Certainty of paternity covaries with paternal care in birds

Summary Male investment in parental care has been hypothesized to be affected or not to be affected by their certainty of paternity, depending on the particular assumptions of theoretical models. We used data on paternal care and extra-pair paternity from 52 bird species to determine whether male parental care was related to certainity of paternity. Paternal care was measured as the relative male contribution to nest building, courtship feeding, incubation, and feeding of nestlings, respectively. Males of avian taxa did not provide less parental care during nest building, courtship feeding and incubation if the frequency of extra-pair paternity was high. However, male participation in feeding of offspring was significantly negatively related to the frequency of extra-pair paternity. This was also the case when the effects of potentially confounding variables such as developmental mode of offspring (which may result in males being freed from parental duties), extent of polygyny (which may result in less paternal care), and the frequency of multiple clutches during one breeding season (which may increase the probability of finding fertile females during the nestling period) were controlled statistically. These results suggest that the extent of paternal care has been affected by certainty of paternity, and that sex roles during the energetically most expensive parts of reproduction have been shaped by sperm competition.     

Sperm competition or sperm selection: no evidence for female influence over paternity in yellow dung flies Scatophaga stercoraria

Recent studies of non-random paternity have suggested that sperm selection by females may influence male fertilization success. Here we argue that the problems originally encountered in partitioning variation in non-random mating between male competition and female choice are even more pertinent to interpreting patterns of non-random paternity because of intense sperm competition between males. We describe an experiment with the yellow dung fly, Scatophaga stercoraria, designed to partition variance in the proportion of offspring sired by the second male, P ₂, between males and females, and to control for sperm competition. Large males were shown to have a higher P ₂ than small males but P ₂ was independent of the size of the female’s first mate. This result might suggest an absolute female preference for large males via sperm selection. However, large males have a higher constant rate of sperm transfer and displacement. After controlling for this effect of sperm competition, large males did not achieve higher paternity than small males. We argue that a knowledge of the mechanism of sperm competition is essential so that male effects can be controlled before conclusions are made regarding the influence of sperm selection by females in generating non-random paternity. Received: 4 April 1995 / Accepted after revision: 17 October 1995     

Female choice on male quantitative traits in lizards — why is it so rare?

Female choice on the basis of male traits has been described in an array of taxa but has rarely been demonstrated in reptiles. In the sand lizard (Lacerta agilis), and possibly in other non-territorial reptiles, a male's contribution to a female's fitness is restricted to his genes. In order to choose males of high genetic quality, females have to trade the fitness gain against the costs of active choice. In a Swedish population of sand lizards, long-lived males sired offspring with higher embryonic survival compared to offspring sired by short-lived males. In spite of this female sand lizards did not mate selectively with older and/or larger males. There appeared to be mo reliable cues to male longevity; age-specific male body size was highly variable. Furthermore, estimates of male nuptial coloration did not covary with ectoparasite load and, hence, females cannot use male coloration as a cue to heritable resistance to pathogenic parasite effects. When cues to male genetic quality are poor, or inaccurate, and males make no parental investment, we predict that female choice will be rare. Sand lizard females mating with many partners lay clutches with higher hatching success. Thus, females may obtain good genes for their young by multiple mating, thereby avoiding costs associated with mate choice.     

Why do male Callosobruchus maculatus beetles inseminate so many sperm?

Male Callosobruchus maculatus F. (Coleoptera: Bruchidae) inseminate more sperm than females can effectively store in their spermathecae. This study examines the adaptive significance of excess sperm transfer by measuring components of male and female reproductive success in response to manipulating the number of sperm inseminated. The number of sperm transferred during copulation was reduced from 56,000 ±4,462 to 8,700±1,194 by sequentially mating males to virgin females. Reducing the number of sperm inseminated by the first male to mate had no effect on the extent of sperm precedence, but reducing the number of sperm inseminated by the second male resulted in a significant reduction in the extent of sperm precedence. When large numbers of sperm are inseminated the remating refractory period of females is increased. These results indicate that males transferring large numbers of sperm during copulation have a two-fold advantage at fertilization; they are more effective at preempting previously stored sperm and they are likely to father more offspring by delaying the time of female remating. The transfer of excess sperm does not appear to serve as nonpromiscuous male mating effort; the number of eggs laid, their fertility and the subsequent survival of zygotes were unaffected by manipulating the number of sperm inseminated. The underlying mechanisms of sperm precedence were also examined. Simple models of sperm displacement failed to accurately predict the patterns of sperm precedence observed in this species. However, the results do not provide conclusive evidence against the models but rather serve to highlight our limited understanding of the movement of sperm within the female's reproductive tract.     

Increased sperm numbers in the vas deferens of meadow voles, <Emphasis Type="Italic">Microtus pennsylvanicus,</Emphasis> in response to odors of conspecific males

Sperm competition occurs when the sperm of two or more males compete to fertilize the egg/s of a particular female. Males of some species respond to a high risk of sperm competition by increasing the number of sperm in their ejaculates. Males may accomplish such a response by increasing the intensity or duration of contraction of the cauda epididymidis and vas deferens. During emission (first phase of the ejaculatory process), the vas deferens receives sperm from the cauda epididymidis and propels the sperm to the urethra. In this paper, we tested the hypothesis that males exposed to a high risk of sperm competition mobilize larger numbers of sperm from the cauda epididymidis to the vas deferens before initiation of copulatory behavior. This accumulation of sperm in the vas deferens would result in a larger number of sperm in the ejaculate. To test this hypothesis, we exposed male meadow voles, Microtus pennsylvanicus, to either low or high risks of sperm competition using soiled bedding of conspecific individuals. At three different times after this exposure (15, 30, or 60 min), we removed both vasa deferentia and counted the sperm within them. We found a significant increase in sperm numbers in the vas deferens of males after 30 min of being exposed to a high risk of sperm competition. The lower sperm numbers after 15 and 60 min of exposure suggest that the observed response is relatively slow and that sperm mobilized to the vasa deferentia may return to the cauda epididymides if ejaculation does not occur some time after the observed response. Our results indicate that the physiological response that may result in high sperm numbers in the ejaculate in relation to high risk of sperm competition can occur before initiation of copulatory behavior.     

Effects of frequent mating on sex allocation in the simultaneously hermaphroditic great pond snail (Lymnaea stagnalis)

Simultaneous hermaphrodites have the opportunity to control the allocation of resources to the male and female function depending on the circumstances. Such flexibility also provides the possibility to influence sex allocation in the mating partner. To investigate this idea, we measured egg production (female investment) and sperm production as well as prostate gland size (both are part of male investment) under different mating regimes in the great pond snail Lymnaea stagnalis. We find no evidence for the prediction from sex allocation theory that sperm production increases with mating frequency. However, we do find that animals with more mating opporunities develop smaller prostate glands, in which seminal fluid is produced. Moreover, repeated mating increases egg production, thus shifting allocation towards the female function, and probably decreases growth. So, our data hint at a three-way trade-off between part of the male function (prostate gland), female function, and growth. Interestingly, sex allocation seems to be shifted in the opposite direction from the one predicted by theory. We discuss how this feminization is suggestive of a direct manipulation by the sperm donor, probably to stimulate immediate sperm use.     

Patterns of sperm use in the scorpionfly Panorpa germanica L. (Mecoptera: Panorpidae)

Sperm competition can be a powerful selective force in the evolution of reproductive strategies and mating systems. In studies on sperm competition, patterns of sperm use are typically reported as the mean species value of P ₂, determined as the proportion of offspring sired by the second male to copulate with a doubly mated female. However, the within-species variance in P ₂ has mostly been ignored, although taking this variance into account may be crucial for understanding the underlying mechanisms of sperm competition. Paternity analysis among the offspring of doubly mated females of Panorpa germanica (Mecoptera, Panorpidae) revealed a relationship between relative copulation durations of both males and the proportion of offspring each male will sire. This correlation between proportional copulation durations and paternity suggests mixing of sperm from different males inside the female’s spermatheca. Yet, sperm mixing appears to be incomplete, as paternity was overall slightly shifted towards the second male on average fathering a higher proportion of the offspring than its relative copulation duration would predict in case of complete sperm mixing. For individual males, however, the outcome of sperm competition is rather unpredictable as the intraspecific variance in P ₂ was found to be very high, irrespective of copulation durations. Possible causes of the observed variance in P ₂ and the partial last male sperm precedence are discussed.     

Begging and provisioning in broods of asynchronously-hatched yellow-headed blackbird nestlings

Studies of begging have found a positive relationship between begging level and provisioning level. Studies of unequal nestlings, however, have found that small nestlings generally beg more but are fed less than their larger siblings. We manipulated the begging levels of yellow-headed blackbird (Xanthocephalus xanthocephalus) chicks to investigate how begging benefits individuals in broods of unequal siblings. Food-deprived chicks begged more and were fed more; satiated chicks begged less and were fed less. When we deprived each chick of a brood in turn, large and small chicks generally increased begging and received more provisioning. Small chicks, however, rarely received more food than their larger siblings even when they behged relatively more. Parent yellow-headed blackbirds increase provisioning to hungry begging chicks, but also allocate food based on relative offspring size.     

低磷胁迫下不同水稻品种根系吸收能力差异的生理与遗传本质

在低磷（1mg L-1)和适磷水平（10mg L^-1)下，分别以混合培养方式研究了粳稻京系17（Oryza sativa ssp.japonica)(JX17)和粘稻窄叶青8号（Oryza sativa ssp.indica)(ZYQ8)对磷营养的反应。结果表明，低磷胁迫下，JX17较ZYQ8吸收更多磷素，干物重也显著增加。相反，ZYQ8则受到更强的磷胁迫，吸磷量和干物重降低，主要是因为JX17具有较高的吸磷速度，单位根长吸磷量大，即根系高亲和力磷酸盐转运蛋白表达强，最大限度地吸收可能利用的磷素，从而改善植株营养状况，同时又有较多的磷被分配到根系，使根第，根表面积和根干重相应提高，进一步增加了其在低磷胁迫条件下的竞争磷营养的优势。图3表2参17     

Between-group encounters among bonnet macaques (<Emphasis Type="Italic">Macaca radiata</Emphasis>)

Socioecological theory suggests that between-group competition is an important factor affecting the nature of primate social relationships. Between-group encounters in macaques may involve female resource defense, male mate defense, and male resource defense. We observed between-group encounters in two groups (a forest group and a temple group) of bonnet macaques (Macaca radiata). We observed 102 encounters in 875 h of observation of the forest group (1.40 per 12-h day) and 58 encounters in 907 h of observation of the temple group (0.77 per 12-h day). Aggressive interactions between groups occurred in 32.4% and 29.3% of encounters in the forest and temple groups, respectively. Overall, we found little support for the female resource defense hypothesis. Females in both groups rarely participated aggressively in between-group encounters. We found support for the male mate defense hypothesis. For example, males of the forest group were more aggressive during encounters in the mating season than in the non-mating season. Males were also aggressive to females from their own group immediately following encounters. We also found partial support for the male resource defense hypothesis. Encounters in the forest group occurred in a feeding context more often than expected based on time budgets. Also, males in the temple group were more often aggressive in food-related encounters than in other encounters. The findings of this study suggest that socioecological models of primate social relationships need to distinguish male and female strategies during between-group encounters and integrate the resulting functional outcomes.Communicated by T. Czeschlik