Developmental aspects of kin recognition

The ability to recognise kin requires the individual to possess a variety of abilities. Individuals must produce a cue which indicates relatedness, they must process this cue to determine relatedness and then must act on this cue. Research has concentrated on the first and final stage of this process, i.e., the cues of kinship and kin correlated behaviour. Little attention has been paid to how individuals process cues to determine relatedness. This paper discusses how individuals ‘recognise’ kin, the exhibition of kin correlated behaviour and considers the role of the MHC in these processes. Two broad theories have emerged to explain how individuals recognise their kin: either a recognition gene(s) or some experiential mechanism. In mammals there is no evidence to suggest that recognition (the processing of the cue) is under genetic control but rather is the result of experience, learning about related individuals during development. Moreover studies on kin recognition in the domestic dog suggest that all kin are not recognised by the same process but different classes of kin, parents, siblings may well be recognised using different means. Studies of kin correlated behaviour suggest that the behaviour exhibited towards kin by Mongolian gerbils is mediated by the environment. Factors of environmental familiarity, sex and developmental age all affect the response of individuals to kin and non‐kin. In these situations the ability to recognise kin does not change but the exhibition of kin correlated behaviour changes according to environmental conditions. These studies indicate that kin recognition may not be the ‘unified’ process previously thought and thus any explanations of the proximate and ultimate causation of kin recognition need to encompass this complexity. The question remains of whether the MHC is complex enough to do so. This revised version was published online in July 2006 with corrections to the Cover Date.     

No evidence that relatedness or familiarity modulates male harm in Drosophila melanogaster flies from a wild population

Sexual selection frequently promotes the evolution of aggressive behaviors that help males compete against their rivals, but which may harm females and hamper their fitness. Kin selection theory predicts that optimal male–male competition levels can be reduced when competitors are more genetically related to each other than to the population average, contributing to resolve this sexual conflict. Work in Drosophila melanogaster has spearheaded empirical tests of this idea, but studies so far have been conducted in laboratory‐adapted populations in homogeneous rearing environments that may hamper kin recognition, and used highly skewed sex ratios that may fail to reflect average natural conditions. Here, we performed a fully factorial design with the aim of exploring how rearing environment (i.e., familiarity) and relatedness affect male–male aggression, male harassment, and overall male harm levels in flies from a wild population of Drosophila melanogaster, under more natural conditions. Namely, we (a) manipulated relatedness and familiarity so that larvae reared apart were raised in different environments, as is common in the wild, and (b) studied the effects of relatedness and familiarity under average levels of male–male competition in the field. We show that, contrary to previous findings, groups of unrelated‐unfamiliar males were as likely to fight with each other and harass females than related‐familiar males and that overall levels of male harm to females were similar across treatments. Our results suggest that the role of kin selection in modulating sexual conflict is yet unclear in Drosophila melanogaster, and call for further studies that focus on natural populations and realistic socio‐sexual and ecological environments.     

Recent advance of kin recognition in plant

Plants have the ability to discriminate kin members from strangers in competitive interactions and show altruistic behavior towards related individuals. Studies have showed that plants recognize their neighbors and adjust their ecological strategy mainly through leaf volatiles, root secretions and photographic carrier. The target plants can modify their morphological traits (root size, root:shoot ratio, seed numbers etc.) or metabolism characteristics (secondary metabolites, defense-related proteins etc.) when groups of plants shared common resources, so as to minimize competition with close relatives. The density of kin recognition is influenced by environmental conditions. The main reasons for controversial experimental results of kin recognition are associated with plant materials, standard of kin selection, ecological factors and measured indices. Further studies are required to understand the mechanisms of kin interactions in plants from physiological, biochemical, molecular and metabolic levels.     

植物亲缘识别的研究进展

植物的亲缘识别(kin recognition)指植物通过识别周边个体与自己的亲缘关系, 调整自身的生长生态策略、促进亲缘个体的生存与繁衍。研究表明, 植物主要通过特定的叶片挥发物、根系分泌物、感光载体等途径, 识别周边个体与自己的亲缘关系, 改变自身形态学策略(如根系大小、根冠比、种子数量等)或者生理代谢策略(次生代谢物质、防御蛋白等), 调整与周边个体的竞争强度, 缓和与近亲缘个体之间的竞争, 加强与远亲缘或非亲缘个体的竞争。同时亲缘识别的强度也受环境因子(养分等)的影响。结合目前的研究进展, 该文分析了导致亲缘识别的研究结果存在差异或争议的主要原因, 认为主要与实验材料的选择、亲缘关系的界定标准、环境条件及测定的指标不统一有关。将来的研究应重点从生理生化、分子、代谢水平上深入研究植物亲缘识别的机理。     

Kin recognition by paternal half-siblings in captive Papio cynocephalus

Our objective in this study was to evaluate whether a group of paternally related, subadult baboons (Papio cynocephalus) would preferentially interact with kin or nonkin when they had been raised apart from kin other than their mothers. Subjects and their mothers were removed from the breeding group and placed in alternate housing within 24 h after birth to ensure that the subjects would not have a social history with either their sire or their half-siblings. At 90 days of age, the 23 subjects were separated from their mothers and assigned to a peer–peer social group. Behavioral performance was measured using focal animal sampling techniques and 12 molecular behavioral criteria. Analyses of the data indicate that in dyadic interactions kin did not interact more frequently than nonkin in performance of affiliative, sociosexual, and agonistic behaviors. The hypothesis that baboons recognize kin in the absence of maternal associations was not supported by the data; moreover, we suggest that social learning and social history are the most likely mechanisms for kin recognition. Am. J. Primatol. 43:147–157, 1997. © 1997 Wiley-Liss, Inc.     

Fine‐scale genetic structure and helping decisions in a cooperatively breeding bird

In animal societies, characteristic demographic and dispersal patterns may lead to genetic structuring of populations, generating the potential for kin selection to operate. However, even in genetically structured populations, social interactions may still require kin discrimination for cooperative behaviour to be directed towards relatives. Here, we use molecular genetics and long‐term field data to investigate genetic structure in an adult population of long‐tailed tits Aegithalos caudatus, a cooperative breeder in which helping occurs within extended kin networks, and relate this to patterns of helping with respect to kinship. Spatial autocorrelation analyses reveal fine‐scale genetic structure within our population, such that related adults of either sex are spatially clustered following natal dispersal, with relatedness among nearby males higher than that among nearby females, as predicted by observations of male‐biased philopatry. This kin structure creates opportunities for failed breeders to gain indirect fitness benefits via redirected helping, but crucially, most close neighbours of failed breeders are unrelated and help is directed towards relatives more often than expected by indiscriminate helping. These findings are consistent with the effective kin discrimination mechanism known to exist in long‐tailed tits and support models identifying kin selection as the driver of cooperation.     

Nepotistic cooperation in non-human primate groups

Darwin was struck by the many similarities between humans and other primates and believed that these similarities were the product of common ancestry. He would be even more impressed by the similarities if he had known what we have learned about primates over the last 50 years. Genetic kinship has emerged as the primary organizing force in the evolution of primate social organization and the patterning of social behaviour in non-human primate groups. There are pronounced nepotistic biases across the primate order, from tiny grey mouse lemurs (Microcebus murinus) that forage alone at night but cluster with relatives to sleep during the day, to cooperatively breeding marmosets that rely on closely related helpers to rear their young, rhesus macaque (Macaca mulatta) females who acquire their mother''s rank and form strict matrilineal dominance hierarchies, male howler monkeys that help their sons maintain access to groups of females and male chimpanzees (Pan troglodytes) that form lasting relationships with their brothers. As more evidence of nepotism has accumulated, important questions about the evolutionary processes underlying these kin biases have been raised. Although kin selection predicts that altruism will be biased in favour of relatives, it is difficult to assess whether primates actually conform to predictions derived from Hamilton''s rule: br > c. In addition, other mechanisms, including contingent reciprocity and mutualism, could contribute to the nepotistic biases observed in non-human primate groups. There are good reasons to suspect that these processes may complement the effects of kin selection and amplify the extent of nepotistic biases in behaviour.     

KIN-MEDIATED MORPHOGENESIS IN FACULTATIVELY CANNIBALISTIC TADPOLES

Inclusive fitness theory predicts that organisms can increase their fitness by helping or not harming relatives, and many animals modify their behavior toward kin in a manner consistent with this prediction. Morphogenesis also may be sensitive to kinship environment, particularly in species where certain individuals facultatively develop structures that can be used against conspecifics as weaponry. We tested this hypothesis by examining whether and how consanguinity affected the probability that a structurally distinctive carnivore phenotype, which is opportunistically cannibalistic, would be produced in plains spadefoot toad tadpoles (Spea bombifrons) and southern spadefoot toad tadpoles (S. multiplicata). For tadpoles of S. multiplicata, individuals were significantly more likely to express the carnivore phenotype in mixed sibship groups than in pure sibship groups. For tadpoles of S. bombifrons, individuals were significantly more likely to express the carnivore phenotype when reared alone than in pure sibship groups. Both outcomes were independent of food availability or sibship specific differences in size or growth rate, and waterborne chemical signals from nonkin were sufficient to trigger expression of the carnivore phenotype. Our results suggest that morphogenesis may be responsive to kinship environment in any species or population that occurs as multiple, environmentally induced forms (polyphenism) that differ in their ability to help or to harm others.     

Indirect genetic effects and kin recognition: estimating IGEs when interactions differ between kin and strangers

Social interactions among individuals are widespread, both in natural and domestic populations. As a result, trait values of individuals may be affected by genes in other individuals, a phenomenon known as indirect genetic effects (IGEs). IGEs can be estimated using linear mixed models. The traditional IGE model assumes that an individual interacts equally with all its partners, whether kin or strangers. There is abundant evidence, however, that individuals behave differently towards kin as compared with strangers, which agrees with predictions from kin-selection theory. With a mix of kin and strangers, therefore, IGEs estimated from a traditional model may be incorrect, and selection based on those estimates will be suboptimal. Here we investigate whether genetic parameters for IGEs are statistically identifiable in group-structured populations when IGEs differ between kin and strangers, and develop models to estimate such parameters. First, we extend the definition of total breeding value and total heritable variance to cases where IGEs depend on relatedness. Next, we show that the full set of genetic parameters is not identifiable when IGEs differ between kin and strangers. Subsequently, we present a reduced model that yields estimates of the total heritable effects on kin, on non-kin and on all social partners of an individual, as well as the total heritable variance for response to selection. Finally we discuss the consequences of analysing data in which IGEs depend on relatedness using a traditional IGE model, and investigate group structures that may allow estimation of the full set of genetic parameters when IGEs depend on kin.     

Individuality,kin similarity and experimental playback of contact calls in cooperatively breeding riflemen

Riflemen/tītipounamu (Acanthisitta chloris) are kin-based cooperatively breeding birds, which appear able to recognise their relatives. Here, we investigate the potential for vocalisations to act as recognition cues in riflemen. We identified an appropriate contact call and recorded it at the nest from 19 adult riflemen. Measurements of call characteristics were individually repeatable. In addition, call similarity was significantly correlated with relatedness among all birds and among males. Thus, in principle, these contact calls contain sufficient information for individual recognition of familiar kin, and some assessment of relatedness between unfamiliar birds. To test whether riflemen responded differently to calls of kin, we broadcast calls of relatives and non-relatives as separate treatments in a playback experiment. Focal birds rarely responded aggressively or affiliatively, and their tendency to do so was unrelated to treatment. We conclude that zip calls are suitable kin recognition cues, but whether they are used as such remains unknown.     

Temporal changes in kin structure through a population cycle in a territorial bird, the red grouse Lagopus lagopus scoticus

Populations of red grouse ( Lagopus lagopus scoticus ) undergo regular multiannual cycles in abundance. The 'kinship hypothesis' posits that such cycles are caused by changes in kin structure among territorial males producing delayed density-dependent changes in aggressiveness, which in turn influence recruitment and regulate density. The kinship hypothesis makes several specific predictions about the levels of kinship, aggressiveness and recruitment through a population cycle: (i) kin structure will build up during the increase phase of a cycle, but break down prior to peak density; (ii) kin structure influences aggressiveness, such that there will be a negative relationship between kinship and aggressiveness over the years; (iii) as aggressiveness regulates recruitment and density, there will be a negative relationship between aggressiveness in one year and both recruitment and density in the next; (iv) as kin structure influences recruitment via an affect on aggressiveness, there will be a positive relationship between kinship in one year and recruitment the next. Here we test these predictions through the course of an 8-year cycle in a natural population of red grouse in northeast Scotland, using microsatellite DNA markers to resolve changing patterns of kin structure, and supra-orbital comb height of grouse as an index of aggressiveness. Both kin structure and aggressiveness were dynamic through the course of the cycle, and changing patterns were entirely consistent with the expectations of the kinship hypothesis. Results are discussed in relation to potential drivers of population regulation and implications of dynamic kin structure for population genetics.     

植物的亲缘选择

亲缘选择是指在一个随机交配群体中的个体基于亲缘关系而以一种非随机性的方式相互作用,其作用结果是亲缘个体得到更大的广义适合度。综述了亲缘选择和亲缘竞争两种观点以及各自的试验支持证据;分析了导致亲缘选择试验结果出现分歧的原因,认为这主要是由于对亲缘选择理解上的模糊以及试验设计的不严谨所致。植物间的亲缘选择研究不仅相对较少,对亲缘选择的机制研究更为欠缺,这就造成了目前对此问题在科学认识上出现不少盲点。综合前期研究,提出今后对亲缘选择的研究应该首先界定"亲缘"程度,同时改良试验设计方案,选择多种不同生境下的物种对亲缘选择进行深入研究,并且考虑环境因子对植物亲缘选择的影响。同时,对植物亲缘识别机制的研究应该从生理生化方面出发,通过定性定量地分析探索植物根系分泌物在植物亲缘识别中的作用和作用途径。     

The social evolution of dispersal with public goods cooperation

Selection can favour the evolution of individually costly dispersal if this alleviates competition between relatives. However, conditions that favour altruistic dispersal also mediate selection for other social behaviours, such as public goods cooperation, which in turn is likely to mediate dispersal evolution. Here, we investigate – both experimentally (using bacteria) and theoretically – how social habitat heterogeneity (i.e. the distribution of public goods cooperators and cheats) affects the evolution of dispersal. In addition to recovering the well‐known theoretical result that the optimal level of dispersal increases with genetic relatedness of patch mates, we find both mathematically and experimentally that dispersal is always favoured when average patch occupancy is low, but when average patch occupancy is high, the presence of public goods cheats greatly alters selection for dispersal. Specifically, when public goods cheats are localized to the home patch, higher dispersal rates are favoured, but when cheats are present throughout available patches, lower dispersal rates are favoured. These results highlight the importance of other social traits in driving dispersal evolution.     

Queen number and genetic relatedness in a neotropical wasp, Polybia occidentalis

We found that genetic relatedness among Polybia occidentalisworkers was .26±0.057, a value high enough to make altruisticbehavior by workers relatively easy to explain. This comparativelyhigh level of relatedness can be attributed to close relatednessamong queens of .57±0.077 and to great variation amongcolonies in numbers of queens. The harmonic mean of queen numberis 3.1 queens per colony, which is much lower than the arithmeticmean of 10.6 queens per colony. These results are consistentwith a colony cycle called cyclical oligogyny, that is characterizedby a reduction in queen number from colony initiation to colonyreproduction. We did not find any evidence that one or a fewqueens monopolized egg laying or that there was any inbreeding,both of which have been hypothesized to increase relatednessamong workers. Another factor that can increase relatednessamong workers and the brood they rear is withincolony segregationon the basis of relatedness. We found that combmate pupae aresignificantly more closely related to each other (r = .41) thanthey are to pupae in other combs (r = .33), but we have notinvestigated whether workers take advantage of these relatednesspatterns. This distribution of relatedness among combs willoccur if queens do not lay eggs randomly throughout the nest,but concentrate their egg laying on one or a subset of the availablecombs.     

Olfactory kin recognition in a songbird

The ability to recognize close relatives in order to cooperate or to avoid inbreeding is widespread across all taxa. One accepted mechanism for kin recognition in birds is associative learning of visual or acoustic cues. However, how could individuals ever learn to recognize unfamiliar kin? Here, we provide the first evidence for a novel mechanism of kin recognition in birds. Zebra finch (Taeniopygia guttata) fledglings are able to distinguish between kin and non-kin based on olfactory cues alone. Since olfactory cues are likely to be genetically based, this finding establishes a neglected mechanism of kin recognition in birds, particularly in songbirds, with potentially far-reaching consequences for both kin selection and inbreeding avoidance.     

Mating structure and genetic relatedness among gynes in the primitively eusocial wasp Polistes snelleni (Hymenoptera: Vespidae)

Mating structure and genetic relatedness among gynes (potential reproductive females) of the paper wasp Polistes snelleni were investigated using DNA microsatellite markers. All colonies had one foundress inseminated by a single male, and no sign of inbreeding was detected. The mean genetic relatedness among gynes was 0.734 ± 0.028 (SE), which is not significantly different from the expected value of 0.75 for full sisters.     

Kin recognition in an annual plant

Kin recognition is important in animal social systems. However, though plants often compete with kin, there has been as yet no direct evidence that plants recognize kin in competitive interactions. Here we show in the annual plant Cakile edentula, allocation to roots increased when groups of strangers shared a common pot, but not when groups of siblings shared a pot. Our results demonstrate that plants can discriminate kin in competitive interactions and indicate that the root interactions may provide the cue for kin recognition. Because greater root allocation is argued to increase below-ground competitive ability, the results are consistent with kin selection.     

Low levels of relatedness on black grouse leks despite male philopatry

In lekking species, males cluster on specific areas for display (the leks) and females generally prefer to copulate with males on large aggregations. The maintenance of leks in which only a few males reproduce might be explained if subordinate males gain indirect fitness benefits. By joining a lek on which relatives are displaying, subordinates might attract more females to the lek thereby increasing the mating opportunities of their kin. In black grouse, a genetic structure among leks has previously been found suggesting that relatives could display together. Using 11 microsatellite loci, we extended this result by testing for the presence of kin structures in nine black grouse leks (101 males). The genetic differentiation among flocks was higher in males than in females, suggesting female-biased dispersal and male philopatry. Because of this genetic structure, males were more related within than among leks. However, the mean relatedness within each lek hardly differed from zero. The lekking males were not more related than random assortments of males from the winter flocks and there were no kin clusters within leks. Thus, black grouse males do not choose to display with and close to relatives. Male philopatry alone was not sufficient to induce elevated levels of relatedness on the leks either because of male partial dispersal or a rapid turnover of the successful males. The indirect fitness benefits associated with males' settlement decision are probably limited compared to the direct benefits of joining large aggregations such as increased current and future mating opportunities.     

Female rhesus macaques discriminate unfamiliar paternal sisters in playback experiments: support for acoustic phenotype matching

Widespread evidence exists that when relatives live together, kinship plays a central role in shaping the evolution of social behaviour. Previous studies showed that female rhesus macaques (Macaca mulatta) recognize familiar maternal kin using vocal cues. Recognizing paternal kin might, however, be more difficult as rhesus females mate promiscuously during the possible conception period, most probably concealing paternity. Behavioural observations indicate that semi free-ranging female rhesus macaques prefer to associate with their paternal half-sisters in comparison to unrelated females within the same group, particularly when born within the same age cohort. However, the cues and mechanism/s used in paternal kin discrimination remain under debate. Here, we investigated whether female rhesus macaques use the acoustic modality to discriminate between paternal half-sisters and non-kin, and tested familiarity and phenotype matching as the underlying mechanisms. We found that test females responded more often to calls of paternal half-sisters compared with calls of unrelated females, and that this discrimination ability was independent of the level of familiarity between callers and test females, which provides, to our knowledge, the first evidence for acoustic phenotype matching. Our study strengthens the evidence that female rhesus macaques can recognize their paternal kin, and that vocalizations are used as a cue.