Queen activation of lazy workers in colonies of the eusocial naked mole-rat.

Evolutionary conflicts of interest are expected to arise in genetically diverse social groups. In eusocial insect societies, a potential conflict exists between the queen and her workers over how active the workers should be, and evidence exists that queen aggression increases activity levels of her lazier workers. Here I provide evidence that queen aggression (shoving) in laboratory colonies of the eusocial mammal, the naked mole-rat (Heterocephalus glaber), is a convergently evolved manifestation of queen-worker conflict over worker activity. Queen-initiated shoves activate inherently lazy workers, which tend to be larger and/or less related to the queen than are infrequently shoved, industrious workers. In addition, queen removal selectively depresses the activity of workers that are larger and less related to her. Finally, queen shoving and worker inactivity are pronounced when colonies are satiated but not when colonies are hungry, indicating that the underlying 'work-conflict' is highly context-specific.     

Fitness benefits of prolonged post-reproductive lifespan in women

Most animals reproduce until they die, but in humans, females can survive long after ceasing reproduction. In theory, a prolonged post-reproductive lifespan will evolve when females can gain greater fitness by increasing the success of their offspring than by continuing to breed themselves. Although reproductive success is known to decline in old age, it is unknown whether women gain fitness by prolonging lifespan post-reproduction. Using complete multi-generational demographic records, we show that women with a prolonged post-reproductive lifespan have more grandchildren, and hence greater fitness, in pre-modern populations of both Finns and Canadians. This fitness benefit arises because post-reproductive mothers enhance the lifetime reproductive success of their offspring by allowing them to breed earlier, more frequently and more successfully. Finally, the fitness benefits of prolonged lifespan diminish as the reproductive output of offspring declines. This suggests that in female humans, selection for deferred ageing should wane when one's own offspring become post-reproductive and, correspondingly, we show that rates of female mortality accelerate as their offspring terminate reproduction.     

Competition induces adaptive shifts in caste ratios of a polyembryonic wasp

An important transition in insect life-history evolution was the shift from a solitary existence to living in groups comprising specialized castes. Caste-forming species produce some individuals that reproduce and others with worker functions that have few or no offspring. Morphologically specialized castes are well known in eusocial species like ants and termites, but castes have also evolved in less-studied groups like thrips, aphids and polyembryonic wasps. Because selection acts at both the individual and the colony level, ratios of investment in different castes are predicted to vary with environmental factors like competition and resources. However, experimental evidence for adaptive shifts in caste ratios is limited owing to the experimental difficulty of manipulating factors thought to influence caste ratios, and because some species produce behaviourally flexible castes that switch tasks in response to colony needs. Unlike other caste-forming species, the broods of polyembryonic wasps develop clonally, so that increased production of one caste probably results in decreased production of the other. Here we show that the polyembryonic wasp Copidosoma floridanum alters caste ratios in response to interspecific competition. Our results reveal a distinct trade-off by C. floridanum between reproduction and defence, and show experimentally that caste ratios shift in an adaptive manner.     

Clonal reproduction by males and females in the little fire ant

Sexual reproduction can lead to major conflicts between sexes and within genomes. Here we report an extreme case of such conflicts in the little fire ant Wasmannia auropunctata. We found that sterile workers are produced by normal sexual reproduction, whereas daughter queens are invariably clonally produced. Because males usually develop from unfertilized maternal eggs in ants and other haplodiploid species, they normally achieve direct fitness only through diploid female offspring. Hence, although the clonal production of queens increases the queen's relatedness to reproductive daughters, it potentially reduces male reproductive success to zero. In an apparent response to this conflict between sexes, genetic analyses reveal that males reproduce clonally, most likely by eliminating the maternal half of the genome in diploid eggs. As a result, all sons have nuclear genomes identical to those of their father. The obligate clonal production of males and queens from individuals of the same sex effectively results in a complete separation of the male and female gene pools. These findings show that the haplodiploid sex-determination system provides grounds for the evolution of extraordinary genetic systems and new types of sexual conflict.     

Parasitic Cape honeybee workers, Apis mellifera capensis, evade policing.

Relocation of the Cape honeybee, Apis mellifera capensis, by bee-keepers from southern to northern South Africa in 1990 has caused widespread death of managed African honeybee, A. m. scutellata, colonies. Apis mellifera capensis worker bees are able to lay diploid, female eggs without mating by means of automictic thelytoky (meiosis followed by fusion of two meiotic products to restore egg diploidy), whereas workers of other honeybee subspecies are able to lay only haploid, male eggs. The A. m. capensis workers, which are parasitizing and killing A. m. scutellata colonies in northern South Africa, are the asexual offspring of a single, original worker in which the small amount of genetic variation observed is due to crossing over during meiosis (P. Kryger, personal communication). Here we elucidate two principal mechanisms underlying this parasitism. Parasitic A. m. capensis workers activate their ovaries in host colonies that have a queen present (queenright colonies), and they lay eggs that evade being killed by other workers (worker policing)-the normal fate of worker-laid eggs in colonies with a queen. This unique parasitism by workers is an instance in which a society is unable to control the selfish actions of its members.     

Insurance-based advantage to helpers in a tropical hover wasp

The origin and maintenance of eusociality is a central problem in evolutionary biology. Eusocial groups contain individuals that forfeit their own reproduction in order to help others reproduce. In facultatively eusocial taxa, offspring can choose whether to found new nests or become helpers in their natal groups. In many facultatively eusocial insects, offspring need continuous care during development, but adult carers have life expectancies shorter than the developmental period. When a lone foundress dies, her partly reared brood are usually doomed. Here, we show that helpers in a tropical hover wasp (Liostenogaster flavolineata) have an insurance-based advantage over lone foundresses because after a helper dies, most of the brood that she has partly reared will be brought to maturity by surviving nest-mates. After some of the helpers are experimentally removed from a multi-female nest, the reduced group is left with more brood than it would normally rear. We found that larger, more valuable extra brood were reared through to maturity, but not smaller, less valuable brood. Smaller brood may be sacrificed to feed larger brood, and reduced groups probably benefited from increased short-term helper recruitment. Rearing extra brood did not increase adult mortality or brood development time.     

Parthenogenesis in Komodo dragons

Parthenogenesis, the production of offspring without fertilization by a male, is rare in vertebrate species, which usually reproduce after fusion of male and female gametes. Here we use genetic fingerprinting to identify parthenogenetic offspring produced by two female Komodo dragons (Varanus komodoensis) that had been kept at separate institutions and isolated from males; one of these females subsequently produced additional offspring sexually. This reproductive plasticity indicates that female Komodo dragons may switch between asexual and sexual reproduction, depending on the availability of a mate--a finding that has implications for the breeding of this threatened species in captivity. Most zoos keep only females, with males being moved between zoos for mating, but perhaps they should be kept together to avoid triggering parthenogenesis and thereby decreasing genetic diversity.     

Females increase offspring heterozygosity and fitness through extra-pair matings

Females in a variety of species commonly mate with multiple males, and there is evidence that they benefit by producing offspring of higher genetic quality; however, the nature of these genetic benefits is debated. Enhanced offspring survival or quality can result from intrinsic effects of paternal genes---'good genes'--or from interactions between the maternal and paternal genomes--'compatible genes'. Evidence for the latter process is accumulating: matings between relatives lead to decreased reproductive success, and the individual level of inbreeding--measured as average heterozygosity--is a strong fitness predictor. Females should thus benefit from mating with genetically dissimilar males. In many birds, social monogamy restricts mate choice, but females may circumvent this by pursuing extra-pair copulations. Here we show that female blue tits, Parus caeruleus, increase the heterozygosity of their progeny through extra-pair matings. Females thereby produce offspring of higher reproductive value, because less inbred individuals have increased survival chances, a more elaborate male secondary sexual trait (crown colour) and higher reproductive success. The cost of inbreeding may therefore be an important factor driving the evolution of female extra-pair mating.     

Intrasexual competition and sexual selection in cooperative mammals

In most animals, the sex that invests least in its offspring competes more intensely for access to the opposite sex and shows greater development of secondary sexual characters than the sex that invests most. However, in some mammals where females are the primary care-givers, females compete more frequently or intensely with each other than males. A possible explanation is that, in these species, the resources necessary for successful female reproduction are heavily concentrated and intrasexual competition for breeding opportunities is more intense among females than among males. Intrasexual competition between females is likely to be particularly intense in cooperative breeders where a single female monopolizes reproduction in each group. Here, we use data from a twelve-year study of wild meerkats (Suricata suricatta), where females show high levels of reproductive skew, to show that females gain greater benefits from acquiring dominant status than males and traits that increase competitive ability exert a stronger influence on their breeding success. Females that acquire dominant status also develop a suite of morphological, physiological and behavioural characteristics that help them to control other group members. Our results show that sex differences in parental investment are not the only mechanism capable of generating sex differences in reproductive competition and emphasize the extent to which competition for breeding opportunities between females can affect the evolution of sex differences and the operation of sexual selection.     

Complex social behaviour derived from maternal reproductive traits

A fundamental goal of sociobiology is to explain how complex social behaviour evolves, especially in social insects, the exemplars of social living. Although still the subject of much controversy, recent theoretical explanations have focused on the evolutionary origins of worker behaviour (assistance from daughters that remain in the nest and help their mother to reproduce) through expression of maternal care behaviour towards siblings. A key prediction of this evolutionary model is that traits involved in maternal care have been co-opted through heterochronous expression of maternal genes to result in sib-care, the hallmark of highly evolved social life in insects. A coupling of maternal behaviour to reproductive status evolved in solitary insects, and was a ready substrate for the evolution of worker-containing societies. Here we show that division of foraging labour among worker honey bees (Apis mellifera) is linked to the reproductive status of facultatively sterile females. We thereby identify the evolutionary origin of a widely expressed social-insect behavioural syndrome, and provide a direct demonstration of how variation in maternal reproductive traits gives rise to complex social behaviour in non-reproductive helpers.     

Female choice selects for a viability-based male trait in pheasants

Recent theory on sexual selection suggests that females in species without paternal care choose mates by their secondary sexual characters because these indicate genotypic quality which will be transmitted to the offspring. These ideas are not yet empirically supported as data quantifying the relationship between female mate choice and female reproductive success are lacking. Only in one case, in Colias butterflies, has it been demonstrated unequivocally that females choose 'good genotypes' as mates and there is only one study, on Drosophila, demonstrating that mate choice increases one component of offspring fitness. Spur length of male pheasants (Phasianus colchicus) correlates with various fitness-related properties. We here present the first experimental field data showing that female pheasants select mates on the basis of male spur length and that female mate choice correlates with female reproductive success.     

Rapid evolution of male reproductive genes in the descent of man

A diverse body of morphological and genetic evidence has suggested that traits pertaining to male reproduction may have evolved much more rapidly than other types of character. Recently, DNA sequence comparisons have also shown a very high level of divergence in male reproductive proteins between closely related Drosophila species, among marine invertebrates and between mouse and rat. Here we show that rapid evolution of male reproductive genes is observable in primates and is quite notable in the lineages to human and chimpanzee. Nevertheless, rapid evolution by itself is not necessarily an indication of positive darwinian selection; relaxation of negative selection is often equally compatible with the DNA sequence data. By taking three statistical approaches, we show that positive darwinian selection is often the driving force behind this rapid evolution. These results open up opportunities to test the hypothesis that sexual selection plays some role in the molecular evolution of higher primates.     

Parents suppress reproduction and stimulate dispersal in opposite-sex juvenile white-footed mice.

Juvenile dispersal is sex-biased in many mammals and birds: one sex often disperses more often or farther than the other. Two hypotheses are generally presented for sex-biased dispersal. The first holds that juvenile dispersal reduces reproductive and/or resource competition between parents and same-sexed offspring. If so, presence of a parent on the natal home range should both promote dispersal of same-sex offspring and suppress reproduction of those that remain. The second is that juvenile dispersal reduces matings between parents and offspring, thus decreasing the likelihood of inbreeding depression. If so, presence of a parent should favour dispersal and reproductive suppression of offspring of the opposite sex. Here I present evidence that juvenile dispersal in white-footed mice, Peromyscus leucopus, is due to inbreeding avoidance. When population density was high, experimental removal of one parent delayed dispersal of opposite-sexed offspring and only the presence of the parents of opposite sex suppressed juvenile reproduction.     

Pre-social benefits of extended parental care

The evolution of helping, in which some individuals forfeit their own reproduction and help others to reproduce, is a central problem in evolutionary biology. Recently proposed insurance-based mechanisms rely on a pre-existing life history with a long period of offspring dependency relative to the short life expectancies of adult carers: a lone mother's offspring are doomed if she dies young, whereas after a helper dies, other group members can finish rearing the offspring. A critical question, however, is how this life history could evolve in ancestral non-social populations, as offspring survival would then depend on a single, short-lived carer. Here, we resolve this paradox by focusing on the extended parental care inherent in prolonged dependency. We show experimentally that in non-social wasps, extended care can significantly reduce the impact of interspecific parasites. Under extended care, offspring are less vulnerable by the time they are exposed to parasites, and costs of parasitism are reduced because mothers have the option to terminate investment in failing offspring. By experimentally simulating aspects of extended care in a species where it is lacking, we demonstrate that neither benefit requires specialized behaviour. Such benefits could therefore offset the disadvantage of prolonged dependency in non-social species, thereby facilitating the evolution of helping.     

光照诱发雪鸡恢复性机能活动

采用联合光照方法，使处于性休止期的雪鸡恢复性机能活动。雄性可产出形态正常的精子；雌性输卵管发育并能产卵，由此揭示：采用联合先照技术，有助于改变雪鸡季节性繁衍后代的习性。     

Kin selection and cooperative courtship in wild turkeys

In the few species of birds in which males form display partnerships to attract females, one male secures most or all of the copulations. This leads to the question of why subordinate males help in the absence of observable reproductive benefits. Hamilton's concept of kin selection, whereby individuals can benefit indirectly by helping a relative, was a crucial breakthrough for understanding apparently altruistic systems. However in the only direct test of kin selection in coordinated display partnerships, partners were unrelated, discounting kin selection as an explanation for the evolution of cooperation. Here I show, using genetic measures of relatedness and reproductive success, that kin selection can explain the evolution of cooperative courtship in wild turkeys. Subordinate (helper) males do not themselves reproduce, but their indirect fitness as calculated by Hamilton's rule more than offsets the cost of helping. This result confirms a textbook example of kin selection that until now has been controversial and also extends recent findings of male relatedness on avian leks by quantifying the kin-selected benefits gained by non-reproducing males.     

Paternal cyclophosphamide treatment of rats causes fetal loss and malformations without affecting male fertility

The use of cytotoxic, mutagenic and carcinogenic agents as treatment for various types of cancer may be particularly hazardous in men of reproductive age as there exists the possibility that this may lead to congenital malformations in the progeny. Such agents can affect fertility and other aspects of male reproductive function, for example, treatment with anti-cancer drugs such as cyclophosphamide has been associated with oligozoospermia, azoospermia and increased levels of serum follicle-stimulating hormone (FSH). Depending on the cumulative dose and the duration of treatment, spermatogenesis often returns but this may take years. The relevance of the effects of such chemicals on the male reproductive system to the offspring is poorly understood. We have set out to determine whether present tests of male reproductive function (that is, endocrine status, numbers of spermatozoa, fertility) can predict deleterious effects of a paternally administered agent on the offspring. Here, we report that chronic administration in rats of low doses of the widely used drug cyclophosphamide had minimal effects on the male reproductive system and fertility, but resulted in malformations and retardation of growth in the surviving fetuses and a high frequency of fetal death. Thus, adverse effects on the fetus cannot be predicted from the effects of a drug on the male reproductive system.     

Infectious parthenogenesis

Parthenogenesis-inducing Wolbachia bacteria are reproductive parasites that cause infected female wasps to produce daughters without mating. This manipulation of the host's reproduction enhances the transmission of Wolbachia to future generations because the bacteria are passed on vertically only from mothers to daughters. Males are dead ends for cytoplasmically inherited bacteria: they do not pass them on to their offspring. Vertical transmission of Wolbachia has been previously considered to be the main mode of transmission. Here we report frequent horizontal transmission from infected to uninfected wasp larvae sharing a common food source. The transferred Wolbachia are then vertically transmitted to the new host's offspring. This natural and unexpectedly frequent horizontal transfer of parthenogensis-inducing Wolbachia intraspecifically has important implications for the co-evolution of Wolbachia and their host.     

变色沙蜥繁殖的研究

作者通过研究证明变色沙蜥Ｐｈｒｙｎｏｃｅｐｈａｌｕｓ ｖｅｒｓｉｃｏｌｏｒ的繁殖活动具有明显的季节性。雄晰性成熟时最小吻－肛长为４５ｍｍ，吻－肛长大的雄晰在一年中成熟精子出现早且维持时间长。雌蜥性成熟时的最小吻－肛长为４１ｍｍ，产卵期为５月末至７月末，每只雌性成蜥每年产卵一次，吻－肛长大的雌蜥产卵多而大且比吻－肛长小的雌晰产卵早。     

Female mating bias results in conflicting sex-specific offspring fitness

Indirect-benefit models of sexual selection assert that females gain heritable offspring advantages through a mating bias for males of superior genetic quality. This has generally been tested by associating a simple morphological quality indicator (for example, bird tail length) with offspring viability. However, selection acts simultaneously on many characters, limiting the ability to detect significant associations, especially if the simple indicator is weakly correlated to male fitness. Furthermore, recent conceptual developments suggest that the benefits gained from such mating biases may be sex-specific because of sexually antagonistic genes that differentially influence male and female reproductive ability. A more suitable test of the indirect-benefit model would examine associations between an aggregate quality indicator (such as male mating success) and gender-specific adult fitness components, under the expectation that these components may trade off. Here, we show that a father's mating success in the cricket, Allonemobius socius, is positively genetically correlated with his son's mating success but negatively with his daughter's reproductive success. This provides empirical evidence that a female mating bias can result in sexually antagonistic offspring fitness.