Do tiny males grow up? Sperm competition and optimal resource allocation schedule of dwarf males of barnacles

Barnacles, marine crustaceans, have three sexual patterns: simultaneous hermaphroditism, dioecy and androdioecy. In dioecy and androdioecy, large individuals (females and hermaphrodites, respectively) are attached by dwarf males. Depending on species, some dwarf males grow up, others do not in their life time. To investigate which environmental conditions affect growth patterns of dwarf males of barnacles, we investigate the evolutionarily stable life history strategy of dwarf males using Pontryagin's maximum principle. Sperm competition among dwarf males and that among dwarf males and large hermaphrodites is taken into account. Dwarf males grow up in food-rich environments, while they do not grow at all in food-poor environments. ESS of the resource allocation schedule between reproduction and growth follows an "intermediate growth strategy" (simultaneous growth and reproduction) for dioecious species, in which sperm competition is not severe. On the other hand, it approaches "bang-bang control" (switching from allocating all resources toward growth then to reproduction), as sperm competition against surrounding large hermaphrodites becomes severe in androdioecious species.     

An evolutionarily stable joining policy for group foragers

For foragers that exploit patchily distributed resources thatare challenging to locate, detecting discoveries made by otherswith a view to joining them and sharing the patch may oftenbe an attractive tactic, and such behavior has been observedacross many taxa. If, as will commonly be true, the time takento join another individual on a patch increases with the distanceto that patch, then we would expect foragers to be selectivein accepting joining opportunities: preferentially joining nearbydiscoveries. If competition occurs on patches, then the profitabilityof joining (and of not joining) will be influenced by the strategiesadopted by others. Here we present a series of models designedto illuminate the evolutionarily stable joining strategy. Weconfirm rigorously the previous suggestion that there shouldbe a critical joining distance, with all joining opportunitieswithin that distance being accepted and all others being declined.Further, we predict that this distance should be unaffectedby the total availability of food in the environment, but shouldincrease with decreasing density of other foragers, increasingspeed of movement towards joining opportunities, increased difficultyin finding undiscovered food patches, and decreasing speed withwhich discovered patches can be harvested. We are further ableto make predictions as to how fully discovered patches shouldbe exploited before being abandoned as unprofitable, with discoveredpatches being more heavily exploited when patches are hard tofind: patches can be searched for remaining food more quickly,forager density is low, and foragers are relatively slow intraveling to discovered patches.     

Patterns of evolutionarily stable strategies: the maximal pattern conjecture revisited

A conflict is defined by a set of pure strategies [1,..., n] and a payoff matrix, and may have many evolutionarily stable strategies (ESSs). A collection of subsets of the set of pure strategies is called a pattern. If there is an n x n matrix which has ESSs whose supports match those of the pattern, then that pattern is said to be attainable. Much of the work on patterns of ESSs relied upon an unproved conjecture. Subject to some relaxation of the definition of attainability, this conjecture is proved.     

Dichogamy, Sex Allocation, and Mating System of Campanula microdonta and C. punctata

Dichogamy and sex allocation in several populations of Campanula microdonta and C. punctata were investigated with regard to their mating systems. Duration of the staminate phase differed among the populations: staminate phase was longer in self-compatible (SC) and largely outcrossing populations than in self-incompatible (SI) and outcrossing populations or in SC and largely inbreeding populations. Duration of the pistillate phase among the populations was less variable than duration of the staminate phase. Male reproductive effort decreased with increase of the estimated selfing rates. Male allocation (weight ratio of androecium to gynoecium or to total flower) may be used as an indicator of the breeding system. Within each population, small flowers allocate proportionately more resources to the androecium than to the gynoecium. Among populations, SC outcrossing populations tend to produce large ovaries, and SC inbreeding populations tend to produce small ovaries.     

Mating behavior of cockerels: Effects of social status,group size and group density

Mating behavior was studied to reexamine the correlation between social status of cockerels and mating frequency. A moderate correlation of 0.30 (P < 0.025) was found between males-dominated and completed-matings-per-hour. This relationship was not as large as expected from the findings by Guhl and Warren (1946), but was closer to the results of Craig et al. (1977). Completed matings, aggressive acts and interferences increased relative to dominance categories in four combinations of group size and density. Males with more area had more (P < 0.05) courting, completed matings and aggressive acts, and smaller flocks showed more behavioral activity. Mating activities peaked between the second and third weeks after flock formation. Mating proficiency increased linearly over time and was adversely affected by interference until the fourth week.     

Adaptive evolution of sexual systems in pedunculate barnacles

How and why diverse sexual systems evolve are fascinating evolutionary questions, but few empirical studies have dealt with these questions in animals. Pedunculate (gooseneck) barnacles show such diversity, including simultaneous hermaphroditism, coexistence of dwarf males and hermaphrodites (androdioecy), and coexistence of dwarf males and females (dioecy). Here, we report the first phylogenetically controlled test of the hypothesis that the ultimate cause of the diverse sexual systems and presence of dwarf males in this group is limited mating opportunities for non-dwarf individuals, owing to mating in small groups. Within the pedunculate barnacle phylogeny, dwarf males and females have evolved repeatedly. Females are more likely to evolve in androdioecious than hermaphroditic populations, suggesting that evolution of dwarf males has preceded that of females in pedunculates. Both dwarf males and females are associated with a higher proportion of solitary individuals in the population, corroborating the hypothesis that limited mating opportunities have favoured evolution of these diverse sexual systems, which have puzzled biologists since Darwin.     

Resource allocation to reproduction and soma in Drosophila: a stable isotope analysis of carbon from dietary sugar

Metabolic resources in adults of holometabolous insects may derive either from larval or adult feeding. In Drosophila melanogaster, reproduction and lifespan are differently affected by larval vs. adult resource availability, and it is unknown how larval vs. adult acquired nutrients are differentially allocated to somatic and reproductive function. Here we describe the allocation of carbon derived from dietary sugar in aging female D. melanogaster. Larval and adult flies were fed diets contrasting in sucrose (13)C/(12)C, from which we determined the extent to which carbon acquired at each stage contributed to adult somatic tissue and to egg manufacture. Dietary sugar is very important in egg provisioning; at every age, roughly one half of the carbon in eggs was derived from sugar, which turned over from predominantly larval to entirely adult dietary sources. Sucrose provided approximately 40% of total somatic carbon, of which adult dietary sucrose came to supply approximately 75%. Unlike in eggs, however, adult acquired sucrose did not entirely replace the somatic carbon from larvally acquired sucrose. Because carbon from larval sucrose appears to be fairly "replaceable", larval sucrose cannot be a limiting substrate in resource allocation between reproduction and lifespan.     

不同刈牧强度对冷蒿生长与资源分配的影响

利用野外实验与盆栽实验,对不同刈牧强度下冷蒿生长与资源分配影响的研究结果表明,按比例刈割冷篙的再生生长大于留茬高度刈割,在生长季前期,不刈割冷蒿净生长高于刈割处理,而进入生长季中后期(8月中旬以后),轻度刈割净生长高于不刈割处理,冷篙种群生物量分配的总体格局是根>叶>茎,刈牧明显影响冷蒿生物量分配格局,尤其是叶和花的分配,3／4刈割或留茬4cm刈割叶生物量分配显著高于其它各处理,而花的生物量及其分配显著低于其它处理,根、茎生物量分配各处理间差异不显著．冷蒿有性生殖分配随刈牧强度的增加而降低,繁殖方式发生了改变,优先将光合产物分配给再生茎以及繁殖方式转向营养繁殖,通过克隆生长维持和扩大种群是冷蒿对强度放牧的生态适应对策。     

Effects of social group size on information transfer and task allocation

Summary Social animals exchange information during social interaction. The rate of interaction and, hence, the rate of information exchange, typically changes with density and density may be affected by the size of the social group. We investigate models in which each individual may be engaged in one of several tasks. For example, the different tasks could represent alternative foraging locations exploited by an ant colony. An individual's decision about which task to pursue depends both on environmental stimuli and on interactions among individuals. We examine how group size affects the allocation of individuals among the various tasks. Analysis of the models shows the following. (1) Simple interactions among individuals with limited ability to process information can lead to group behaviour that closely approximates the predictions of evolutionary optimality models, (2) Because per capita rates of social interaction may increase with group size, larger groups may be more efficient than smaller ones at tracking a changing environment, (3) Group behaviour is determined both by each individual's interaction with environmental stimuli and by social exchange of information. To keep these processes in balance across a range of group sizes, organisms are predicted to regulate per capita rates of social interaction and (4) Stochastic models show, at least in some cases, that the results described here occur even in small groups of approximately ten individuals.     

Mating system and brain size in bats

The contribution of sexual selection to brain evolution has been little investigated. Through comparative analyses of bats, we show that multiple mating by males, in the absence of multiple mating by females, has no evolutionary impact on relative brain dimension. In contrast, bat species with promiscuous females have relatively smaller brains than do species with females exhibiting mate fidelity. This pattern may be a consequence of the demonstrated negative evolutionary relationship between investment in testes and investment in brains, both metabolically expensive tissues. These results have implications for understanding the correlated evolution of brains, behaviour and extravagant sexually selected traits.     

Superinfections can induce evolutionarily stable coexistence of pathogens

Parasites reproduce and are subject to natural selection at several different, but intertwined, levels. In the recent paper, Gilchrist and Coombs (Theor. Popul. Biol. 69:145–153, 2006) relate the between-host transmission in the context of an SI model to the dynamics within a host. They demonstrate that within-host selection may lead to an outcome that differs from the outcome of selection at the host population level. In this paper we combine the two levels of reproduction by considering the possibility of superinfection and study the evolution of the pathogen’s within-host reproduction rate p. We introduce a superinfection function φ = φ(p,q), giving the probability with which pathogens with trait q, upon transmission to a host that is already infected by pathogens with trait p, “take over” the host. We consider three cases according to whether the function q → φ(p,q) (i) has a discontinuity, (ii) is continuous, but not differentiable, or (iii) is differentiable in q = p. We find that in case (i) the within-host selection dominates in the sense that the outcome of evolution at the host population level coincides with the outcome of evolution in a single infected host. In case (iii), it is the transmission to susceptible hosts that dominates the evolution to the extent that the singular strategies are the same as when the possibility of superinfections is ignored. In the biologically most relevant case (ii), both forms of reproduction contribute to the value of a singular trait. We show that when φ is derived from a branching process variant of the submodel for the within-host interaction of pathogens and target cells, the superinfection functions fall under case (ii). We furthermore demonstrate that the superinfection model allows for steady coexistence of pathogen traits at the host population level, both on the ecological, as well as on the evolutionary time scale.      

An evolutionarily stable strategy for aggressiveness in feeding groups

Animals searching for food in groups display highly variabledegrees of aggressiveness. In this paper I present an individual-basedgame theoretical model of how gregarious animals should adjusttheir level of aggressiveness to their environmental conditions.In accordance with behavioral observations, the predicted levelof aggressiveness increases progressively with decreasing foodavailability and increasing animal density. The proposed modelalso predicts a positive influence of food energy value andhandling time on the level of aggressiveness within the group.In addition, the model provides information about the influenceof aggressive behavior on individual foraging success, interference,and population dynamics. Adaptive behavioral rules for aggressivenessin consumers are predicted to respond to both competitors and food density in a way that contributes to stabilization of thedynamics of population systems.     

Influence of social mixing and group size on skin lesions and mounting in organic entire male pigs

Alternatives to surgical castration are needed, due to stress and pain caused by castration of male pigs. One alternative is production of entire male pigs. However, changed behaviour of entire males compared with castrated males might adversely affect the welfare of entire males and changes in management procedures and production system might be needed. Elements from the organic pig production system might be beneficial in this aspect. The aim of this article is to investigate the effect of grouping strategy including social mixing and group size on levels of mounting behaviour and skin lesions, hypothesising that procedures that disrupt the social stability (e.g. regrouping) will have a larger negative effect in small groups compared with large groups. Approximately 1600 organic entire male pigs of the breed (Landrace×Yorkshire)×Duroc were reared in parallel in five organic herds, distributed across four batches in a 2×2 factorial design in order to test the influence of social mixing (presence or absence of social mixing at relocation) and group size (15 and 30 animals). Animals were able to socialise with piglets from other litters during the lactation period, and were all mixed across litters at weaning. A second mixing occurred at insertion to fattening pens for pigs being regrouped. Counting of skin lesions (1348 or 1124 pigs) and registration of mounting behaviour (1434 or 1258 pigs) were done on two occasions during the experimental period. No interactive effects were found between social mixing and group size on either skin lesions or mounting frequency. Herd differences were found for both mounting frequency and number of skin lesions. No association between skin lesions and mounting were revealed. Social mixing and group size were shown as interacting effects with herds on mounting frequency (P<0.0001), but with no consistent pattern across all herds. In addition, no effect of social mixing was found on mean number of skin lesions, but more lesions were observed in large groups (P<0.036). This could indicate that keeping entire male pigs in groups of 30 animals as compared with smaller groups of 15 may marginally decrease the welfare of these animals.     

Mating tactics and male-male courtship in the lek-breeding cichlid Oreochromis mossambicus

Data are presented on the breeding behaviour of Oreochromis mossambicus under captive conditions. Males tended to synchronize their occupation of territories and breeding activities. Different male mating tactics were observed, namely establishing a breeding territory, acting as a floater, or behaving as a sneaker. The majority of spawnings observed involved dominant males and were subjected to interference from other males. Males were found to court other males that frequently responded to these attempts by adopting a female-like behaviour. Results are discussed in terms of a probable time constraint in territoriality, which promotes male-male competition and a low level of sex discrimination by territorial fish.     

Asynchronous pupation of univoltine insects as evolutionarily stable phenology

Evolutionarily stable seasonal timing of larval feeding stages is studied theoretically for univoltine insects. In the evolutionarily stable (or ESS) population, each individual maximizes its own lifetime reproductive success by choosing the hatching and pupation dates, given the resource availability curve with a peak in the middle of a year, a higher daily mortality in the feeding stages, and the growth rate decreasing with the larval biomass in the population. If growth rate is proportional to the body size, the population at the ESS is composed of a mixture of phenotypes differing in hatching and pupation, but pupation interval over which some popation occur every day is much longer than hatching interval. If growth rate increases with the body size at a speed slower than linearly, large sized larvae should pupate earlier than small ones.     

Genital cutting and western discourses on sexuality

This article explores dominant discourses surrounding male and female genital cutting. Over a similar period of time, these genital operations have separately been subjected to scrutiny and criticism. However, although critiques of female circumcision have been widely taken up, general public opinion toward male circumcision remains indifferent. This difference cannot merely be explained by the natural attributes and effects of these practices. Rather, attitudes toward genital cutting reflect historically and culturally specific understandings of the human body. In particular, I suggest that certain problematic understandings of male and female sexuality are deeply implicated in the dominant Western discourses on genital surgery.     

Strong stability and evolutionarily stable strategies with two types of players

Static ESS conditions are developed for the frequency evolution of a two-species haploid system by analyzing the stability of the corresponding dynamics for two pairs of strategies. A dynamic strong stability concept is introduced and shown to be equivalent to the ESS conditions in all cases where a regularity assumption is satisfied.