Degree Of Sperm Competition Research Articles

Strategic male mating behaviour in Argiope lobata

In mating systems with strictly limited male mating opportunities, a male's reproductive success depends strongly on the quality of his mate. Female quality is often linked to body size and mass. Hence, males with limited mating opportunities should choose large and heavy females to maximize their reproductive success. However, high-quality females are likely to attract more than one male, which consequently results in male–male competition. This scenario can be found in the spider species Argiope lobata. Males can copulate twice at most and eventually die after mating, but can monopolize females through the application of mating plugs. These plugs are costly because males block the female's genital openings with parts of their sperm-transferring organs (pedipalps), rendering them dysfunctional. Plugging moreover increases the risk of sexual cannibalism. We tested whether male A. lobata adjust their mating investment to the expected gains, which are a function of female fecundity and the risk of sperm competition. We presented males with virgin females of different fecundity, in either the presence or absence of a rival. Indeed, we found that males were selective and preferred larger females under competitive conditions. When a rival was present and the female was small, males frequently performed ultrashort copulations of less than 3 s without sperm transfer. Ultrashort copulations may imply that males were not willing to invest in paternity under these conditions. In contrast, when two rivals competed for heavy and thus fecund females, males were significantly more likely to monopolize paternity by depositing a plug. In conclusion, our results suggest that males try to adapt their copulatory behaviour to female quality and the degree of sperm competition even in ‘dangerous’ mating systems.

Post-Coital Sperm Competence in Polygamous Animals: The Role of Sperm Traits in Species?Specific Strategies

In some species, the females may mate with more than one male. In these polygamous animals, both pre-coital and post-coital strategies are required for reproductive success. This mini review aims at providing an update of the role of sperm traits in species–specific strategies for post-coital sperm competence. According to the spermcompetition theory, in such species the reproductive success of the males then depends upon both the magnitude of inter-male rivalry and covert sperm competition. The number of inseminated sperm, the percentage of sperm without morphological abnormalities in the ejaculates, and the sperm velocity variables play a significant role in sperm competition. Therefore, males that transfer more sperm per ejaculate or who copulate more often, are more likely to sire offspring. The transfer of large numbers of sperm is also guaranteed by ejaculates with high sperm concentrations. Usually, males of polygamous species have large testes and produce ejaculates with high sperm concentrations. Sperm competition acts as a selective force driving evolutionary change in sperm morphology and function. The degree of sperm pleiomorphy shown by a species varies according to the degree of sperm competition its behaviour entails. For example, the ejaculates of very polygamous species have much more homogenous sperm with lower sperm abnormalities. Finally, recent studies in birds have shown that the straight-line velocity (VSL) value, the most accurate estimate of sperm cell velocity, is of great importance in competition scenarios. Thus, sperm with a higher VSL affords a male fertilization advantages over his rivals.

Testis asymmetry in birds: the influences of sexual and natural selection

Gonad size and shape asymmetries are particularly common in birds. Although some obvious size and shape differences between the left and right testes in birds were first documented more than a century ago, little is known about what influences the variation across species in either the degree or the direction of these asymmetries. Here we show that a left bias in size is the most likely ancestral state in most orders and families, and that there is a weak but significant negative relation between the degree of size and shape asymmetries. In extant species, testis size and shape symmetries increase with the degree of sperm competition (relative testes mass), but those relations are significant only in those species with left bias in each of these traits. When space is particularly constrained on the left side of the body cavity due to a large gizzard (e.g. in granivores), the left testis is more elongated and the degree of size symmetry is larger. Despite significant patterns, relative testes mass and gizzard complexity together explain < 10% of the variation in testis asymmetry in species with left biases. Thus our analyses suggest that some other factor is largely responsible for the evolution of gonad asymmetry in birds: 1) that a left bias in testis size might be a correlated response to selection for a left bias in the development of a single (left) ovary in females of most bird species, and/or 2) that physiological efficiency due to the dramatic and rapid increase in testis size of most species during the breeding season might favour enlargement of one testis. Our conclusions highlight the need to rethink and improve our understanding of the physiological processes underlying the investment in gonads by male birds.

Population density does not influence male gonadal investment in the Least Killifish,Heterandria formosa

Comparative studies documenting a relationship between male gonadal investment and the degree of sperm competition (SC) have usually considered the association between these traits to be driven by qualitative differences in the mating system, such as whether spawning occurs in pairs or groups. However, ecological and demographic differences between conspecific populations may also generate variation in the importance of SC that can drive the evolution of male gonadal investment. In this study, we examined whether variation in population density, which is predicted to influence the level of SC in many animals, is correlated with male gonadal investment among populations of the least killifish, Heterandria formosa, a species with internal fertilization in which multiple mating is common. We complemented this field study by testing whether males respond plastically to experimentally increased levels of SC by increasing investment in testis. This experiment involved two treatments. In the first, we eliminated the potential for sperm competition (NSC) by housing a single male with a single female. In the second, we created a high risk of SC by housing five males with two females. In the field survey, we found significant differences among populations in density and relative testis mass. However, there was no evidence for a correlation between population density and relative testis mass. In our lab experiment, males did not adjust their gonadal investment in response to experiencing different levels of SC for 4 weeks. Our combined results indicate that gonadal investment in male H. formosa is not related to variation in population density.

Information content of female copulation calls in wild long-tailed macaques (Macaca fascicularis)

Primates are unusual in that many females display sexual signals, such as sex skin swellings/colorations and copulation calls, without any sex role reversal. The adaptive function of these signals remains largely unclear, although it has been suggested that they provide males with information on female reproductive status. For sex skin swellings, there is increasing evidence that they represent a graded signal indicating the probability of ovulation. Data on the functional significance of copulation calls are much scarcer. To clarify the information content of such calls, we recorded copulation calls in wild long-tailed macaques (Macaca fascicularis) and analysed the structure of these calls during the ovarian cycle. Specifically, we correlated selected call parameters with the female oestrogen to progestogen ratio (obtained from faecal samples), which are known to be elevated during the female's fertile phase. In addition, we ran a general linear mixed model for these call parameters, testing factors (cycle phase, occurrence/absence of ejaculation, male dominance status, occurrence/absence of mate guarding) which potentially influence female copulation calls in primates. Our results show that copulation calls of female long-tailed macaques signal mating outcome and rank of the mating partner, but not female reproductive status. They also show for the first time on primates that copulation calls can convey information on whether a female is mate guarded or not. We suspect that the function of these calls is manipulation of male mating and mate-guarding behaviour and that in this way the degree of sperm competition and ultimately male reproductive success is influenced.

Speciation and the evolution of gamete recognition genes: pattern and process

Proteins on gamete surfaces are major determinants of fertilization success, particularly in free-spawning animals. Molecular analyses of these simple genetic systems show rapid evolution, positive selection, accelerated coalescence and, sometimes, extensive polymorphism. Careful analysis of the behavior of sperm produced by males with different gamete alleles shows that these alleles can deliver significant functional differences. Three forms of allele-specific fertilization advantage have been shown: assortative mating based on gamete type, rare allele advantage and heterozygote superiority. Models suggest that sperm and egg proteins may be coevolutionary partners that can alternate between directional selection for high fertilization ability and cyclic adaptation of eggs and sperm driven by sexual conflict. These processes act within allopatric populations and may accelerate their divergence if gamete adaptations in separate demes reduce cross-fertilization. Reproductive character displacement by reinforcement may play a diversifying role when previously allopatric populations rejoin. In circumstance that might prove to be common, divergence in sympatry can be driven by sexual conflict or by association of mating types with ecological differences. The ecology of fertilization, especially the degree of sperm competition and egg death via polyspermy, are important determinants of the strength and direction of selection on gametes. Free-spawning animals allow careful analysis of gamete recognition -from the behavior of adults and interactions of gametes, to molecular patterns of allele divergence. Future research efforts on the evolutionary consequences of fertilization ecology, and the interaction between extensive variation in egg surface proteins and sperm fertilization ability, are particularly needed.

Sperm Allocation Strategies and Female Resistance: A Unifying Perspective

The classical viewpoint in sperm competition theory, which holds that males evolve sperm allocation strategies in response to a given degree of sperm competition, has recently been challenged by an alternative viewpoint, which holds that the degree of sperm competition is itself a consequence of these same strategies. Here, we present a game theory model that unites these alternative views as the endpoints of a continuum. Based on the recognition that female control over mating may limit the extent to which male strategies affect the degree of sperm competition, we investigate sperm allocation strategies in a setting where females can resist excessive matings more or less successfully. We discuss how conflicting predictions made by previous theory relate to implicit assumptions about female resistance behavior. Moreover, we show that female resistance, while being highly relevant to the predicted relationship between ejaculate size and the degree of sperm competition, has little effect on the predicted positive correlation between relative testis size and the degree of sperm competition. This result strengthens one of the central predictions of sperm competition theory and is in accordance with empirical findings from a wide range of taxa.

Sex‐Specific Spawning Behavior and Its Consequences in an External Fertilizer

Identifying the target of sexual selection in externally fertilizing taxa has been problematic because species in these taxa often lack sexual dimorphism. However, these species often show sex differences in spawning behavior; males spawn before females. I investigated the consequences of spawning order and time intervals between male and female spawning in two field experiments. The first involved releasing one female sea urchin's eggs and one or two males' sperm in discrete puffs from syringes; the second involved inducing males to spawn at different intervals in situ within a population of spawning females. In both, fertilization success was measured as the fraction of eggs fertilized and the paternity share of each male. The results indicate that spawning after females imposes a cost on males but only during sperm competition. Further, the optimal interval between the initiations of male and female spawning depends on degree of sperm competition, distance between males and females, and water velocity. The results show that sex differences in spawning timing of marine invertebrates can be explained on the basis of the differential costs and benefits of spawning out of synchrony with the other sex and that the result of sexual selection on external fertilizers may be behavioral rather than morphological differentiation of the sexes.

Sex-Specific Spawning Behavior and Its Consequences in an External Fertilizer

Identifying the target of sexual selection in externally fertilizing taxa has been problematic because species in these taxa often lack sexual dimorphism. However, these species often show sex differences in spawning behavior; males spawn before females. I investigated the consequences of spawning order and time intervals between male and female spawning in two field experiments. The first involved releasing one female sea urchin’s eggs and one or two males’ sperm in discrete puffs from syringes; the second involved inducing males to spawn at different intervals in situ within a population of spawning females. In both, fertilization success was measured as the fraction of eggs fertilized and the paternity share of each male. The results indicate that spawning after females imposes a cost on males but only during sperm competition. Further, the optimal interval between the initiations of male and female spawning depends on degree of sperm competition, distance between males and females, and water velocity. The results show that sex differences in spawning timing of marine invertebrates can be explained on the basis of the differential costs and benefits of spawning out of synchrony with the other sex and that the result of sexual selection on external fertilizers may be behavioral rather than morphological differentiation of the sexes.

Male Competition over Access to Females in a Spider with Last‐Male Sperm Precedence

AbstractAgonistic behaviour between male cellar spiders (Pholcus phalangioides) was investigated to test whether (1) size difference determines which male achieves access to the female, (2) males are able to monopolize access to the female until egg laying and whether (3) female resource value increases before egg laying because of last‐male sperm precedence. We further investigated whether (4) there is variation in time and energy spent on courtship and copulation depending on the degree of sperm competition, i.e. with or without rival present. In three experimental settings we introduced two males of either different or similar sizes, or a single male to a female. The mating units were constantly video‐observed until the females produced their first egg sac. Experience, ownership and female resource value in terms of body size was controlled. Our results show that larger males achieve almost exclusive access to females. Size symmetrical settings resulted in increased fighting activity and duration but dominance did not influence mating success. If copulations were disturbed by the rival male, copulations were terminated earlier in symmetrical settings compared with asymmetrical settings. In 94.8% of trials only one copulation took place, suggesting that the copulating male successfully monopolized access to the female. Males confronted with a rival copulated longer but courted significantly shorter than lone males. Although the last male to copulate sires 88% of the offspring in P. phalangioides, neither fighting nor courtship activity increased before the female laid a batch of eggs. This suggests that males have no indication of the timing of oviposition.

Reproductive Anatomy of the Reed Bunting: A Species Which Exhibits a High Degree of Sperm Competition through Extra-Pair Copulations

The morphology of the reproductive organs of both male and female Reed Buntings (Emberiza schoeniclus) are described in relation to the high level of sperm competition exhibited by the species as a result of extra-pair copulation behavior. In males, the volume of the cloacal protuberance in relation to body weight was 26.7 mm 3 g -1 , which is relatively large for a socially monogamous passerine. The weight of the testes was 23% greater than that expected for a bird of comparable size, and the spermatozoa were among the longest recorded for any avian species (mean length = 291 μm). The morphology of the female tract also showed adaptations associated with a high level of sperm competition: the sperm storage tubules were extremely long (mean length = 865 μm).

Sperm allocation, sperm production and fertilization rates in the bucktooth parrotfish

Males and females of pelagically spawning marine fishes face different problems in maximizing reproductive success. Males must allocate sperm over frequent, successive spawns. Females must ensure successful fertilization of their egg clutches. This study examined sperm allocation between individual spawns, the daily ‘standing crop’ of sperm just prior to spawning, gonadal investment (testis weight/body weight) in males and fertilization rates of females of the coral reef fish Sparisoma radians. Males released a median of 8.94×10 6sperm per pair spawn, with the number increasing with increasing clutch size and body size of the female mate. Standing crop of sperm correlated with male spawning frequency, and testicular investment correlated with degree of sperm competition by streaking males. Fertilization efficiency (number of eggs fertilized by a given number of sperm) increased with the body size of the female partner. Fertilization rate, i.e. the proportion of the clutch successfully fertilized, was over 90% in most spawns, but significantly increased with the number of sperm released by the male. The probability that a spawn was joined by a second, competing male (‘streaker’) increased with the body size of the spawning female. Overall, the sperm allocation pattern can be explained both as a form of sperm economy in which males release the minimum number of sperm needed to fertilize most of the eggs in clutches of different size, and as a response to an increased probability of sperm competition with females of increasing size.

Mass and production rate of ejaculates in relation to monandry/polyandry in butterflies

The mating system maintained in a species has a strong effect on the degree of sperm competition, and certainty of paternity should accordingly influence the optimal sperm content, nutrient content, and mass of the ejaculate. We investigated how ejaculate mass relates to the degree of polyandry in 20 species of butterflies belonging to the families Pieridae and Satyridae. We found that the degree of polyandry has a substantial effect on the reproductive performance of males. The allometric line between ejaculate mass and male body mass has a higher elevation in the pierids compared to the satyrids. The mean number of matings performed by the pierid species is also higher compared to the mean of the satyrids. Thus, the relative ejaculate mass is larger in the family in which polyandry is more pronounced. A within family effect of degree of polyandry on relative ejaculate mass was also detected in the pierids. Since males of polyandrous species on average mate more often than males of monandrous species, they should be expected to have a higher capacity for producing many ejaculates. We investigated how this capacity was influenced by the degree of polyandry, by allowing males of seven different species (Danaus plexippus, Lasiommata megera, Papilio machaon, Pararge aegeria, Pieris napi, Pieris rapae, and Polygonia c-album) to mate twice, with different time intervals between matings. The results showed that not only is the mass of the ejaculate greater in more polyandrous species, but also the rate at which males are able to produce sperm and accessory substances is greater. Hence our data indicate that sperm competition is important for explaining variation in ejaculate mass in butterflies.