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1.
The number of mating partners an individual has within a population is a crucial parameter in sex allocation theory for simultaneous hermaphrodites because it is predicted to be one of the main parameters influencing sex allocation. However, little is known about the factors that determine the number of mates in simultaneous hermaphrodites. Furthermore, in order to understand the benefits obtained by resource allocation into the male function it is important to identify the factors that predict sperm‐transfer success, i.e. the number of sperm a donor manages to store in a mate. In this study we experimentally tested how social group size (i.e. the number of all potential mates within a population) and density affect the number of mates and sperm‐transfer success in the outcrossing hermaphroditic flatworm Macrostomum lignano. In addition, we assessed whether these parameters covary with morphological traits, such as body size, testis size and genital morphology. For this we used a method, which allows tracking sperm of a labelled donor in an unlabelled mate. We found considerable variation in the number of mates and sperm‐transfer success between individuals. The number of mates increased with social group size, and was higher in worms with larger testes, but there was no effect of density. Similarly, sperm‐transfer success was affected by social group size and testis size, but in addition this parameter was influenced by genital morphology. Our study demonstrates for the first time that the social context and the morphology of sperm donors are important predictors of the number of mates and sperm‐transfer success in a simultaneous hermaphrodite. Based on these findings, we hypothesize that sex allocation influences the mating behaviour and outcome of sperm competition. 相似文献
2.
Evolutionary theory predicts an influence of mating group size on sex allocation in simultaneous hermaphrodites. We experimentally manipulated the social situation during reproduction in a simultaneous hermaphrodite parasite, the tapeworm Schistocephalus solidus, by placing worms as singles, pairs or triplets into an in vitro system that replaces the final host. We then determined the reproductive allocation patterns after 24 h (i.e. before the start of egg release) and after 72 h (i.e. around the peak of egg release rate) using stereology. After 24 h, sex allocation strongly depended on worm volume (which is determined in the second intermediate host), but was not significantly affected by the social situation experienced during reproduction. After 72 h, worms in groups had less vesicular sperm (i.e. sperm to be used in future inseminations) than singles. They also stored significantly more received sperm in their seminal receptacles than singles, suggesting that more sperm had been transferred in groups. Moreover, worms in triplets stored significantly more received sperm than worms in pairs, suggesting that they either mated more often and/or transferred more sperm per mating. This suggests a behavioural response to the increased risk of sperm competition in triplets. We further discuss the relative importance of sex allocation decisions at different life‐history stages. 相似文献
3.
Most models of sex allocation distinguish between sequential and simultaneous hermaphrodites, although an intermediate sexual pattern, size‐dependent sex allocation, is widespread in plants. Here we investigated sex allocation in a simultaneous hermaphrodite animal, the tapeworm Schistocephalus solidus, in which adult size is highly variable. Sex allocation was determined using stereological techniques, which allow measuring somatic and reproductive tissues in a common currency, namely volume. We investigated the relationships between individual volume and allocation to different reproductive tissues using an allometric model. One measure of female allocation, yolk gland volume, increased more than proportionally with individual volume. This is in contrast to the measure of male allocation, testis volume, which showed a strong tendency to increase less than proportionally with individual volume. Together these patterns led to sex allocation being strongly related to individual volume, with large individuals being more biased towards female allocation. We discuss these findings in the light of current ideas about size‐dependent sex allocation in, primarily, plants and try to extend them to simultaneous hermaphrodite animals. 相似文献
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5.
Pemberton AJ Sommerfeldt AD Wood CA Flint HC Noble LR Clarke KR Bishop JD 《Journal of evolutionary biology》2004,17(3):506-518
The importance of sexual compatibility between mates has only recently been realized in zoological research into sexual selection, yet its study has been central to botanical research for many decades. The reproductive characteristics of remote mating, an absence of precopulatory mate screening, internal fertilization and embryonic brooding are shared between passively pollinated plants and a phylogenetically diverse group of sessile aquatic invertebrates. Here, we further characterize the sexual compatibility system of one such invertebrate, the colonial ascidian Diplosoma listerianum. All 66 reciprocal pairings of 12 genetic individuals were carried out. Fecundities of crosses varied widely and suggested a continuous scale of sexual compatibility. Of the 11 animals from the same population c. 40% of crosses were completely incompatible with a further c. 20% having obvious partial compatibility (reduced fecundity). We are unaware of other studies documenting such high levels of sexual incompatibility in unrelated individuals. RAPD fingerprinting was used to estimate relatedness among the 12 individuals after a known pedigree was successfully reconstructed to validate the technique. In contrast to previous results, no correlation between genetic similarity and sexual compatibility was detected. The blocking of many genotypes of sperm is expected to severely modify realized paternity away from 'fair raffle' expectations and probably reduce levels of intra-brood genetic diversity in this obligatorily promiscuous mating system. One adaptive benefit may be to reduce the bombardment of the female reproductive system by outcrossed sperm with conflicting evolutionary interests, so as to maintain female control of somatic : gametic investment. 相似文献
6.
Baeza JA 《Evolution; international journal of organic evolution》2007,61(10):2360-2373
Two fundamental questions dealing with simultaneous hermaphrodites are how resources are optimally allocated to the male and female function and what conditions determine shifts in optimal sex allocation with age or size. In this study, I explored multiple factors that theoretically affect fitness gain curves (that depict the relationship between sex-specific investment and fitness gains) to predict and test the overall and size-dependent sex allocation in a simultaneously hermaphroditic brooding shrimp with an early male phase. In Lysmata wurdemanni, sperm competition is absent as hermaphrodites reproducing in the female role invariably mated only once with a single other shrimp. Shrimps acting as females preferred small over large shrimps as male mating partners, male mating ability was greater for small compared to large hermaphrodites, and adolescent males were predominant in the population during the breeding season. In addition, brooding constraints were not severe and varied linearly with body size whereas the ability to acquire resources increased markedly with body size. Using sex allocation theory as a framework, the findings above permitted to infer the shape of the male and female fitness gain curves for the hermaphrodites. The absence of sperm competition and the almost unconstrained brooding capacity imply that both curves saturate, however the male curve levels off much more quickly than the female curve with increasing level of investment. In turn, the predominance of adolescent males in the population implies that the absolute gain of the female curve is greater than that of the male curve. Last, the size-dependent female preference and male mating ability of hermaphrodites determines that the absolute gain of the male curve is greater for small than for large hermaphrodites. Taking into consideration the inferred shape of the fitness gain curves, two predictions with respect to the optimal sex allocation were formulated. First, overall sex allocation should be female biased; it permits hermaphrodites to profit from the female function that provides a greater fitness return than the male function. Second, sex allocation should be size-dependent with smaller hermaphrodites allocating more than proportionally resources to male reproduction than larger ones. This size-dependent sex allocation permits hermaphrodites to profit from male mating opportunities that are the greatest at small body sizes. Size-dependent sex allocation is also expected because the male fitness gain curve decelerates more quickly than the female gain curve and experiments indicated that resources are greater for large than small hermaphrodites. These two predictions were tested when determining the sex allocation of hermaphrodites by dissecting their gonad and quantifying ovaries versus testes mass. Supporting the predictions above, hermaphrodites allocated, on average, 118 times more to the female than to the male gonad and the proportion of resources devoted to male function was higher in small than in large hermaphrodites. A trade-off between male and female allocation is assumed by theory but no negative correlation between male and female reproductive investment was observed. In L. wurdemanni, the relationship between sex-specific investment and fitness changes during ontogeny in a way that is consistent with an adjustment of sex allocation to improve size-specific reproductive success. 相似文献
7.
Sex allocation theory predicts that mating frequency and long‐term sperm storage affect the relative allocation to male and female function in simultaneous hermaphrodites. We examined the effect of mating frequency on male and female reproductive output (number of sperm delivered and eggs deposited) and on the resources allocated to the male and female function (dry mass, nitrogen and carbon contents of spermatophores and eggs) in individuals of the simultaneous hermaphrodite land snail Arianta arbustorum. Similar numbers of sperm were delivered in successive copulations. Consequently, the total number of sperm transferred increased with increasing number of copulations. In contrast, the total number of eggs produced was not influenced by the number of copulations. Energy allocation to gamete production expressed as dry mass, nitrogen or carbon content was highly female‐biased (>95% in all estimates). With increasing number of copulations the relative nitrogen allocation to the male function increased from 1.7% (one copulation) to 4.7% (three copulations), but the overall reproductive allocation remained highly female‐biased. At the individual level, we did not find any trade‐off between male and female reproductive function. In contrast, there was a significant positive correlation between the resources allocated to the male and female function. Snails that delivered many sperm also produced a large number of eggs. This finding contradicts current theory of sex allocation in simultaneous hermaphrodites. 相似文献
8.
Physiological trade-offs between life-history traits can constrain natural selection and maintain genetic variation in the face of selection, thereby shaping evolutionary trajectories. This study examines physiological trade-offs in simultaneously hermaphroditic banana slugs, Ariolimax dolichophallus. These slugs have high heritable variation in body size, which strongly predicts the number of clutches laid, hatching success and progeny growth rate. These fitness components were associated, but only when examined in correlation with body size. Body size mediated these apparent trade-offs in a continuum where small animals produced rapidly growing progeny, intermediate-sized animals laid many clutches and large animals had high hatching success. This study uses a novel statistical method in which the components of fitness are analysed in a mancova and related to a common covariate, body size, which has high heritability. The mancova reveals physiological trade-offs among the components of fitness that were previously masked by high variation in body size. 相似文献
9.
《Ethology, Ecology and Evolution》2012,24(2):187-195
The relationship between body size and hermaphroditic mating strategies was studied by following a population of the sea slug, Aplysia vaccaria, for one year. Fourteen individuals were tagged with internal microchips, and all mating pairs were weighed. Tagged individuals mated repeatedly both as sperm recipients and sperm donors. There was a trend toward size-assortative mating and size-assortative spatial clustering, and animals acting as females were larger than the average mass for the population. These findings support prior studies showing an association between larger relative size and sperm recipient roles in hermaphroditic gastropods, and highlight the need for critical experiments to determine the mechanisms of role choice and mate choice. 相似文献
10.
N. Vellnow L. Marie‐Orleach K. S. Zadesenets L. Schärer 《Journal of evolutionary biology》2018,31(2):180-196
Hermaphroditic animals face the fundamental evolutionary optimization problem of allocating their resources to their male vs. female reproductive function (e.g. testes and sperm vs. ovaries and eggs), and this optimal sex allocation can be affected by both pre‐ and post‐copulatory sexual selection. For example, local sperm competition (LSC) – the competition between related sperm for the fertilization of a partner's ova – occurs in small mating groups and can favour a female‐biased sex allocation, because, under LSC, investment into sperm production is predicted to show diminishing fitness returns. Here, we test whether higher testis investment increases an individual's paternity success under sperm competition, and whether the strength of this effect diminishes when LSC is stronger, as predicted by sex allocation theory. We created two subsets of individuals of the simultaneously hermaphroditic flatworm Macrostomum lignano – by sampling worms from either the highest or lowest quartile of the testis investment distribution – and estimated their paternity success in group sizes of either three (strong LSC) or eight individuals (weak LSC). Specifically, using transgenic focal individuals expressing a dominant green‐fluorescent protein marker, we showed that worms with high testis investment sired 22% more offspring relative to those with low investment, corroborating previous findings in M. lignano and other species. However, the strength of this effect was not significantly modulated by the experienced group size, contrasting theoretical expectations of more strongly diminishing fitness returns under strong LSC. We discuss the possible implications for the evolutionary maintenance of hermaphroditism in M. lignano. 相似文献
11.
Masami M. Tamechika Kohei Matsuno Satoshi Wada Yoichi Yusa 《Ecology and evolution》2020,10(5):2492-2498
Sex allocation theory predicts that the optimal sexual resource allocation of simultaneous hermaphrodites is affected by mating group size (MGS). Although the original concept assumes that the MGS does not differ between male and female functions, the MGS in the male function (MGSm; i.e., the number of sperm recipients the focal individual can deliver its sperm to plus one) and that in the female function (MGSf; the number of sperm donors plus one) do not always coincide and may differently affect the optimal sex allocation. Moreover, reproductive costs can be split into “variable” (e.g., sperm and eggs) and “fixed” (e.g., genitalia) costs, but these have been seldom distinguished in empirical studies. We examined the effects of MGSm and MGSf on the fixed and variable reproductive investments in the sessilian barnacle Balanus rostratus. The results showed that MGSm had a positive effect on sex allocation, whereas MGSf had a nearly significant negative effect. Moreover, the “fixed” cost varied with body size and both aspects of MGS. We argue that the two aspects of MGS should be distinguished for organisms with unilateral mating. 相似文献
12.
S. K. Emms D. A. Stratton A. A. Snow 《Evolution; international journal of organic evolution》1997,51(5):1481-1489
We studied the relationship between inflorescence size and male fitness in the andromonoecious lily Zigadenus paniculatus, using experimentally manipulated inflorescences to eliminate possible correlations between flower number, resource availability, and other floral traits. Allozyme markers were used to determine the siring success of large versus small plants in 14 arrays of plants, each array containing five large and five small plants. The inflorescence size of small plants was held constant both within and among arrays; the size of large plants was held constant within an array but was varied among arrays. Large plants sired more than half the seeds in 12 of the 14 arrays, and significantly more than half in six of these 12. However, in eight of the arrays, large plants sired significantly fewer seeds than expected on the basis of their size advantage. Furthermore, there was no significant relationship between relative size and relative siring success in comparisons among arrays. A maximum-likelihood model estimated that 28% of seeds were sired by imported pollen, with 95% confidence limits of 13% and 50%. Within these limits, high import rates tended to mask the relative success of large plants in several arrays. These results suggest that the evolution of inflorescence size in Z. paniculatus is at least partly driven by selection for increased male success, assuming genetic variation for flower number. However, the data also support a growing body of evidence that estimates of male fitness in plants can be highly variable. We discuss the sources of this variability and the possible effects of inflorescence design on the relationship between inflorescence size and fitness. 相似文献
13.
Body size-dependent gender role in a simultaneous hermaphrodite freshwater snail, Physa acuta 总被引:1,自引:0,他引:1
We examined whether gender role in the simultaneous hermaphroditefreshwater snail, Physa acuta, is determined by relative bodysize in a manner predicted by the size-advantage model. We observedthe body-size combinations of pairs in the laboratory by usingfield-collected populations. Smaller individuals tended to playthe "male" role (sperm donor), and larger snails the "female"(sperm recipient). Next, we analyzed the mating behaviors involvedin gender-role decision in snail pairs of three different body-sizecombinations, using "large" and "small" snails. Smaller snailswere more likely to approach the partner as a male in different-sizecombination (large/small), whereas frequent initial approachesas a male and rejection behavior as a female were observed inthe large/large combination. Third, we examined the body sizepreference when a snail can freely choose the partner from twoother individuals of different body sizes (large/large/smallor large/small/small). Small individuals had a significant tendencyto act as the male and positively selected large snails as thefemale partner in both triple combinations. However, the largeindividual acted as both the male and the female with nearlyequal frequency. In the size-differing pairings, copulationsoccurred after fewer male approaches and fewer rejections thanin pairings involving two large snails, suggesting that bodysize difference is one of the behavioral solutions in genderconflict. Clear gender-role switching associated with body sizewas not seen. Smaller snails thus have a tendency to play themale role more frequently but adopt both gender roles when theirbody size is sufficiently large. 相似文献
14.
The cestode Schistocephalus solidus is a facultatively self-fertilising simultaneous hermaphrodite. Here we test for differences in the starting point, the rate,
and the magnitude of egg production between individuals allowed to reproduce alone (only self-fertilisation possible) or in
pairs (both self- and cross-fertilisation possible). Specifically, we want to distinguish between alternative processes responsible
for the lower egg production in paired individuals observed in an earlier study (Wedekind et al., 1998). We designed an improved in vitro system, replacing the bird final host that allows us to measure, with high temporal resolution, the timing and magnitude
of lifetime egg production of worms in these two social situations. We found that the experimental groups did not differ significantly
in the starting point of egg production. However, the temporal pattern in egg production differed between them, in that paired
individuals had a lower rate of egg production. This, however, did not lead to a significant reduction in lifetime egg production,
as pairs compensated for the lower rate by producing eggs longer than single individuals. We argue that the lower rate of
egg production may nevertheless lead to a time cost of pairing in the study species, and that this cost is likely to represent
a cost of outcrossing due to sexual selection.
This revised version was published online in July 2006 with corrections to the Cover Date. 相似文献
15.
Individuals within a population often differ considerably in size or resource status as a result of environmental variation. In these circumstances natural selection would favour organisms not with a single, genetically determined allocation, but with a genetically determined allocation rule specifying allocation in relation to size or environment. Based on a graphical analysis of a simple evolutionarily stable strategy (ESS) model for herbaceous perennial plants, we aim to determine how cosexual plants within a population should simultaneously adjust their reproductive allocation and sex allocation to their size. We find that if female fitness gain is a linear function of resource investment, then a fixed amount of resources should be allocated to male function, and to post‐breeding survival as well, for individuals above a certain size threshold. The ESS resource allocation to male function, female function, and post‐breeding survival positively correlate if both male and female fitness gains are a saturating function of resource investment. Plants smaller than the size threshold are expected to be either nonreproductive or functionally male only. 相似文献
16.
Multiple mating, paternity, and body size in a simultaneous hermaphrodite, Aplysia californica 总被引:2,自引:0,他引:2
Sperm displacement and sperm competition prove difficult tomeasure, but are crucial elements in predicting sex allocationstrategies of sperm-storing hermaphrodites. Body size is predictedto affect sex allocation so that within a population, largeanimals invest a greater proportion of resources in female functionthan do small animals. These mating strategies depend on spermdisplacement abilities and lead to similar levels of paternityacross body sizes despite differences in resource level. Thepresent study investigated mating patterns, multiple paternity,and sperm competition in a field population of a simultaneouslyhermaphroditic sea slug, Aplysia californica (California seahare). Animals mating in the female role were larger than themean for the population, indirectly supporting theoretical predictionsfor increased investment in female function with body size.However, contrary to predictions, animals mating in the malerole were not different in size from the population mean orthe animals they inseminated. Individual tagging revealed thatsea slugs are capable of moving across distances that allowfor the sampling of many potential mates, and that they materepeatedly in both sexual roles. Microsatellite paternity analysisdemonstrated that multiple mating in the field leads to multiplepaternity, and last-sperm donors achieve high levels of paternity.There was no effect of body size on paternity. Further paternitystudies are needed to reveal the mechanisms of sperm precedencepatterns in A. californica. 相似文献
17.
Social group size may affect the potential for sperm competition, and this in turn may favour ontogenetic adjustments in testicular mass according to the likely requirements for sperm and spermatophore production. In a number of comparative analyses of testis mass among vertebrate species that differ in mating system or social organization, increasing potential for sperm competition is associated with larger testis size. Intraspecific phenotypic plasticity should be able to produce the same pattern if social group size is heterogenous and reflects differing degrees of average sperm competition, but this intraspecific effect is less well studied. We tested the effect of social groups on both male and female investment in the simultaneously hermaphroditic leech, Helobdella papillornata. Leeches were placed in groups of one, two, four or eight. Sexual investment at the onset of reproductive maturity was quantified as the total testisac volume for male function and total egg volume for female function. We found that testisac volume (statistically adjusted for body size) showed a significant increase with increasing group size. Total egg volume (also adjusted for body size) was unaffected by group size. Our findings indicate adaptive developmental plasticity in male gonad investment in response to the potential for sperm competition. 相似文献
18.
Sex allocation theory for simultaneous hermaphrodites assumes a direct trade-off between the allocation of resources to the male and female reproductive functions. Empirical support for this basic assumption is scarce, possibly because studies rarely control for variation in individual reproductive resource budgets. Such variation, which can have environmental or genetic sources, can generate a positive relationship between male and female investment and can thus obscure the trade-off. In this study on the hermaphroditic flatworm Macrostomum sp. we tried to control for budget effects by restricting food availability in a standardized way and by using an inbred line. We then manipulated mating group size in a two-way design (two group sizes x two enclosure sizes) in order to induce phenotypic variation in male allocation, and expected to find an opposing correlated response in female allocation. The results suggest that we only managed to control the budget effects under some conditions. Under these the sex allocation trade-off emerged. Under the other conditions we found a strongly positive correlation between male and female allocation. We discuss possible causes for the observed differences. 相似文献
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Although species with both male and female sexual functionsare often dichotomized into simultaneous and sequential hermaphrodites,many simultaneous hermaphrodites also exhibit sequential changesin sex allocation. In a field experiment using one such species,the gobiid fish, Lythrypnus dalli, female-biased individualsreallocated to male function in relation to their relative bodysize: consistent with the sizeadvantage hypothesis, large femaleswere more likely to reallocate and large fish had the highestspawningrates. Individuals, despite internal allocation to bothsexual functions, adopted only one behavioral gender. Behavioralmales had higher reproductive rates than behavioral females,and laboratory experiments showed that females preferred tomate with large males. Behavioral males grew more rapidly anddid notdiffer from behavioral females in survivorship. In addition,individuals who adopted male behavior but did not receive eggsin their nests maintained high levels of female tissue, whereasmales that received eggs did not. Laboratory experiments showedthat, unlike most hermaphroditic animals, L. dalli canchangeallocation either from female to maleor from male to female. Thus, L.dalli shares haracteristics of both sequential and simultaneoushermaphrodites. Simultaneous hermaphroditism maybe maintained,in this species, to facilitate rapid sex change from femaleto male and to retain flexibility o that unsuccessful malescan revert to reproduction as females. 相似文献