2020. It is pro ...
is positively correlated with maternal blood pressure.


A tripartite (fetus–mother–father) immune conflict in humans and other placentals

During pregnancy, there is a two-way traffic of immunologically active cell lines through the placenta. Fetal
lymphocyte A lymphocyte is a type of white blood cell (leukocyte) in the immune system of most vertebrates. Lymphocytes include T cells (for cell-mediated and cytotoxic adaptive immunity), B cells (for humoral, antibody-driven adaptive immunity), an ...
lines may survive in women even decades after giving birth.


See also

* Intrauterine cannibalism * The kinship theory of genomic imprinting * Intragenomic and intrauterine conflict in humans


References


External links


Genetic conflicts in human pregnancyParent-offspring conflict
{{DEFAULTSORT:Parent-offspring conflict Evolutionary biology Reproduction