The copulatory plug delays ejaculation by rival males and affects sperm competition outcome in house mice

Females of many species mate with multiple males (polyandry), resulting in male–male competition extending to post‐copulation (sperm competition). Males adapt to such post‐copulatory sexual selection by altering features of their ejaculate that increase its competitiveness and/or by decreasing the risk of sperm competition through female manipulation or interference with rival male behaviour. At ejaculation, males of many species deposit copulatory plugs, which are commonly interpreted as a male adaptation to post‐copulatory competition and are thought to reduce or delay female remating. Here, we used a vertebrate model species, the house mouse, to study the consequences of copulatory plugs for post‐copulatory competition. We experimentally manipulated plugs after a female's first mating and investigated the consequences for rival male behaviour and paternity outcome. We found that even intact copulatory plugs were ineffective at preventing female remating, but that plugs influenced the rival male copulatory behaviour. Rivals facing intact copulatory plugs performed more but shorter copulations and ejaculated later than when the plug had been fully or partially removed. This suggests that the copulatory plug represents a considerable physical barrier to rival males. The paternity share of first males increased with a longer delay between the first and second males' ejaculations, indicative of fitness consequences of copulatory plugs. However, when males provided little copulatory stimulation, the incidence of pregnancy failure increased, representing a potential benefit of intense and repeated copulation besides plug removal. We discuss the potential mechanisms of how plugs influence sperm competition outcome and consequences for male copulatory behaviour.     

Competitive exclusion through reproductive interference

A simple differential equation model was developed to describe the competitive interaction that may occur between species through reproductive interference. The model has the form comparable to Volterra's competition equations, and the graphical analysis of the outcome of the two-species interaction based on its zero-growth isoclines proved that: (1) The possible outcome in this model, as in usual models of resource competition, is either stable coexistence of both species or gradual exclusion of one species by the other, depending critically upon the values of the activity overlapping coefficient c_ij; (2) but, for the same c_ij-values, competitive exclusion is much more ready to occur here than in resource competition; (3) and moreover, the final result of the competition is always dependent on the initial-condition due to its non-linear isoclines, i.e., even under the parameter condition that generally allows both species to coexist, an extreme bias in intial density to one species can readily cause subsequent complete exclusion of its counterparts. Thus, it may follow that the reproductive interference is likely to be working in nature as an efficient mechanism to bring about habitat partitioning in either time or space between some closely related species in insect communities, even though they inhabit heterogeneous habitats where resource competition rarely occurs so that they could otherwise attain steady coexistence.     

Competition in a turbidostat for an inhibitory nutrient

A model of competition in a turbidostat between two species for an inhibitory growth-limiting nutrient is considered. It is shown that the model has rich dynamics. A coexistence equilibrium and the washout equilibrium can be asymptotically stable simultaneously so that coexistence may depend on initial conditions. Under certain conditions, periodic coexistence of the two species occurs. There is a possibility that two species coexist, whereas one species dies out in the absence of its rival.     

Analysis of species coexistence co-mediated by resource competition and reproductive interference

Reproductive interference is any interspecific sexual interaction that reduces the reproductive success of females through promiscuous reproductive activities of heterospecific individuals. This phenomenon is ubiquitous in nature in both plants and animals, and is frequently observed in biological invasions. However, its effects on interspecific competition remain incompletely understood despite growing concern. To study the interactive effects of resource competition and reproductive interference on species coexistence and exclusion, we analyzed a unified competition model including both processes in symmetric and asymmetric scenarios. The results of our model showed that resource competition and reproductive interference act synergistically to promote competitive exclusion. We also found that when the two processes are asymmetric, the species that is superior in reproductive interference can coexist with or even exclude the species that is superior in resource competition. Therefore, coexistence is possible via an unbalanced trade-off between resource use and reproduction. Our results suggest that integration of reproductive interference and resource competition will contribute to a better understanding of interspecific competition and to more effective biodiversity conservation against management of biological invasions.     

A model for the evolutionary maintenance of monogyny in spiders

Sexual selection theory predicts that males should attempt to mate with several females, unless the benefits of male promiscuity are trumped by alternative benefits associated with male monogamy (monogyny). Here we use a game theory model to address the adaptive value of a monogynous strategy, which has the sole benefit of enhancing a male's paternity share in the context of competition with other males. We consider two ways in which monogynists might enhance their paternity: by outcompeting rival ejaculates in sperm competition, and by reducing the probability that a female remates with rival males. The model is based on the biology of some particularly well-studied spider species, in which males are morphologically restricted to mate with either one or at most two females in their lifetime. Our results suggest that, regardless of the mechanism of paternity enhancement involved, a male-biased sex ratio is generally required for the evolution and maintenance of monogyny. Moreover, we show that there is a large region of parameter space where monogyny and bigyny can coexist as alternative mating strategies under negative frequency dependent selection. There is also a narrow range of conditions where either monogyny or bigyny can be evolutionarily stable. Our results are in qualitative agreement with empirical findings in spiders.     

Me against who? Male guppies adjust mating behaviour according to their rival’s presence and attractiveness

Sexual selection theory suggests that males need to constantly reappraise their mating decisions to take account of the presence and the phenotypes of their rivals. Here we examine this expectation by asking: (i) If the presence of a rival influences male mating behaviour; (ii) How important is the attractiveness of the rival (absolute attractiveness) in shaping male behaviour; and (iii) How does a male's attractiveness in comparison to his rival (relative attractiveness) influence a male's mating decisions. Using the Trinidadian guppy, a species in which female mate choice (based on males’ attractive traits) plays an important role in male mating outcomes, we recorded the frequency of courtship displays and unsolicited attempts by focal males. First, we quantified focal male mating behaviour with and without a rival. Since the probability of a successful mating is, on average, halved by the presence of a rival, we predicted that under competition the focal male would invest more in less costly mating tactic—unsolicited attempts. Second, we examined how the rival's standard length and area of orange coloration mediated focal male mating behaviour. We found that rival presence influenced how focal males responded to females in terms of both mating tactics. However, the rival attractiveness elicited changes only in male courtship display. Focal males increased courtship display rate if his rival was small or if possessed large amounts of orange, regardless of considering rival absolute or relative attractiveness. Our results show that males invest in the costlier mating tactic when there is no rival or in the presence of a smaller rival. Interestingly, they make a similar investment in the presence of an attractive orange rival. Overall, this study highlights the importance of fine‐grained male decisions in mating encounters and shows that mating tactics are differentially shaped by multiple competition risk cues.     

Sexual Interference and Sexual Defense in the Smooth Newt,Triturus vulgaris (Amphibia,Urodela, Salamandridae)

• 1 When a male smooth newt encounters a ♀ who is already engaged in courtship, he may mimic her behaviour during the spermatophore deposition and transfer stages of the courtship. He thereby usurps the courting ♂ and may inseminate the ♀ himself. Such sexual interference depresses the short-term, and perhaps long-term, mating success of the courting ♂.
• 2 In the presence of a potential rival, the courting ♂ alters certain aspects of his sexual behaviour. He displays more intensely to the ♀ and attempts to draw her away from the rival by increasing the duration of his display. He may also “check” that it is the ♀, and not the rival, who will elicit the deposition of a spermatophore from him. These changes in the behaviour of the courting ♂ are interpreted as defense against sexual interference.
• 3 Female smooth newts may be multiply inseminated as a consequence of sexual interference; this may result in sperm competition. However, ♀♀ seem to find competitive interactions between ♂ ♂ “aversive”.
• 4 Sexual interference by ♀-mimicry and associated defensive behaviour patterns are common in the urodele amphibians. Interference can be thought of as a “side-payment” conditional mating strategy.

     

The effects of interference competition on stability,structure and invasion of a multi-species system

Summary An interference competition model for a many species system is presented, based on Lotka-Volterra equations in which some restrictions are imposed on the parameters. The competition coefficients of the Lotka-Volterra equations are assumed to be expressed as products of two factors: the intrinsic interference to other individuals and the defensive ability against such interference. All the equilibrium points of the model are obtained explicitly in terms of its parameters, and these equilibria are classified according to the concept of sector stability. Thus survival or extinction of species at a stable equilibrium point can be determined analytically.The result of the analysis is extended to the successional processes of a community. A criterion for invasion of a new species is obtained and it is also shown that there are some characteristic quantities which show directional changes as succession proceeds.     

Rival male relatedness does not affect ejaculate allocation as predicted by sperm competition theory

When females are sexually promiscuous, the intensity of sperm competition for males depends on how many partners females mate with. To maximize fitness, males should adjust their copulatory investment in relation to this intensity. However, fitness costs associated with sperm competition may not only depend on how many males a female has mated with, but also how related rival males are. According to theoretical predictions, males should adjust their copulatory investment in response to the relatedness of their male rival, and transfer more sperm to females that have first mated with a non-sibling male than females that have mated to a related male. Here, for the first time, we empirically test this theory using the Australian field cricket Teleogryllus oceanicus. We expose male crickets to sperm competition from either a full sibling or non-sibling male, by using both the presence of a rival male and the rival male's actual competing ejaculate as cues. Contrary to predictions, we find that males do not adjust ejaculates in response to the relatedness of their male rival. Instead, males with both full-sibling and non-sibling rivals allocate sperm of similar quality to females. This lack of kin biased behaviour is independent of any potentially confounding effect of strong competition between close relatives; kin biased behaviour was absent irrespective of whether males were raised in full sibling or mixed relatedness groups.     

Interference competition and species coexistence

Interference competition is ubiquitous in nature. Yet its effects on resource exploitation remain largely unexplored for species that compete for dynamic resources. Here, I present a model of exploitative and interference competition with explicit resource dynamics. The model incorporates both biotic and abiotic resources. It considers interference competition both in the classical sense (i.e. each species suffers a net reduction in per capita growth rate via interference from, and interference on, the other species) and in the broad sense (i.e. each species suffers a net reduction in per capita growth rate via interference from, but can experience an increase in growth rate via interference on, the other species). Coexistence cannot occur under classical interference competition even when the species inferior at resource exploitation is superior at interference. Such a trade-off can, however, change the mechanism of competitive exclusion from dominance by the superior resource exploiter to a priority effect. Now the inferior resource exploiter can exclude the superior resource exploiter provided it has a higher initial abundance. By contrast, when interference is beneficial to the interacting species, coexistence is possible via a trade-off between exploitation and interference. These results hold regardless of whether the resource is biotic or abiotic, indicating that the outcome of exploitative and interference competition does not depend on the exact nature of resource dynamics. The model makes two key predictions. First, species that engage in costly interference mechanisms (e.g. territoriality, overgrowth or undercutting, allelopathy and other forms of chemical competition) should not be able to coexist unless they also engage in beneficial interference mechanisms (e.g. predation or parasitism). Second, exotic invasive species that displace native biota should be superior resource exploiters that have strong interference effects on native species with little or negative cost. The first prediction provides a potential explanation for patterns observed in several natural systems, including plants, aquatic invertebrates and insects. The second prediction is supported by data on invasive plants and vertebrates.     

Linked exploitation and interference competition drives the variable behavior of a classic predator–prey system

The potential connection between exploitation and interference competition was recognized long ago but has not been evaluated. We measured the levels of both forms of competition for the protist Didinium preying upon Paramecium. Across populations, exploitation intensity was tightly linked to interference intensity, and the form of this relationship follows from a simple model of interaction speeds. The variation in interference competition was as large across populations of Didinium as has been observed previously across species from a variety of taxa including birds, mammals, insects, crustaceans, flatworms and protists. The link between exploitation and interference competition alters our understanding of how interference competition influences population dynamics. Instead of simply stabilizing systems, variation in interference levels can shift population dynamics through qualitatively different regimes because of its association with exploitation competition. Strong interference competition pushes a system to a regime of deterministic extinction, but intermediate interference generates a system that is stable with a high competitive ability. This may help to explain why the distribution of interference values is unimodal and mostly intermediate in intensity. Synthesis Exploitation and interference competition are typically viewed as separate processes. Exploitation is described with a functional response in which the inclusion of interference competition – the effect of predator density on foraging rates – is optional. Although recent work indicates that interference competition is widespread, there is little work linking the two forms of competition. In this article we present evidence that exploitation and interference competition are linked mechanistically through movement patterns that simultaneously generate beneficial interactions of consumers with resources and detrimental interactions with other consumers. This connection alters our view of the role that interference plays in ecological dynamics.     

Host specialization by reproductive interference between closely related herbivorous insects

Host specialization among closely related herbivores is common and is therefore a major theme in ecology. Despite its ecological importance, no general explanatory framework for host specialization is currently available. We introduce a new model of the evolution of host specialization in herbivorous insects. We use a two-dimensional lattice comprising 32 × 32 cells. Moreover, the model incorporates reproductive interference, defined as any negative outcomes resulting from interspecific mating, between two species of herbivores feeding on two different host plants as well as ordinary resource competition and differences in host suitability. Our simulation showed that reproductive interference, together with other factors such as host-related performance and resource competition, can actually drive the evolution of host specialization and thereby host partitioning between herbivorous insects during secondary contact. Host specialization arises particularly when levels of both reproductive interference and resource competition are intermediate. In contrast, host specialization seldom occurs if reproductive interference is absent. Thus, reproductive interference is as key to specialization as is spatial partitioning, which was not regarded as a realistic outcome when only resource competition was considered.     

Mutual assessment via visual status signals in Polistes dominulus wasps

Many animals use signals to assess the fighting ability of rivals and reduce the cost of aggressive competition. However, little is known about how an individual''s own quality influences their signal assessment decisions. Polistes dominulus wasps have visual signals of fighting ability that provide a good model for testing the dynamics of rival choice. We found that rival assessment behaviour was influenced by the advertised quality of the individual, their rivals, and the interaction between individual and rival quality. Individuals of high advertised quality were more likely to challenge rivals and individuals of low advertised quality were more likely to be challenged. However, when choosing among two rivals with different advertised quality, individuals did not simply choose the lower quality rival. Instead, they only preferred the lower quality rival when there was a small difference between their own advertised quality and that of their rivals. Individuals were not choosy when both rivals advertised relatively high or relatively low quality. Therefore, although P. dominulus facial patterns function as conventional signals of fighting ability that provide valuable information about their bearer''s behavioural strategy, there is substantial variation in signal responses based on the relative intensity of the senders'' and receivers'' signals.     

Female Sexual Interference in the Smooth Newt,Triturus vulgaris vulgaris

Competition between unmated females for males was observed in the smooth newt, T. vulgaris, during the early part of the breeding season. Females interfered at the spermatophore-transfer stage of courtship, preventing a rival female from picking up the sperm mass. Interference often resulted in the rival female abandoning the courtship encounter. In many of the encounters (70%), the interfering female was inseminated. However, interference between females was not observed after the commencement of ovulation. By contrast, the incidence of sexual interference between males was low during the preovulatory period and increased significantly post-ovulation. Sexual interference in newts and salamanders has previously only been observed between males. During the greater part of the breeding season, when females are ovipositing, eggs are the limiting resource and males compete for access to receptive females. However, this study suggests that early in the breeding season, prior to commencement of ovulation, the limiting resource may temporarily be sperm, due to the physiological constraint of spermatophore production. This may lead to female competition for mates.     

Soil microbial biomass C and symbiotic processes associated with soybean after sulfentrazone herbicide application

Resource competition and chemical interference are mechanisms of interaction among plants that may occur simultaneously. However, both mechanisms are rarely considered together when modelling plant growth. We propose a new empirical model that estimates biologically significant parameters on both plant competition and chemical interference. The model is tested with data sets from different density-dependent experiments done with two species (the grass Lolium rigidum Gaud. and the legume Glycine max soya L.) subjected to a noxious chemical environment when growing (allelochemicals and herbicides, respectively). Hypotheses on the effect of allelochemicals and its interaction with density are tested using maximum likelihood ratio tests in order to ask, for these species, whether chemical interference is playing a significant role in the interactions among plants or on the contrary, whether interactions among plants are sufficiently explained by the resource competition. In all cases a significant interaction between chemicals and density is observed. This interaction is inconsistent with the hypothesis of only resource competition having an influence of plant biomass and suggests a significant density-dependent effect of chemicals on plant growth.     

Male crickets adjust ejaculate quality with both risk and intensity of sperm competition

Sperm competition theory predicts that males should increase their expenditure on the ejaculate with increasing risk of sperm competition, but decrease their expenditure with increasing intensity. There is accumulating evidence for sperm competition theory, based on examinations of testes size and/or the numbers of sperm ejaculated. However, recent studies suggest that ejaculate quality can also be subject to selection by sperm competition. We used experimental manipulations of the risk and intensity of sperm competition in the cricket, Teleogryllus oceanicus. We found that males produced ejaculates with a greater percentage of live sperm when they had encountered a rival male prior to mating. However, when mating with a female that presented a high intensity of sperm competition, males did not respond to risk, but produced ejaculates with a reduced percentage of live sperm. Our data suggest that males exhibit a fine-tuned hierarchy of responses to these cues of sperm competition.     

The evolution of strategic male mating effort in an information transfer framework

Sperm competition theory predicts that males should use cues indicating the risk and intensity of sperm competition to tailor their sperm investment accordingly. Rival males are an important source of social information regarding sperm competition risk. However, revealing such information may not be in the rival males' interest. Here, we use a theoretical approach based on informed and uninformed games to investigate when information transfer about sperm competition risk to competitors is beneficial for a male, and when it is not. The results show that signalling to potential future mates that a female has already mated is beneficial when the signalling male has a sperm competition disadvantage, whereas it is unfavourable when the signaller has an advantage. The reason for this counterintuitive result is that the rival males' optimal response is to reduce sperm investment when the signaller has a disadvantage and, conversely, to increase investment when the signaller has an advantage. Furthermore, we analysed scenarios where males use alternative reproductive tactics. In this situation, signalling the awareness of sperm competition risk rarely pays; instead, it is beneficial to maintain an information advantage. Thus, it may be beneficial for bourgeois males to accept cuckoldry instead of revealing their sperm competition awareness to reproductive parasites. These results provide new insight into the evolution of communication between rivals in the context of sperm competition.     

Ejaculate size varies with socio-sexual situation in an insect

Abstract.

• 1 Males operate within a finite energy budget and cannot produce limitless supplies of sperm. On the other hand, when a female mates with a second male while still containing fertile sperm from a rival male, selection should favour the male that inseminates more sperm. Optimal strategy should thus be for males to exercise discretion in the allocation of sperm to individual females. Assuming the outcome of sperm competition to be based on either the raffle or kamikaze principles, the sperm competition hypothesis predicts a positive association between the probability that the sperm will find themselves in competition with sperm from rival males and the number of sperm inseminated.
• 2 The beetle, Tenebrio molitor L., behaves in accordance with this hypothesis. Males accompanied by a rival male before and during mating inseminate more sperm per ejaculate than unaccompanied males. Accompanied males are also faster to initiate mating and more likely to show post-copulatory guarding. Adjustment of number of sperm inseminated was shown by males subjected to both long-term (5 days) and short-term (5 min) exposure to potential intrasexual competitors. Individual males exposed to both levels also demonstrated the ability to adjust their ejaculate according to socio-sexual situation.
• 3 We conclude that male T.molitor adjust the number of sperm they inseminate according to some perception of the risk of sperm competition.

     

Effects of competitive asymmetry on a local density model of plant interference

Although competition between plants is usually asymmetric (i.e. larger plants have a disproportionate effect on smaller plants) almost all models of plant competition at the local level have assumed symmetric competition. We add a simple version of competitive asymmetry to the local density neighborhood models of plant interference and population dynamics developed by Pacala & Silander (1985, Am. Nat. 125, 385-411; 1987, Oikos 48, 217-224) by assuming that plants within a neighborhood can be put in a linear dominance hierarchy based upon their initial size. The size of a focal plant is a function of the number of dominant and the number of subordinate neighbors within its neighborhood, with subordinate neighbors having less of an effect than dominant ones. Asymmetry prevents precipitous changes in focal plant size with changes in local density, making the relationship between focal plant size and local density hyperbolic, even if the symmetric model is not hyperbolic. Thus, asymmetry makes the model conform to the law of constant final yield, irrespective of the form of the relationship between plant size and local crowding. Asymmetry also prevents population dynamic oscillations in the model in cases in which it would occur in the absence of asymmetry. The results show that asymmetry has major effects on a model of local interference in plants, and point to the importance of including it in such models.     

The effects of male social environment on sperm phenotype and genome integrity

Sperm function and quality are primary determinants of male reproductive performance and hence fitness. The presence of rival males has been shown to affect ejaculate and sperm traits in a wide range of taxa. However, male physiological conditions may not only affect sperm phenotypic traits but also their genetic and epigenetic signatures, affecting the fitness of the resulting offspring. We investigated the effects of male‐male competition on sperm quality using TUNEL assays and geometric morphometrics in the zebrafish, Danio rerio. We found that the sperm produced by males exposed to high male–male competition had smaller heads but larger midpiece and flagellum than sperm produced by males under low competition. Head and flagella also appeared less sensitive to the osmotic stress induced by activation with water. In addition, more sperm showed signals of DNA damage in ejaculates of males under high competition. These findings suggest that the presence of a rival male may have positive effects on sperm phenotypic traits but negative effects on sperm DNA integrity. Overall, males facing the presence of rival males may produce faster swimming and more competitive sperm but this may come at a cost for the next generation.