Male Investment Research Articles

Variability of GnRH secretion in two goby species with socially controlled alternative male mating tactics

Male reproductive phenotypic plasticity related to environmental–social conditions is common among teleost fish. In several species, males adopt different mating tactics depending on their size, monopolizing mates when larger, while parasitizing dominant male spawns when smaller. Males performing alternative mating tactics are often characterized by a strong dimorphism in both primary and secondary reproductive traits. According to studies on sex-changing species and on species where only one male morph is reproductively active, male alternative phenotypes are expected to vary also in gonadotropin-releasing hormone (GnRH) neurons in forebrain preoptic area (POA). Here, we compared the intra- and inter-sexual variations in number and size of GnRH neurons, along with gonads and male accessory structure investment, in two goby species, the grass goby, Zosterisessor ophiocephalus, and the black goby, Gobius niger, characterized by male alternative mating phenotypes. In both species, older and larger males defend nests, court and perform parental care, while younger and smaller ones try to sneak territorial male spawning. We found that grass goby and black goby have different patterns of GnRH expression. Grass goby presents a clear intra-sexual dimorphism in GnRH expression, related to the occurrence of alternative mating tactics, while in the black goby, only inter-sexual differences are observed. The inter- and intra-specific variability in the GnRH neurons in these two goby species is discussed in light of the differences in migratory behavior, nest type, and mating system.

The role of testosterone in male downy woodpeckers in winter home range use, mate interactions and female foraging behaviour

Studies of the role of testosterone (T) in birds have typically focused on sexual or aggressive behaviours of males during the breeding period, but males of nonmigratory species may invest in mate and territory long before breeding, and the influence of T in facilitating nonbreeding-season behaviours is poorly understood. We gave free-living male downy woodpeckers, Picoides pubescens, T-implants during the winter to determine whether elevated levels of T increased a male's ability to exclusively occupy territory-based resources, and whether elevated T strengthened a male's investment in an existing pair bond relationship. We also explored how a female's foraging efficiency might be affected by her mate's behaviour if he had elevated T. We found little difference between control and T-implanted males with regard to home range exclusivity. Surprisingly, male-male display rates were significantly lower in T-implanted males than in controls. Regarding male-female interactions, T-implanted males that experienced high incursion rates from other males maintained more frequent spatial association with their mate, suggesting that T facilitates male behaviours that could restrict the mate's access to other male birds. Female mates of T-males showed reduced foraging rates, but because male-female aggression was similar between treatment groups, the cause for this reduction is unknown. The results indicate that exogenous T during winter affects a variety of behaviours in male woodpeckers, and proximate influences on pair bond maintenance in winter may be a fruitful avenue for future research.

The impact of male-killing bacteria on the evolution of aphidophagous coccinellids

Many aphidophagous coccinellids harbour male-killing bacteria. These maternally inherited bacteria kill males early in embryogenesis, female offspring of infected mothers gaining a large resource advantage from the consumption of their dead brothers. In this paper, the diversity of male-killing bacteria and their coccinellid hosts will be briefly reviewed. Thereafter, the impact that invasion by male-killers has on coccinellid hosts will be addressed in two ways. First, the selective effects due to the loss of male progeny, and the intra-genomic conflict resulting from the cytoplasmic inheritance of the bacteria will be considered. Reduc- tions in mitochondrial diversity, effects on clutch sizes and the evolution of male-killer suppresser systems will be discussed. Second, the impact of female biased population sex ratios on the evolution of reproductive strategies and the primary sex ratio will be investigated. Preliminary results suggesting changes in male investment per copulation, reductions in female testing behaviour of males, and male biases in the progenic sex ratios of some uninfected male lines will be presented. Avenues for future work will be outlined.

Sex differences in rates of territory joining and inheritance in a cooperatively breeding cichlid fish

In social groups, subordinates may gain dominant breeding status either by inheriting the top position in their current group or by dispersing to join a new group. The pathway to breeder status is likely to vary between males and females as a result of sex differences in the costs of dispersal and inbreeding. We report results from a field study conducted to explore sex differences in the rates of territory joining and inheritance in a cooperatively breeding cichlid, Neolamprologus pulcher. We experimentally created 34 breeding vacancies and investigated the effects of breeder removal on the degree of cooperative behaviour and how changes in status influenced reproductive physiology. We found that 71% of male vacancies were filled by joiners (incoming fish, not previously members of the group) entering the territory. In contrast, only 15% of female vacancies were filled by joiners entering the group from elsewhere. Helpers increased their frequency of cooperative behaviour following the removal of a female breeder, but not following removal of a male breeder. Our results suggest that female breeder vacancies are typically filled by subordinate helpers who inherit from within the group and that male breeder vacancies are commonly filled by joining individuals (existing breeders or former helpers from other groups). Male social status and gonadal investment were positively correlated. This study represents one of the first experimental attempts to examine sexual differences in the pathway to breeding status in a cooperatively breeding species.

Mate Choice in Male Mandrills (Mandrillus sphinx)

AbstractMale primates that attempt to monopolize access to receptive females by mate‐guarding expend time and energy and risk injury, making reproduction costly. Males should therefore show mate choice and preferentially allocate mating effort to females that are likely to be fertile and those that will produce high‐quality offspring. Specifically, males should preferentially mate‐guard high‐ranking females rather than low‐ranking females, as such females are more likely to be fertile and are able to invest more in offspring. Males should also prefer parous females to nullipares, for similar reasons. Finally, males should avoid mating with close relatives, to avoid the deleterious effects of inbreeding. We investigated 13 group‐years of mate‐guarding observations for two semi‐free‐ranging groups of mandrills to examine the influence of these factors on male investment in mate‐guarding. We found that males mate‐guarded higher‐ranking females more than lower‐ranking females, and parous females more than nullipares. Female age, true relatedness and maternal kinship did not influence male mate‐guarding. Our results suggest that male mandrills do exercise mate choice for higher‐quality females, in the form of higher‐ranking and parous females. As alpha males are responsible for the great majority of mate‐guarding, this can lead to assortative mating, where high‐ranking males reproduce with high‐ranking females, and has important implications for social relationships and kin selection.

Regulation of immunocompetence by different androgen metabolites in a blenny with alternative reproductive tactics

In Parablennius parvicornis, small reproductive males with relatively low expression of secondary sexual characters (M- morphotype) parasite on the parental investment of the larger nest-holder males which have fully developed secondary sexual characters (M+ morphotype). In comparison with M+ males, M- males have relatively low levels of androgens while having high blood cell percentages of lymphocytes and antigen responsiveness. Here we test the hypothesis that androgens are a causal factor for these differences in immunocompetence between morphotypes. After drawing an initial blood sample, males received a silastic implant containing either oil only (C), or oil with testosterone (T) or 11-ketotestosterone (KT). Males were re-caught 2 weeks later for drawing of the final blood sample. KT but not T induced the development of secondary sexual characters in M- males. M- males treated with KT showed lower swimming activity than the males treated with T or C implants, suggesting that KT also mediates behavioral changes in M- males. As expected, blood cell percentages of lymphocytes, but not of granulocytes, were higher in M- males than in M+ males. Overall, lymphocyte percentages increased in the C group which might have been a response to the surgery/treatment. In concordance with the hypothesis, lymphocyte percentages were suppressed in males treated with T in comparison with controls. However, no significant change was found in KT-treated males. This suggests that androgens modulate central, morphological and immunological traits by partly independent androgen mechanisms in P. parvicornis.

COMPARATIVE PHYLOGENETIC ANALYSIS OF MALE ALTERNATIVE REPRODUCTIVE TACTICS IN RAY-FINNED FISHES

Using comparative phylogenetic analysis, we analyzed the evolution of male alternative reproductive tactics (MARTs) in ray-finned fishes (Actinopterygii). Numerous independent origins for each type of MART (involving sneaker males, female mimics, pirates, and satellite males) indicate that these behaviors have been highly labile across actinopterygiian evolution, consistent with a previous notion that convergent selection in fishes can readily mold the underlying suites of reproductive hormones into similar behaviors. The evolutionary appearance of MARTs was significantly correlated with the presence of sexually selected traits in bourgeois males (P = 0.001) but not with the presence of male parental care. This suggests that MARTs often arise from selection on some males to circumvent bourgeois male investment in mate monopolization, rather than to avoid male brood care per se. We found parsimony evidence for an evolutionary progression of MARTs wherein sneaking is usually the evolutionary precursor to the presumably more complex MARTs of female mimicry and cooperative satellite behavior. Nest piracy appears not to be part of this evolutionary progression, possibly because its late onset in the life cycle of most ray-finned fishes reduces the effects of selection on this reproductive tactic.

Análisis ecológico de la flora de dos parcelas pirenaicas de seguimiento de los ecosistemas forestales.

Plants with hermaphrodite flowers show a trade-off in resource allocation to male and female functions of reproduction. Such a balance can be conditioned by several factors. One of them seems to be pollination vector. Thalictrum is a genus with species that use different ways of pollination. So that we have investigated the resource allocation patterns to each function (male and female) among five species with different pollination vectors. Allocation has been mesured as dry weight of the different flower structures. We have found a significative trend, male function relative investment is bigger in anemophilous than entomophilous among the flve studied species: Th. macrocarpum (wind pollinated), Th. Minus (zoind pollinated), Th. Aquilegiifolium (insect pollinated), Th. tuberosum (insect pollinated) and Th. flavum (insect pollinated). During bloom, the androecium has the the greater biomass in these five species, but when all the reproductive process is taken in account, fruits reach higher values. Furthermore, this temporal separation of the maximum allocation to each function would be a way to make competence for the same limited resources weaker.

Terminal Investment Strategies and Male Mate choice: Extreme Tests of Bateman

Bateman's principle predicts the intensity of sexual selection depends on rates of increase of fecundity with mating success for each sex (Bateman slopes). The sex with the steeper increase (usually males) is under more intense sexual selection and is expected to compete for access to the sex under less intense sexual selection (usually females). Under Bateman and modern refinements of his ideas, differences in parental investment are key to defining Bateman slopes and thus sex roles. Other theories predict sex differences in mating investment, or any expenditures that reduce male potential reproductive rate, can also control sex roles. We focus on sexual behaviour in systems where males have low paternal investment but frequently mate only once in their lifetimes, after which they are often killed by the female. Mating effort (=terminal investment) is high for these males, and many forms of investment theory might predict sex role reversal. We find no qualitative evidence for sex role reversal in a sample of spiders that show this extreme male investment pattern. We also present new data for terminally-investing redback spiders (Latrodectus hasselti). Bateman slopes are relatively steep for male redbacks, and, as predicted by Bateman, there is little evidence for role reversal. Instead, males are competitive and show limited choosiness despite wide variation in female reproductive value. This study supports the proposal that high male mating investment coupled with low parental investment may predispose males to choosiness but will not lead to role reversal. We support the utility of using Bateman slopes to predict sex roles, even in systems with extreme male mating investment.

Hormonal induction of undescribed males resolves cryptic species of cladocerans

Cyclic parthenogens have a mixed breeding system with both meiotic and ameiotic eggs. Although investment in sexual stages is often synchronized with seasonal cycles, the degree of investment is a quantitative trait associated with habitat instability. Populations of cyclic parthenogens from stable environments, such as large lakes and oceans, generally show reduced or undetectable investment in males. Indeed, males of many species of lacustrine cyclic parthenogens are unknown to science. Methyl farnesoate (MF), a crustacean juvenile hormone, has been implicated as an inducer of male formation in Daphnia magna (Crustacea: Cladocera), a denizen of unstable habitats with marked sexual recruitment. Here, we show experimentally that MF induces male production in four distantly related lacustrine species of cladocerans under growth conditions unfavourable for male production. The males of three species are new to science. Unlike females, the anatomy of the previously unknown males of Bosmina (Lunobosmina) oriens permitted ready morphological diagnosis of sibling species and subfossils. The results suggest that the role for MF in the sex determination of cladocerans is general.

Chance, Time Allocation, and The Evolution of Adaptively Flexible Sex Role Behavior

An alternative to classic sexual selection hypotheses for sex differentiated pre-mating behavior is that time available for mating-as individuals experience it-along with fitness differences among alternative potential mates, induces choosy versus indiscriminate mating behavior. This alternative hypothesis says that selection has acted so that all individuals flexibly express fitness-enhancing choosy, indiscriminate, and competitive mating behavior, induced by time-varying life histories, environmental and social cues. Key predictions of DYNAMATE, the formal model of adaptively flexible sex role behavior of individuals of both sexes within dynamically changing populations, include: (1) All individuals regardless of sex assess likely fitness outcomes from mating with alternative potential mates before expressing choosy or indiscriminate behavior. (2) Males and females express adaptively flexible, choosy and indiscriminate behavior so that individuals may change their behavior-from moment to moment-to fit dynamically changing circumstances. (3) Indiscriminate behavior of males and (4) choosy behavior of females would often be maladaptive even in species with greater female than male parental investment, when females have longer latencies to receptivity to re-mating than males, and when the relative reproductive rate of males is greater than in females. (5) Whether or not females show choosy behavior will not affect whether or not males exhibit choosy or indiscriminate behavior, and vice versa. (6) When other model parameters are equal, the proportion of individuals of a given sex expressing choosy or indiscriminate mating behavior is a function of the distribution of fitness ratios (a distribution of all fitness differences that would be conferred on an individual by mating with any two sequentially or simultaneously encountered alternative potential mates). (7) Whether same-sex individuals behaviorally compete is a function of the fitness that would be conferred if the strategist won access to a potential mate, but not a function of relative reproductive rate or its proxy, the operational sex ratio. We call for re-evaluation of sex differences in choosy, indiscriminate, and competitive behavior under strong experimental controls that level the ecological playing fields of males and females, i.e., under experimental conditions informing the mechanisms of phenotypic expression. We end with comments on the classic question of questions: why are the sexes as they are?

Do a male garter snake's energy stores limit his reproductive effort?

The storage and utilization of energy plays a critical role in reproductive output for females of many species, including snakes. However, links between energy and reproductive effort in males are less clear. Studies at a communal den of red-sided garter snakes (Thamnophis sirtalis parietalis (Say, 1823)) in Manitoba suggest that energy is critical to reproduction for males as well as females. Males vary substantially in body condition (mass relative to body length) at the time they emerge from winter inactivity. The energy to be expended in courtship is stored in the muscles rather than the "conventional" sites for energy storage in snakes (abdominal fat bodies or liver). A male's reproductive effort (the duration of his residency at courting aggregations near the den) was linked to his energy stores and to the rate of depletion of those stores. Male snakes that emerged from hibernation in better condition, and that lost mass slowly thereafter, remained in courting aggregations near the den for longer periods than did males that emerged in poor condition (i.e., with less energy resources) and (or) lost mass more rapidly. In outdoor arenas, males that engaged in courtship lost mass more rapidly than did males with no courtship opportunities. These data suggest that courtship is energetically expensive for male garter snakes and that the amount of effort that a male invests in reproduction is determined by his energy stores.

Outcrossing hermaphroditic polychaete worms adjust their sex allocation to social conditions

Sex allocation theory predicts that simultaneous hermaphrodites shift sex allocation facultatively in response to variation in local group size. This study was performed to evaluate the relative investment in each sex function by the simultaneously hermaphroditic polychaete worm Ophryotrocha diadema and to test whether allocation to each sex depends on the number of reproductive competitors. Four experimental groups were set up (in a 2 x 2 factorial design) with small or large group size and with small or large enclosures to control for potential confounding effects of density. We measured the proportion of female and male investment in focal individuals. Results revealed that individuals regulated their reproductive output so that when reproductive competitors were present, the number of female gametes was strongly reduced and the male function increased. In contrast, under monogamy, individuals in small groups produced lower numbers of sperm but had a higher egg output than worms in large groups. Density did not affect sex allocation in our experiment. Our findings provide qualitative support for Local Mate Competition theory, but also show that the pattern of sex allocation specific to this species is more complex than expected by current theory.

Survival and reproduction when food is scarce: implications for a lekking Hawaiian Drosophila

Abstract. 1. In insects, larval diet can have a major impact on development, survival, and reproductive success. However, resource availability at the adult phase of the life cycle is also likely to have strong effects in species where there is an extended period of sexual maturation following adult eclosion.2. The effect of diet on the survival and reproductive success of the lekking Hawaiian fruit fly, Drosophila grimshawi, was explored. Two generations of emerging adults were exposed to one of two feeding regimes: ‘constant’ and ‘varied’ (corresponding to food ‘each day’ or ‘every other day’ respectively). The impact of resource availability on survival and reproductive success in each generation was then investigated.3. The probability of survival to 5 weeks old was higher for individuals fed a constant diet than individuals fed a varied diet, but was comparable for males and females.4. There was a significant maternal effect on offspring survival. Offspring whose mothers were reared on a constant feeding regime had higher survival than offspring whose mothers were reared on a varied diet.5. There was no relationship between feeding regime and the quantity of pheromones deposited by males (a measure of male reproductive investment); however F2 sons were more likely to deposit pheromones and deposited a larger quantity of pheromone than their F1 sires. The number and sex ratio of offspring (a measure of female reproductive effort) emerging from the F1 generation was unrelated to maternal or paternal feeding regime.6. The implications of variation in the foraging environment for mate choice in D. grimshawi are discussed.

Intra-sexual variation in male reproduction in teleost fish: a comparative approach

The occurrence of intra-sexual variation in reproduction is a widespread phenomenon in teleosts. One such form of variation consists in the occurrence of alternative male types: males that invest resources in mate attraction and males that exploit the investment of the former males, by trying to sneak fertilizations during spawning. These alternative reproductive tactics can be classified according to their plasticity during the life span of the individuals (i.e., fixed vs. sequential vs. reversible). Furthermore, the differences between morphs within a given species may involve a set of different traits, including reproductive behavior, the differentiation of male morphological traits, and the patterns of gonad tissue allocation and the differentiation of gonadal accessory glands. In this paper, we review the available data on four species exhibiting different types of intra-sexual plasticity in reproduction that have been studied in our lab. The data on the proximate mechanisms, androgens and forebrain arginine-vasotocin (AVT), underlying these alternative tactics suggest that between-morph differences in androgen levels, especially in 11-ketotestosterone, are especially present in species where the alternative male types have evolved morphological traits that are tactic-specific (i.e., sexual polymorphisms) and that differences in AVT appear to be related to between-morph differences in the expression of courtship behavior. Therefore, this comparative approach leads us to propose that the different endocrine systems are involved in the differentiation of different sets of traits that make up alternative phenotypes, and that the differentiation of alternative tactics is not controlled by a single endocrine system (e.g., androgens).

Nocturnal Breeding Behavior and Related Parental Investment of the Black Tern

Abstract Nocturnal incubating and brooding behavior of the Black Tern (Chlidonias niger) was studied at two colonies in successive years in The Netherlands. In late evening and early morning, nests were checked regularly for female or male nest attendance until or from complete darkness. During daytime females incubated eggs 60% of the time. At night, males took over completely, while females spent the night at a communal roost more than three km away from the breeding colonies. Few females stayed with their downy young at night and more females brooded young, as chicks became older. The total parental breeding investment (day and night) was equal (51% for males), but with higher nocturnal investment by males (79%). From six days after hatching, chicks were left unattended at night. The difference between male and female nocturnal investment might be explained by the preference for self-survival instead of clutch and brood survival. Males probably have greater interest in clutch survival than females, as ...

A Cross-Cultural Investigation of the Role of Foot Size in Physical Attractiveness

Disparate cultural practices suggest that small foot size may contribute to female attractiveness. Two hypotheses potentially explain such a pattern. Sexual dimorphism in foot size may lead observers to view small feet as feminine and large feet as masculine. Alternately, because small female feet index both youth and nulliparity, evolution may have favored a male preference for this attribute in order to maximize returns on male reproductive investment. Whereas the observational hypothesis predicts symmetrical polarizing preferences, with small feet being preferred in women and large feet being preferred in men, the evolutionary hypothesis predicts asymmetrical preferences, with the average phenotype being preferred in men. Using line drawings that varied only in regard to relative foot size, we examined judgments of attractiveness in nine cultures. Small foot size was generally preferred for females, while average foot size was preferred for males. These results provide preliminary support for the hypothesis that humans possess an evolved preference for small feet in females.

Mass loss during the mating season in male common brushtail possums ( Trichosurus vulpecula ): implications for age‐specific variation in mating strategy

Little is known of patterns of male reproductive effort and investment, and their relationship with age-related differences in male mating strategy, in mammals. In this study, the age-specific patterns of reproductive expenditure in a male marsupial, the common brushtail possum Trichosurus vulpecula were investigated. Possums were studied for two breeding seasons on Magnetic Island, north Queensland, Australia. Energy expenditure in male possums during the mating season was estimated from changes in mass taken monthly during live-trapping. Male possums lost an average of 4.2% of their mass during the main mating season from January to May, suggesting that mating and consort behaviour is energetically expensive and may constrain foraging activity. The percentage of body mass lost by individual males during the breeding season was negatively related to age; older males directed less somatic investment into reproduction than young males. Older males seemed to target their energy expenditure toward the period when oestrus synchrony was highest. The pattern of age-specific energy expenditure for reproduction in T. vulpecula differs markedly from that reported previously for male cervids, probably owing to the spatial dispersal of females leading to differences in mating strategy. The results are discussed in terms of their relevance to potential age-related differences in reproductive strategy and behaviour.

Male house sparrows deliver more food to experimentally subsidized offspring

We explored behavioural adjustments made by parent house sparrows, Passer domesticus, when the nutritional condition of their dependent nestlings was improved experimentally. Male parents responded to artificially supplemented broods by increasing food deliveries, whereas female parents continued matching the already high rate of control females. Thus, parental care was not truncated in the face of fortified offspring, but actually escalated (ca. 17% more adult food deliveries overall). Supplemented nestlings showed a nonsignificant tendency to recruit into the adult breeding population more than controls. We propose that one important reason why male parents responded more strongly than their female partners centres on the lower marginal costs for additional male posthatching investment, specifically by demonstrating that increasing paternal investment is likely to confer higher fitness than alternative male activities.

Male bill colour and age are associated with parental abilities and breeding performance in blackbirds

In monogamous bird species, male parental investment may influence offspring fitness and females may gain advantages through mating with males providing extensive paternal care. However, paternal care is a benefit that can only be assessed indirectly because mate choice precedes paternal activities. Individual quality and age, both signalled by morphological characteristics, may reflect parental abilities. Because they may reflect individual foraging abilities, carotenoid-based colorations have been proposed to honestly signal parental quality. The blackbird (Turdus merula), a socially monogamous species, exhibits biparental care and males show bills that vary from pale yellow to orange due to carotenoid pigments. In this study, we investigated whether male bill colour and age are associated with parental ability. Our results suggest that males with more orange bills and older males are better fathers. Indeed, male visit rate increased with their bill colour index independently of age, and brood condition was higher for adult males, compared to yearlings, independently of bill colour. Overall, the number of fledglings produced was positively influenced by both the age of males and the colour intensity of their bills. Males with more orange bills and adults had a greater number of fledglings and these males also had higher levels of prolactin, a hormone known to promote parental care. This latter finding suggests that prolactin may be the link between carotenoid based colorations and the intensity of paternal effort. Thus, male bill colour seems to honestly reveal male physiological adjustment to paternal activities.