Models of parent-offspring conflict. III. Intra-brood conflict

A genetic model of parent-offspring conflict is developed for the situation where conflictors cause the parent to redirect resources from contemporaneous siblings to themselves, by increased solicitation. It is shown that there is no restriction on the spread of genes causing such conflict if there is no direct cost associated with it. We examine the effect of imposing on the conflictors a cost of the extra solicitation when the cost is felt (a) by the individual conflictor only, and (b) by all members of the brood equally. It is shown that the Evolutionarily Stable Strategies for the two situations allow a greater degree of solicitation when its cost is borne by the whole brood. Multipaternity of broods also increases the degree of conflict.     

Models of parent-offspring conflict. II. Promiscuity

The population genetics of Trivers (1974) concept of parent-offspring is examined for species in which the effects of the conflict are felt by future half-sibs, as in promiscuous mating systems in which the male shows no parental care. Whether or not a rare conflictor gene will spread in a non-conflictor population depends on f(m) greater than (m + 1)/(0.5m + 1.5) for a dominant gene, and on f(m) greater than 1/4(7 + 3) for a recessive gene; f(m) is the fitness gained by a conflictor relative to a non-conflictor offspring [f(m) greater than 1], and m is the amount of parental investment taken by a conflictor relative to m = 1 for a non-conflictor. The ESS value for conflict (mo) in promiscuous species with zero male parental care has mo = f(mo)/4[df(mo)/dmo]. However, where the male maintains the same harem for several breeding seasons, or where there is promiscuity but both sexes contribute equally to parental care, conditions for conflict are equivalent to monogamy.     

Models of parent-offspring conflict. I. Monogamy

Theoretical models for Trivers (1974) concept of parent-offspring conflict are examined for species in which the effects of the conflict are felt by full sibs. A rare conflictor gene will spread if (f(m) greater than 1/2(m + 1), where f(m) is the fitness gained by a conflictor relative to a non-conflictor offspring (f(m) greater than 1), and m is the amount of parental investment taken by a conflictor relative to m = 1 for a non-conflictor. The range of m alleles which can spread against the parent optimum decreases as the cost to the parent increases until a point is reached where there is no conflict of evolutionary interests. There would be no polymorphism for conflictor: non-conflictor alleles unless special conditions prevail. The conflictor allele which spreads most rapidly as a rare mutant against the parental optimum is not an evolutionarily stable strategy (ESS). The ESS for parent-offspring conflict in monogamous species has mo = f(mo)/2[df(mo)/dmo]. The analytical solutions are confirmed throughout by simulations.     

Models of parent-offspring conflict. V. Effects of the behaviour of the two parents

In the absence of any parent-offspring conflict, the total parental investment per offspring should be less when two parents collaborate in caring for the offspring than when only one parent invests. This does not necessarily mean that offspring fare less well when both parents invest. The ‘ideal’ amount of parental investment for an offspring to take is always greater than is ‘ideal’ for the parent to allocate (Trivers 1974). The offspring's optimum is higher if the offspring's action affects the reproductive success of only one parent and lower if both parents are affected (e.g. two-parent investment, or lifelong monogamy). The difference between the parental optimum and the offspring optimum depends on the mating system and on the form of conflict (between successive broods, or within broods), and prescribes a ‘conflict range’. The extent of conflict cannot be deduced solely from a knowledge of the average relatedness between siblings. The conflict is likely to be resolved by an ESS in which intermediate (compromise) levels of investment are paid out to offspring, which nevertheless continue to make costly demands for yet more investment. The degree of conflict can be measured by the extent to which offspring subject their parents to aggressive demands for extra investment, and is likely to be greater when two parents collaborate equally over investment than when only one parent invests. When only one parent invests, conflict is higher if sibling-competition is between siblings in the same broods (intra-brood) than when it is between progeny in successive broods (inter-brood). However, the reverse will tend to be the case when both parents invest equally.     

Packaging of offspring by nests of the ant,Leptothorax longispinosus: parent-offspring conflict and queen-worker conflict

Models of the packaging of offspring predict that parental fitness is maximized by following a set of rules, including the rule to invest the minimal amount in each offspring. Offspring can maximize their fitness by demanding more resources than the parent is selected to give, leading to parent-offspring conflict over packaging. Social insect nests may also experience queen-worker conflict over packaging. Experiments were conducted, using two populations of the ant Leptothorax longispinosus, in order to determine the role of both parent-offspring conflict and queen-worker conflict in packaging. Parent-offspring conflict over packaging was detected towards males and workers, but not to females. This may be because both parental and offspring fitness are maximized by investing as much in possible in females so both parties benefit by cooperating over packaging of females. Queen-worker conflict over packaging was detected for females, males, and workers. The direction taken by the queen-worker conflict is best explained by asymmetries in genetic relatedness among nestmates.     

A model of triploid endosperm evolution driven by parent-offspring conflict

The parental investment in angiosperms comprises the endosperm, a nutrient reserve that is used during seed development. The endosperm contains genes from both parents. The most common endosperm form is the 3n Polygonum -type with more maternal genetic influence than paternal, i.e. with two maternal nuclei and one paternal nucleus. The evolutionary original state is thought to be a diploid endosperm with equal influence of the parents. We focus on the evolution of the triploid endosperm and show that a gene for triploid endosperm would have an initial advantage in a population of diploid endosperm type plants, and increase to fixation. We assume that endosperm amount is controlled by endosperm genes. Then a gene causing triploid endosperm will increase the influence of the mother plant on parental investment. The production of endosperm with two copies of the maternal genes will modify the inheritance of endosperm amount and cause an increased production of seeds.     

Sibling competition stabilizes signalling resolution models of parent-offspring conflict

Young of altricial birds use conspicuous displays to solicit food from their parents. There is experimental evidence that the intensity of these displays is correlated with the level of food deprivation of young, and that parents respond to increased levels of solicitation by increasing the rate of food delivery to the nest. Game-theoretical models based on the handicap principle show that, when solicitation is costly, there is a signalling equilibrium at which there is a one-to-one correspondence between the condition of the young and the intensity of their display. Parents use this information to adjust their levels of investment on the current offspring. However, the models also have a non-signalling equilibrium, and computer simulations show that only the non-signalling equilibrium is stable. Here I show that when direct sibling competition is introduced into the model, in such a way that parents have control on the amount of food provided to the nest, but not on the way the food is allocated among siblings, the non-signalling equilibrium disappears and the signalling equilibrium becomes stable.     

A theoretical model of the evolution of maternal effects under parent-offspring conflict

The evolution of maternal effects on offspring phenotype should depend on the extent of parent-offspring conflict and costs and constraints associated with maternal and offspring strategies. Here, we develop a model of maternal effects on offspring dispersal phenotype under parent-offspring conflict to evaluate such dependence. In the absence of evolutionary constraints and costs, offspring evolve dispersal rates from different patch types that reflect their own, rather than the maternal, optima. This result also holds true when offspring are unable to assess their own environment because the maternal phenotype provides an additional source of information. Consequently, maternal effects on offspring diapause, dispersal, and other traits that do not necessarily represent costly resource investment are more likely to maximize offspring than maternal fitness. However, when trait expression was costly, the evolutionarily stable dispersal rates tended to deviate from those under both maternal and offspring control. We use our results to (re)interpret some recent work on maternal effects and their adaptive value and provide suggestions for future work.     

The influence of mating system on the intensity of parent-offspring conflict in primates

An evolutionary conflict of interest exists between parents and their offspring over the partitioning of parental investment (PI) among siblings. When the direct fitness benefits to offspring of increased PI, outweigh the inclusive fitness costs from lost future sibling fitness, selection should favour the evolution of offspring selfishness over altruism. In theory, this conflict is heightened when females are not strictly monogamous, as current offspring should be less altruistic towards future half-siblings than they would be towards full-siblings. Using data collected on foetal growth rate (representing prenatal PI) in primates, I test the prediction from theory that the resolution of the parent-offspring conflict will be closer to the offspring's evolutionary optima in polyandrous species than in more monandrous species. Using phylogenetic comparative analysis, and controlling for allometry, I show that offspring are able to obtain more PI when the probability of future full-siblings decreases, and that this is most pronounced in taxa where there is the opportunity for direct foetal access to the maternal bloodstream. These results support the hypothesis that the resolution of prenatal PI conflict is influenced by both a species' mating system and by its placental structure.     

Testing the viviparity-driven-conflict hypothesis: parent-offspring conflict and the evolution of reproductive isolation in a poeciliid fish

The evolution of viviparity increases the potential for genomic conflicts between mothers and offspring over the level of maternal investment. The viviparity-driven-conflict hypothesis predicts that such conflicts will drive the evolution of asymmetrical reproductive isolation between populations with divergent mating systems. We tested this hypothesis using crosses between populations of a poeciliid fish that differ in their level of polyandry. Our results support the prediction of an asymmetry in the rate of spontaneous abortion in reciprocal crosses, with the highest rate occurring in crosses between females from a relatively monandrous population and males from a relatively polyandrous population. The patterns of offspring size were not consistent with the pattern predicted by the viviparity-driven-conflict hypothesis: crosses between a monandrous female and a polyandrous male did not produce larger offspring than the reciprocal cross. This discrepancy was due to the presence of an effect of the maternal population on offspring size: polyandrous females produced larger offspring than monandrous females. In addition, offspring size was positively correlated with maternal size in crosses involving a polyandrous male. We discuss these results in light of models for intra- and intergenomic epistasis and the rapid origin of asymmetric reproductive isolation in viviparous taxa.     

Factors influencing family rupture and parent-offspring conflict in the Black Kite Milvus migrans

Juvenile and adult behaviour was studied at eight nests of Black Kites Milvus migrans within the Doñana Biological Reserve, Spain. Parental investment in vigilance and defence of offspring progressively decreased during the post-fledging dependence period. The number of feeds was also slightly reduced towards the end of the period. However, this does not seem to be the main factor which leads to juvenile independence. The fact that the family rupture is sudden and that the post-fledging dependence period tends to shorten as the season progresses suggests that juvenile and adult migratory urgency may be as important a factor as reduced parental investment in breaking the family ties.     

Evolutionary conflict: sperm wars, phantom inseminations

A new experimental study has provided the first definitive evidence for conditional punishment of 'cheats' in a sperm-trading simultaneous hermaphrodite: the sea slug Chelidonura hirundinina. This also provides a rare unequivocal example of conditional reciprocity averting a 'tragedy of the commons' in biology.     

Games among cannibals: competition to cannibalize and parent-offspring conflict lead to increased sibling cannibalism

Sibling cannibalism occurs in many species, yet understanding of sibling cannibalism as an adaptation currently lags behind understanding of other antagonistic interactions among siblings. Observed sibling cannibalism phenotypes likely reflect the interaction between competitive games among siblings and parent-offspring conflict. Using a game-theoretic approach, we derive optimal offspring cannibalism behaviour and parental modifiers that limit or facilitate cannibalism. The results are compared to contemporary frequency-independent analysis. With the addition of game interactions among siblings or parent-offspring co-evolution, our model predicts increased cannibalism (compared to the frequency-independent prediction), as offspring compete to eat siblings. When infertile eggs are present--strengthening competition--offspring risk eating viable siblings in order to gain access to infertile eggs, intensifying parent-offspring conflict. We use the results to make new predictions about the occurrence of sibling cannibalism. Additionally, we demonstrate the utility of trophic egg laying as a maternal mechanism to promote egg eating.     

Sexual reproduction and parent-offspring conflict in the RNA "Tumour" virus/host relationship: implications for vertebrate oncogene evolution.

Longmuir and co-workers have reported that respiration of certain tissue slices is approximated by Michaelis-Menten kinetics. From this and other experimental findings, Longmuir proposed that a carrier is involved in tissue oxygen transport. Gold developed a deterministic model to examine this hypothesis. This report presents a stochastic model for a fixed site carrier in a more general framework that includes the stochastic counter-part to Gold's deterministic model as a special case. The kinetics of tissue oxygen consumption predicted by the model are examined for various cases.     

Suppression of reaging antibody formation. IV. Suppression of reaginic antibodies to penicillin in the mouse.

Reaginic antibodies to the benzylpenicilloyl determinant (BPO) and ovalbumin (OA) were induced readily in B6D2F1 mice by a single i.p. injection of either 1 or 10 mug of BPO4-OA suspended with 1 mg of Al(OH)3 in 0.5 ml of saline. Administration of conjugates consisting of the hapten coupled to the isologous, nonimmunogenic murine gamma-globulins (MgammaG), i.e., BPO9-MgammaG, BPO11-MgammaG, or BPO12-MgammaG, resulted in complete and specific suppression of the induction of the anti-BPO reaginic antibody response without affecting, however, the level of reaginic antibodies to OA. Further study of the effect of epitope density on the immunologic properties of BPOX-MgammaG revealed that a) the lightly haptenated conjugated, BPO1-MgammaG and BPO2.9-MgammaG, were not immunosuppressive, b) the conjugates, BPO4.3-MgammaG and BPO19-MgammaG, were partially tolerogenic, and c) the heavily haptenated conjugate, BPO31-MgammaG, was nontolerogenic. Moreover, most importantly, the ongoing anti-BPO response in sensitized mice was readily abrogated by either four daily or four weekly injections of BPO9-MgammaG. The immunosuppressive effect of BPO12-MgammaG conjugates was dose dependent, complete suppression being achieved with 200 mug of the tolerogen. The unresponsiveness to BPO of spleen cells from immunosuppressed donors was also maintained in adoptive cell transfer experiments in spite of the additional administration of the immunizing antigen under conditions expected to yield a secondary IgE response. Hence, it is suggested that, with special precautions to prevent unleashing an anaphylactic shock, treatment of penicillin-sensitive individuals with polyvalent conjugates of an appropriate number of BPO groups per human gamma-globulin molecule would constitute a rational immunotherapeutic procedure for the abrogation of the allergic response to BPO.     

Evolutionary morphology of the caecilian urogenital system. IV. The cloaca

The gross and microscopic anatomy of male and female cloacae of caecilians (Amphibia: Apoda or Gymnophiona) is described and analyzed in terms of structure and function. The arrangement of musculature and cloacal accessory structures is species-specific in males. Contraction of certain cloacal and body wall musculature facilitates eversion of the male cloaca for use as an intromittent organ. The cloacae of females show less marked morphological differences from species to species, and are modified as receptors of male phallodea.     

Evolutionary conflict over the control of offspring sex ratio

There are many theories which predict how animals should control the sex ratio of their offspring. In diploids, however, such control is rarely seen. Two explanations have been suggested for this. One is that parents are simply unable to control the sex ratio of their offspring. The other is that sperm actively oppose such control. This paper examines the possibilities and consequences of parent-gamete conflict over the sex ratio. Such conflict may occur between any of the parties concerned--sperm, ova, fathers, mothers, offspring. It is concluded that gametes are indeed almost always opposed to any parental manipulation of the sex ratio. However, it is probable that the rarity of adaptive parental control of progeny sex ratio in diploids is because parents are physiologically incapable of altering the sex ratio.     

Estimation of heritability by parent-offspring regression

Summary The implications of bias due to previous inbreeding of parents and genotype x environmental interaction on narrow sense heritability (h²) estimates by parent-offspring regression are enumerated. To remove the bias caused by genotype x environment interaction, an analysis of covariance model could be used. In special cases, where phenotypic expression is a result of two organisms interacting, such as in symbiotic N₂ fixation, an analysis of covariance model with a test of heterogeneity of slopes is recommended. When host genotype x strain interactions are significant, separate heritability estimates for each strain are suggested to take advantage of genotype x strain interaction, which may be a major factor contributing to the expression of N₂ fixation traits.