Nepotism vs Tit-For-Tat,or, why should you be nice to your rotten brother?

Summary It is well known that interactions among relatives facilitate the evolution of altruistic behaviours. Game theoretic models show, however, that guarded altruism (such as Tit-For-Tat) can evolve among non-relatives when individuals interact many times and cheating behaviours can be punished. Strangely, no one has yet asked whether the guarded altruism that evolves among non-relatives might also evolve among close relatives, supplanting unconditional altriusm. We present a series of one-locus sexual haploid models in which Tit-For-Tat, unconditional altruists and selfish individuals interact in groups of full siblings. Tit-For-Tat frequently (but not always) replaced unguarded altruism, in which case the strategic rules for interacting with kin vs non-kin are identical. Even when Tit-For-Tat is selected at a single locus, however, by withholding altruism for non-reciprocating relatives it may qualify as an outlaw from the standpoint of modifier genes at other loci.     

Selective pressures for accurate altruism targeting: evidence from digital evolution for difficult-to-test aspects of inclusive fitness theory

Inclusive fitness theory predicts that natural selection will favour altruist genes that are more accurate in targeting altruism only to copies of themselves. In this paper, we provide evidence from digital evolution in support of this prediction by competing multiple altruist-targeting mechanisms that vary in their accuracy in determining whether a potential target for altruism carries a copy of the altruist gene. We compete altruism-targeting mechanisms based on (i) kinship (kin targeting), (ii) genetic similarity at a level greater than that expected of kin (similarity targeting), and (iii) perfect knowledge of the presence of an altruist gene (green beard targeting). Natural selection always favoured the most accurate targeting mechanism available. Our investigations also revealed that evolution did not increase the altruism level when all green beard altruists used the same phenotypic marker. The green beard altruism levels stably increased only when mutations that changed the altruism level also changed the marker (e.g. beard colour), such that beard colour reliably indicated the altruism level. For kin- and similarity-targeting mechanisms, we found that evolution was able to stably adjust altruism levels. Our results confirm that natural selection favours altruist genes that are increasingly accurate in targeting altruism to only their copies. Our work also emphasizes that the concept of targeting accuracy must include both the presence of an altruist gene and the level of altruism it produces.     

Selfish long-term benefits of hoarding in the Siberian jay

We followed the consumption of cached food in Siberian jay flocksduring winter. Siberian jay flocks contain kin associationscomposed of parents in company with mature offspring. Further,flocks often contain immigrants not closely related to the groupmembers. We studied the extent to which Siberian jay offspringkept caches to themselves or whether they showed a kin biasand shared cached food mutualistically with relatives (parents/siblings)thus making an inclusive fitness gain. We recorded the retrievalof cached food using radio-ptilochronology, a technique thatdetects the consumption of a radioactively labeled food itemin the growth bar laid down in a growing feather the day ofconsumption. Food caching entailed a selfish benefit to thehoarder. The hoarding bird showed a substantial "recovery advantage"and retrieved its own cached food seven times as often as anyof its flock mates. There was no evidence for mutualistic sharingof caches among relatives. This selfish benefit persisted throughoutthe 7-week period for which we could follow the consumptionof labeled food.     

Selective recovery of founder genetic diversity in aquacultural broodstocks and captive, endangered fish populations

Hatchery broodstocks used for genetic conservation or aquaculture may represent their ancestral gene pools rather poorly. This is especially likely when the fish that found a broodstock are close relatives of each other. We re-analysed microsatellite data from a breeding experiment on red sea bream to demonstrate how lost genetic variation might be recovered when gene frequencies have been distorted by consanguineous founders in a hatchery. A minimal-kinship criterion based on a relatedness estimator was used to select subsets of breeders which represented the maximum number of founder lineages (i.e., carried the fewest identical copies of ancestral genes). UPGMA clustering of Nei's genetic distances grouped these selected subsets with the parental gene pool, rather than with the entire, highly drifted offspring generation. The selected subsets also captured much of the expected heterozygosity and allelic diversity of the parental gene pool. Independent pedigree data on the same fish showed that the selected subsets had more contributing parents and more founder equivalents than random subsets of the same size. The estimated mean coancestry was lower in the selected subsets, meaning that inbreeding in subsequent generations would be lower if they were used as breeders. The procedure appears suitable for reducing the genetic distortion due to consanguineous and over-represented founders of a hatchery gene pool.     

The evolution of senescence in multi-component systems

Actuarial senescence is characterized by an increase in mortality rate with increasing chronological age. The reliability theory of senescence proposes that organisms’ vital functions can be modelled as a suite of damageable, irreplaceable elements (typically genes or their products) that protect their bearer from condition-dependent death so long as at least one of the elements remains intact. Current incarnations of the reliability theory of senescence are continuous-time models with no explicit evolutionary component. Here, we use elementary probability theory and evolutionary dynamics analysis to derive a discrete-time version of the reliability theory of senescence. We include three variations on this theme: the ‘Series’ model in which damage to any of n elements results in death, the ‘Parallel’ model, in which damage accumulates in random order and damage to all n elements results in death, and the ‘Cascade’ (multi-stage) model, which is like the Parallel model, except the irreparable damage necessarily follows a strict sequence. For simplicity, we refer to the state of having multiple elements as ‘redundancy’, but this does not imply that the elements are necessarily identical. We show that redundancy leads to actuarial senescence in the Parallel and Cascade models but not in the Series model. We further demonstrate that in the Parallel and Cascade models, lifetime reproductive output (a potential proxy for fitness in populations with discrete generations) is a positive but decelerating function of redundancy. The positive nature of the fitness function leads to the prediction that redundancy and senescence should evolve from non-redundant, non-senescing ancestral populations; however, the deceleration of the fitness function leads to the prediction that this evolution towards increased redundancy will eventually be limited by mutation-selection balance. Using evolutionary dynamics analysis involving the discrete-generation quasispecies equation, we confirm these two predictions. Finally, we show that a population's equilibrium redundancy is sensitive to the environmental conditions that prevailed during its evolution, such as the rate of extrinsic mortality.     

Polyandrous females found fitter populations

Multiple mating by females (polyandry) requires an evolutionary explanation, because it carries fitness costs in many species. When mated females disperse alone to a new habitat, their offspring may have no option but to mate with their siblings and incur inbreeding depression. However, some of the offspring of polyandrous females may only be half siblings, reducing inbreeding depression when isolated groups of siblings only have each other as mates. We investigated this putative benefit of polyandry over monandry by initiating multiple genetically isolated populations of Callosobruchus maculatus beetles, each founded by a single female, who received a complete ejaculate from either one or two males. The early generations had comparable fitness, but the F4 and F5 descendants of doubly inseminated females were more numerous and had higher egg‐to‐adult survival than the descendants of singly inseminated females. This fitness benefit was of similar magnitude whether beetles were reared on their standard food plant, or on a less favourable food source. Our results suggest that polyandrous females produce fitter descendants in inbred founder populations and therefore that polyandry may affect movement ecology and invasion biology.     

Maternal condition and facultative sex ratios in populations with overlapping generations

Facultative investment in offspring sex is related to maternal condition in many organisms. In mammals, empirical support for condition-dependent sex allocation is equivocal, and there is some doubt as to theoretical expectations. Much theory has been developed to make predictions for condition-dependent sex ratios in populations with discrete generations. However, the extension of these predictions to populations with overlapping generations (OLGs; e.g., mammals) has been limited, leaving doubt as to the specific prediction for maternal-condition-dependent sex ratios in mammals. We develop a population genetics model that incorporates maternal effects on multiple offspring fitness components in a population with OLGs. Using a rare-gene and evolutionarily stable strategy approach, we demonstrate that sex ratio predictions of this model are identical to those for equivalent discrete generations models. We show that the predicted sex ratios depend on the sex-specific ratio of R(o) (offspring lifetime fitness) for offspring of good and poor mothers. This offspring lifetime fitness rule indicates that empirical research on conditional sex ratios should consider all three components of offspring R(o) (juvenile survival, adult life span, and fertility).     

Population biology of a selfish sex ratio distorting element in a booklouse (Psocodea: Liposcelis)

Arthropods harbour a variety of selfish genetic elements that manipulate reproduction to be preferentially transmitted to future generations. A major ongoing question is to understand how these elements persist in nature. In this study, we examine the population dynamics of an unusual selfish sex ratio distorter in a recently discovered species of booklouse, Liposcelis sp. (Psocodea: Liposcelididae) to gain a better understanding of some of the factors that may affect the persistence of this element. Females that carry the selfish genetic element only ever produce daughters, although they are obligately sexual. These females also only transmit the maternal half of their genome. We performed a replicated population cage experiment, varying the initial frequency of females that harbour the selfish element, and following female frequencies for 20 months. The selfish genetic element persisted in all cages, often reaching very high (and thus severely female‐biased) frequencies. Surprisingly, we also found that females that carry the selfish genetic element had much lower fitness than their nondistorter counterparts, with lower lifetime fecundity, slower development and a shorter egg‐laying period. We suggest that differential fitness plays a role in the maintenance of the selfish genetic element in this species. We believe that the genetic system in this species, paternal genome elimination, which allows maternal control of offspring sex ratio, may also be important in the persistence of the selfish genetic element, highlighting the need to consider species with diverse ecologies and genetic systems when investigating the effects of sex ratio manipulators on host populations.     

Kin selection,kin avoidance and correlated strategies

Summary Kin selection of correlated strategies is examined for both weak and strong altruism under simple haploid inheritance. While kin assortment enhances the range of evolutionary stability for (strongly altruistic) correlated strategies (defined herein), kin avoidance is possible under a weakly altruistic correlated strategy. When social competition induces role assignments of variable fitness, group mates may prefer association with non-relatives. Even when group life is mandatory, an individual may accept the risk of abandonment (and reproductive death) rather then associate with kin: a competitive superior may behave altruistically by permitting competitively inferior kin to emigrate. Thus, kin selection and social competition are not necessarily mutually supportive processes within groups. I conclude by interpreting dominance as a strongly altruistic correlated strategy in two social hymenopteran contexts.     

Parent and offspring genotypes influence gene expression in early life

Parents can have profound effects on offspring fitness. Little, however, is known about the mechanisms through which parental genetic variation influences offspring physiology in natural systems. White‐throated sparrows (Zonotrichia albicollis, WTSP) exist in two genetic morphs, tan and white, controlled by a large polymorphic supergene. Morphs mate disassortatively, resulting in two pair types: tan male × white female (T × W) pairs, which provide biparental care and white male × tan female (W × T) pairs, which provide female‐biased care. To investigate how parental composition impacts offspring, we performed RNA‐seq on whole blood of WTSP nestlings sampled from nests of both pair types. Parental pair type had a large effect on nestling gene expression, with 881 genes differentially expressed (DE) and seven correlated gene coexpression modules. The DE genes and modules expressed at higher levels in W × T nests with female‐biased parental care function in metabolism and stress‐related pathways resulting from the overrepresentation of proteolysis and stress‐response genes (e.g., SOD2, NR3C1). These results show that parental genotypes and/or associated behaviours influence nestling physiology, and highlight avenues of further research investigating the ultimate implications for the maintenance of this polymorphism. Nestlings also exhibited morph‐specific gene expression, with 92 differentially expressed genes, comprising immunity genes and genes encompassed by the supergene. Remarkably, we identified the same regulatory hub genes in these blood‐derived expression networks as were previously identified in adult WTSP brains (EPM2A, BPNT1, TAF5L). These hub genes were located within the supergene, highlighting the importance of this gene complex in structuring regulatory networks across diverse tissues.     

Reproductive bribing and policing as evolutionary mechanisms for the suppression of within-group selfishness

We show that a new, simple, and robust general mechanism for the social suppression of within-group selfishness follows from Hamilton's rule applied in a multilevel selection approach to asymmetrical, two-person groups: If it pays a group member to behave selfishly (i.e., increase its share of the group's reproduction, at the expense of group productivity), then its partner will virtually always be favored to provide a reproductive "bribe" sufficient to remove the incentive for the selfish behavior. The magnitude of the bribe will vary directly with the number of offspring (or other close kin) potentially gained by the selfish individual and inversely with both the relatedness r between the interactants and the loss in group productivity because of selfishness. This bribe principle greatly extends the scope for cooperation within groups. Reproductive bribing is more likely to be favored over social policing for dominants rather than subordinates and as intragroup relatedness increases. Finally, analysis of the difference between the group optimum for an individual's behavior and the individual's inclusive fitness optimum reveals a paradoxical feedback loop by which bribing and policing, while nullifying particular selfish acts, automatically widen the separation of individual and group optima for other behaviors (i.e., resolution of one conflict intensifies others).     

Selection for altruism through random drift in variable size populations

ABSTRACT: BACKGROUND: Altruistic behavior is defined as helping others at a cost to oneself and a lowered fitness. The lower fitness implies that altruists should be selected against, which is in contradiction with their widespread presence is nature. Present models of selection for altruism (kin or multilevel) show that altruistic behaviors can have 'hidden' advantages if the 'common good' produced by altruists is restricted to some related or unrelated groups. These models are mostly deterministic, or assume a frequency dependent fitness. RESULTS: Evolutionary dynamics is a competition between deterministic selection pressure and stochastic events due to random sampling from one generation to the next. We show here that an altruistic allele extending the carrying capacity of the habitat can win by increasing the random drift of "selfish" alleles. In other terms, the fixation probability of altruistic genes can be higher than those of a selfish ones, even though altruists have a smaller fitness. Moreover when populations are geographically structured, the altruists advantage can be highly amplified and the fixation probability of selfish genes can tend toward zero. The above results are obtained both by numerical and analytical calculations. Analytical results are obtained in the limit of large populations. CONCLUSIONS: The theory we present does not involve kin or multilevel selection, but is based on the existence of random drift in variable size populations. The model is a generalization of the original Fisher-Wright and Moran models where the carrying capacity depends on the number of altruists.     

Parental effort and response to nestling begging in the house sparrow: repeatability,heritability and parent–offspring co‐evolution

Parental effort has a direct impact on individual fitness. Theoretical models exploring how parental effort evolves to cope with offspring demand and sexual conflicts may differ in the assumptions they make in respect to the genetic heritability of parental behaviours. Only a few attempts, however, have been made to estimate the heritability of parental behaviours and their possible co‐evolution with offspring solicitation behaviour. Analysing parent and offspring behaviours in four generations of cross‐fostered broods of house sparrows, we found that parental effort (food delivery rate) was repeatable across consecutive broods and heritable across generations. In contrast, parental response to experimentally induced changes in nestling begging was neither repeatable across broods nor heritable across generations or correlated to nestling begging. Thus, the results give no indication for genetic covariance between begging intensity and parental response, but provide the first cross‐fostering‐based evidence for the heritability of parental investment levels across generations.     

Male house mice produce fewer offspring with lower viability and poorer performance when mated with females they do not prefer

We report experimental results consistent with the hypothesis that constraints on the expression of male mating preferences affect breeder fitness, offspring viability and performance. If constraints on the expression of mating preferences are common, tests of fitness variation associated with mate preferences must eliminate as many constraints on mate preferences as possible. We tested whether male mate preferences influenced breeder fitness, offspring performance and viability in typically polygynous house mice, Mus domesticus, from a feral source population. Our ‘free mate choice’ trials not only eliminated female preferences, male-male and female-female competition, but also our best guesses of the traits mediating choosers' preferences. Males mated with their preferred (P) females sired more litters than males mated with their nonpreferred (NP) females. Offspring viability was significantly lower when males' reproduced with females they did not prefer compared with females they did prefer. Adult sons of males that mated with their P females were socially dominant to sons of males that mated with their NP females. Adult offspring from P pairings built better nests than offspring from NP pairings. The slope of the survivorship curve for P offspring was significantly higher than for NP offspring. These results showed (1) males' mate preferences affected their fitness, (2) males that mated with females they preferred produced more litters than males that mated with females they did not prefer, and (3) their offspring were significantly more viable and performed significantly better on standardized performance tests. This is the first demonstration of fitness benefits of male choice behaviour in a mammal species with typical paternal investment. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

The complete mitochondrial genome of the leafminer Liriomyza sativae (Diptera: Agromyzidae): great difference in the A+T-rich region compared to Liriomyza trifolii

The complete mitochondrial genome sequence of Liriomyza sativae Blanchard (15,551 bp) was determined and analyzed in this study. The circular genome contained 37 genes including 13 protein-coding genes, 22 tRNA genes, 2 rRNA genes and an A + T-rich region. The initiation codons of COI and ND1 were ‘ATCA’ and ‘GTG’, respectively. ND2 gene used the truncated termination codon ‘T’. All the tRNA genes had the typical cloverleaf secondary structures except for tRNA^Ser(AGN) gene, which was found with the absence of a DHU arm. In addition, a tRNA-like secondary structure (tRNA^Met) was found in the A + T-rich region. The great difference was that the length of L. sativae A + T-rich region was 597 bp shorter than that of Liriomyza trifolii (Burgess). Meanwhile, some minor differences such as ‘TATA’ block were also observed in L. sativae in contrast to ‘TACA’ block in L. trifolii. There were also some essential structure elements such as ‘TATA’ block, ‘G(A)_nT’ block, poly-T stretch and stem-and-loop structure in the A + T-rich region of L. sativae mitochondrial genome.     

Molecular characterization and expression analysis of ubiquitin-activating enzyme E1 gene in Citrus reticulata

Ubiquitin-activating enzyme E1 (UBE1) catalyzes the first step in the ubiquitination reaction, which targets a protein for degradation via a proteasome pathway. UBE1 plays an important role in metabolic processes. In this study, full-length cDNA and DNA sequences of UBE1 gene, designated CrUBE1, were obtained from ‘Wuzishatangju’ (self-incompatible, SI) and ‘Shatangju’ (self-compatible, SC) mandarins. 5 amino acids and 8 bases were different in cDNA and DNA sequences of CrUBE1 between ‘Wuzishatangju’ and ‘Shatangju’, respectively. Southern blot analysis showed that there existed only one copy of the CrUBE1 gene in genome of ‘Wuzishatangju’ and ‘Shatangju’. The temporal and spatial expression characteristics of the CrUBE1 gene were investigated using semi-quantitative RT-PCR (SqPCR) and quantitative real-time PCR (qPCR). The expression level of the CrUBE1 gene in anthers of ‘Shatangju’ was approximately 10-fold higher than in anthers of ‘Wuzishatangju’. The highest expression level of CrUBE1 was detected in pistils at 7 days after self-pollination of ‘Wuzishatangju’, which was approximately 5-fold higher than at 0 h. To obtain CrUBE1 protein, the full-length cDNA of CrUBE1 genes from ‘Wuzishatangju’ and ‘Shatangju’ were successfully expressed in Pichia pastoris. Pollen germination frequency of ‘Wuzishatangju’ was significantly inhibited with increasing of CrUBE1 protein concentrations from ‘Wuzishatangju’.     

The role of kin recognition in the evolution of conspecific brood parasitism

Conspecific brood parasitism (CBP) is a common strategy in several species of birds. Currently, some studies suggest that relatedness between host and parasite enhances CBP, since indirect fitness benefits could select for acceptance of related eggs by hosts. Conversely, parasites should avoid laying eggs in nests of relatives if this is costly for the host. Based on the latter argument, kinship should not promote brood parasitism. A recent model clarified this relationship, and showed that kinship can promote brood parasitism, assuming kin recognition. However, in that model kin recognition was assumed perfect. Here we present a model that addresses the role of relatedness and kin selection in CBP, when kin recognition is not perfect and hosts do not always detect parasitism. We consider both the indirect fitness of the parasite and the possible responses of the host. Our results indicate that the existence and accuracy of a kin recognition system is crucial to the final outcome. When CBP represents a cost to the host, a parasitic female that has the choice should avoid parasitizing relatives, unless (1) the costs are not too high and (2) hosts can accurately enough recognize eggs laid by relatives, rejecting them less often than eggs laid by nonkin. But if ‘parasitism’ enhances the direct fitness of the host (which is possible in species with precocial young) parasites should choose relatives whenever possible, even if hosts do not recognize kin eggs. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Conditions for sympatric speciation: A diploid model incorporating habitat fidelity and non-habitat assortative mating

Summary Three types of genes have been proposed to promote sympatric speciation: habitat preference genes, assortative mating genes and habitat-based fitness genes. Previous computer models have analysed these genes separately or in pairs. In this paper we describe a multilocus model in which genes of all three types are considered simultaneously. Our computer simulations show that speciation occurs in complete sympatry under a broad range of conditions. The process includes an initial diversification phase during which a slight amount of divergence occurs, a quasi-equilibrium phase of stasis during which little or no detectable divergence occurs and a completion phase during which divergence is dramatic and gene flow between diverging habitat morphs is rapidly eliminated. Habitat preference genes and habitat-specific fitness genes become associated when assortative mating occurs due to habitat preference, but interbreeding between individuals adapted to different habitats occurs unless habitat preference is almost error free. However, nonhabitat assortative mating, when coupled with habitat preference can eliminate this interbreeding. Even when several loci contribute to the probability of expression of non-habitat assortative mating and the contributions of individual loci are small, gene flow between diverging portions of the population can terminate within less than 1000 generations.     

An evolutionary analysis of the relationship between spite and altruism

We investigate the selective pressures on a social trait when evolution occurs in a population of constant size. We show that any social trait that is spiteful simultaneously qualifies as altruistic. In other words, any trait that reduces the fitness of less related individuals necessarily increases that of related ones. Our analysis demonstrates that the distinction between "Hamiltonian spite" and "Wilsonian spite" is not justified on the basis of fitness effects. We illustrate this general result with an explicit model for the evolution of a social act that reduces the recipient's survival ("harming trait"). This model shows that the evolution of harming is favoured if local demes are of small size and migration is low (philopatry). Further, deme size and migration rate determine whether harming evolves as a selfish strategy by increasing the fitness of the actor, or as a spiteful/altruistic strategy through its positive effect on the fitness of close kin.