Changes In Sex Allocation Research Articles

Male reproductive success is not strongly affected by phenological changes in mate availability in monoecious Sagittaria latifolia.

Many plants express their female and male sex roles at different times (dichogamy), with important consequences for mating. Dichogamy can yield mate limitation via biased floral sex ratios, particularly at the beginning and end of the flowering season when many plants simultaneously function as the same sex. This form of mate limitation should be reduced if plants adjust their allocations to female versus male sex functions in a manner that tracks seasonal variability in mating opportunities. For example, under protogyny (i.e. dichogamy with female function expressed first) plants with male-biased sex expression should have enhanced mating opportunities early in the flowering season as other plants begin to flower (in female sex phase). We quantified seasonal changes in sex allocation, patterns of mate availability and realized siring success in a population of protogynous Sagittaria latifolia. Our results were consistent with previous findings that seasonal changes in sex allocation should compensate for lost mating opportunities under the temporally variable mating environments caused by dichogamy. However, patterns of siring success in the population were inconsistent with this interpretation. We suggest that realized siring success might depend more strongly on spatial than on temporal aspects of mate availability.

Egg provisioning explains the penetrance of symbiont-mediated sex allocation distortion in haplodiploids.

Maternally transmitted symbionts such as Wolbachia can alter sex allocation in haplodiploid arthropods. By biasing population sex ratios towards females, these changes in sex allocation may facilitate the spread of symbionts. In contrast to symbiont-induced cytoplasmic incompatibility (CI), the mechanisms that underpin sex allocation distortion remain poorly understood. Using a nuclear genotype reference panel of the haplodiploid mite Tetranychus urticae and a single Wolbachia variant that is able to simultaneously induce sex allocation distortion and CI, we unraveled the mechanistic basis of Wolbachia-mediated sex allocation distortion. Host genotype was an important determinant for the strength of sex allocation distortion. We further show that sex allocation distortion by Wolbachia in haplodiploid mites is driven by increasing egg size, hereby promoting egg fertilization. This change in reproductive physiology was also coupled to increased male and female adult size. Our results echo previous work on Cardinium symbionts, suggesting that sex allocation distortion by regulating host investment in egg size is a common strategy among symbionts that infect haplodiploids. To better understand the relevance that sex allocation distortion may have for the spread of Wolbachia in natural haplodiploid populations, we parametrized a model based on generated phenotypic data. Our simulations show that empirically derived levels of sex allocation distortion can be sufficient to remove invasion thresholds, allowing CI to drive the spread of Wolbachia independently of the initial infection frequency. Our findings help elucidate the mechanisms that underlie the widespread occurrence of symbionts in haplodiploid arthropods and the evolution of sex allocation.

Male-biased sex allocation in ageing parents; a longitudinal study in a long-lived seabird.

Optimal sex allocation is frequency-dependent, but senescence may cause behaviour at old age to be suboptimal. We investigated whether sex allocation changes with parental age, using 16 years of data comprising more than 2500 molecularly sexed offspring of more than 600 known-age parents in common terns (Sterna hirundo), slightly sexually size-dimorphic seabirds. We decomposed parental age effects into within-individual change and sex allocation-associated selective (dis)appearance. Individual parents did not differ consistently in sex allocation, but offspring sex ratios at fledging changed from female- to male-biased as parents aged. Sex ratios at hatching were not related to parental age, suggesting sons to outperform daughters after hatching in broods of old parents. Our results call for the integration of sex allocation theory with theory on ageing and demography, as a change in sex allocation with age per se will cause the age structure of a population to affect the frequency-dependent benefits and the age-specific strength of selection on sex allocation.

Influence of temperature on patch residence time in parasitoids: physiological and behavioural mechanisms.

Patch time allocation has received much attention in the context of optimal foraging theory, including the effect of environmental variables. We investigated the direct role of temperature on patch time allocation by parasitoids through physiological and behavioural mechanisms and its indirect role via changes in sex allocation and behavioural defences of the hosts. We compared the influence of foraging temperature on patch residence time between an egg parasitoid, Trichogramma euproctidis, and an aphid parasitoid, Aphidius ervi. The latter attacks hosts that are able to actively defend themselves, and may thus indirectly influence patch time allocation of the parasitoid. Patch residence time decreased with an increase in temperature in both species. The increased activity levels with warming, as evidenced by the increase in walking speed, partially explained these variations, but other mechanisms were involved. In T. euproctidis, the ability to externally discriminate parasitised hosts decreased at low temperature, resulting in a longer patch residence time. Changes in sex allocation with temperature did not explain changes in patch time allocation in this species. For A. ervi, we observed that aphids frequently escaped at intermediate temperature and defended themselves aggressively at high temperature, but displayed few defence mechanisms at low temperature. These defensive behaviours resulted in a decreased patch residence time for the parasitoid and partly explained the fact that A. ervi remained for a shorter time at the intermediate and high temperatures than at the lowest temperature. Our results suggest that global warming may affect host-parasitoid interactions through complex mechanisms including both direct and indirect effects on parasitoid patch time allocation.

Oviposition but Not Sex Allocation Is Associated with Transcriptomic Changes in Females of the Parasitoid Wasp Nasonia vitripennis.

Linking the evolution of the phenotype to the underlying genotype is a key aim of evolutionary genetics and is crucial to our understanding of how natural selection shapes a trait. Here, we consider the genetic basis of sex allocation behavior in the parasitoid wasp Nasonia vitripennis using a transcriptomics approach. Females allocate offspring sex in line with the local mate competition (LMC) theory. Female-biased sex ratios are produced when one or a few females lay eggs on a patch. As the number of females contributing offspring to a patch increases, less female-biased sex ratios are favored. We contrasted the transcriptomic responses of females as they oviposit under conditions known to influence sex allocation: foundress number (a social cue) and the state of the host (parasitized or not). We found that when females encounter other females on a patch or assess host quality with their ovipositors, the resulting changes in sex allocation is not associated with significant changes in whole-body gene expression. We also found that the gene expression changes produced by females as they facultatively allocate sex in response to a host cue and a social cue are very closely correlated. We expanded the list of candidate genes associated with oviposition behavior in Nasonia, some of which may be involved in fundamental processes underlying the ability to facultatively allocate sex, including sperm storage and utilization.

Variable flight distance to resources results in changing sex allocation decisions, Megachile rotundata

Resource availability and location are expected to vary, based on use by individuals and changing ecological conditions; the response of animals to those changes is critical to their fitness. To investigate the effects of changing conditions on fitness, we conducted field experiments using the central-place-foraging alfalfa leafcutting bee (Megachile rotundata), examining the changes in sex allocation when presented with changes in flight distance required to obtain resources. The results suggest that changes in flight distance to resources during the year’s flight season altered subsequent allocation decisions: mothers experiencing long flight distance to resources early in the season and short flight distance later in the season generated a greater proportion of female offspring than mothers experiencing the opposite. During the second half of the season, however, it was current experimental conditions significantly impacting decisions with those mothers residing near the resources producing a greater portion of female offspring than those far from resources, regardless of flight distances experienced earlier in the year. These results show that sex allocation decisions may change dynamically during a female’s lifetime in response to ecological changes. We also looked at these results through the contradicting hypotheses marginal value theorem and sex ratio theory, finding that the latter may be in play depending on an individual’s past and/or present condition.

Within-plant variation in reproductive investment: consequences for selection on flowering time.

Variation among the leaves, flowers or fruit produced by a plant is often regarded as a nuisance to the experimenter and an impediment to selection. Here, we suggest that within-plant variation can drive selection on other plant-level traits. We examine within-plant variation in floral sex allocation and in fruit set and predict that such variation generates variation in male success among plants, thereby driving selection on flowering time. We tested this prediction in a simulation model estimating selection on flowering time through male fitness when floral sex allocation and/or fruit set vary directionally among flowers on plants. We parameterized the model through a quantitative literature survey of within-plant change in sex allocation. As predicted, within-plant variation in floral sex allocation and in fruit set probability can generate selection on flowering time through male fitness. Declining fruit set from first to last flowers on plants, as occurs in many species, selected for early flowering onset through male fitness. This result was robust to self-incompatibility and to varying returns on male versus female investment. Selection caused by declining fruit set was strong enough to reverse the selection for late flowering that can be caused by intrafloral protandry. Our model provides testable predictions regarding selection on flowering time through male fitness. The model also establishes the intriguing possibility that within-plant variation may influence selection on other traits, regardless of whether that variation is under selection itself.

Sex ratio variations with temperature in an egg parasitoid: behavioural adjustment and physiological constraint

Sex allocation in haplodiploid arthropods is a central fitness-related decision that has received much attention in insect parasitoids. The effect of temperature on the reproductive strategy of female parasitoids has rarely been addressed, despite evidence of its influence on other fitness-related traits. We explored mechanisms inducing the higher production of males typically observed at low and high temperature in parasitic wasps, considering that this pattern may result from both behavioural adjustment and physiological constraint. By observing the oviposition behaviour of an egg parasitoid, Trichogramma euproctidis , we were able to distinguish the sex ratio intended by the female from the secondary sex ratio, thereby discriminating between a change in behaviour and a physiological constraint on egg fertilization. More males emerged from eggs laid at low (+45%) or high (+80%) temperature than at medium temperature, but the underlying mechanisms differed between the two conditions. We observed a behavioural change in sex allocation at high temperature, suggesting that laying of sons may be advantageous at high temperature. At low temperature, the females' intended sex ratio was similar to that at medium temperature, but physiological constraints prevented egg fertilization during oviposition, resulting in more males emerging from eggs intended to be females. To our knowledge this is the first experimental evidence that temperature modulates both sex allocation and physiological constraints in egg fertilization in parasitoids. • We explored mechanisms inducing higher production of male parasitoids at low and high temperature. • We compared intended vs secondary sex ratios to discriminate between behavioural change and physiological constraint. • More males emerged at low and high temperature than at intermediary temperature, but the mechanisms differed. • We observed a behavioural change in sex allocation at high temperature. • We observed that physiological constraints prevented egg fertilization at low temperature.

Nectar secretion strategy in three Japanese species: changes in nectar volume and sugar concentration dependent on flower age and flowering order

We explored how changes in nectar volume and nectar sugar concentration depend on flower age and flowering order in Gentiana triflora Pallas var. japonica (Kusnez.) Hara, Lobelia sessilifolia Lamb., and Hemerocallis middendorffii Trautv. et Mey. var. esculenta (Koidz.) Ohwi. In G. triflora var. japonica and H. middendorffii var. esculenta, change in nectar volume was small, whereas larger changes in nectar sugar concentration occurred depending on flower age. In L. sessilifolia, both the mean nectar volume and nectar sugar concentration clearly decreased with flower age. In all species, the mean nectar volume of the early-flowering group per plant was high. In terms of the change in sex allocation, the investment in male and female organs of the early-flowering group was high in G. triflora var. japonica and L. sessilifolia. We suggest that plants of G. triflora var. japonica and H. middendorffii var. esculenta secrete floral nectar for up to several days to sustain nectar volume to keep attracting pollinators while simultaneously reducing resource usage. Greater nectar secretion in the early-flowering group, in which flowers have more pollen and ovules, may contribute to greater reproductive success and may be effective for pollinators in learning the location of the plants and flowers. In addition, learning by pollinators should result in increased reproductive success of the later blooming flowers.

Temporal variation in the pollen:ovule ratios of Clarkia (Onagraceae) taxa with contrasting mating systems: field populations

Among plants, pairs of selfing vs. outcrossing sister taxa provide interesting systems in which to test predictions concerning the magnitude and direction of temporal changes in sex allocation. Although resource availability typically declines towards the end of the growing season for annual taxa, temporal changes in mating opportunities depend on mating system and should change less in selfing taxa. Consequently, given that the pollen:ovule (P:O) ratio of flowers reflects the investment in (and potential fitness pay-off due to) male vs. female function, we predicted that the P:O ratio should also be less variable among and within selfers than in closely related outcrossers. To test these predictions, we measured temporal changes in sex allocation in multiple field populations of two pairs of sister taxa in the annual flowering plant genus Clarkia (Onagraceae). In the outcrossing Clarkia unguiculata and the selfing Clarkia exilis, ovule production declined similarly from early to late buds, whereas pollen production remained constant or increased in the outcrosser but remained constant or decreased in the selfer. Consequently, the P:O ratio increased within unguiculata populations but marginally increased or stayed constant in exilis populations. In all populations of the selfing Clarkia xantiana spp. parviflora and the outcrossing C. x. spp. xantiana, both ovule and pollen production per flower declined over time. The effects of these declines on the P:O ratio, however, differed between subspecies. In each xantiana population, the mean P:O ratio did not differ between early and late flowers, although individuals varied greatly in the direction and magnitude of phenotypic change. By contrast, parviflora populations differed in the mean direction of temporal change in the P:O ratio. We found little evidence to support our initial predictions that the P:O ratio of the selfing taxa will consistently vary less than in outcrossing taxa.

Change in movement and subdivision of Myrmica punctiventris (Hymenoptera, Formicidae) colonies in north temperate forests is related to a long-term shift in social organization

We assessed temporal and spatial patterns of nest site use in the cavity dwelling ant, Myrmica punctiventris in a well-studied temperate forest site in central New York State. We evaluated changes in nest site use by repeatedly censusing the ants inhabiting artificial nests (hollow dowels) for three consecutive growing seasons. We confirm a shift towards more polydomy in this population of M. punctiventris, first reported by DeHeer et al. (2001), but extend these findings by demonstrating that this shift has occurred gradually over several years. We are unable to determine if this polydomy is seasonal or year round. We explore various explanations for the occurrence of polydomy and suggest that long-term changes in sex allocation are indirectly driving the shift in nesting strategies.

Wind pollination, sexual dimorphism, and changes in floral traits of Schiedea (Caryophyllaceae)

Both changes in sex allocation and pollination mode may promote the separation of sexes in plant populations. Simultaneous evolution of wind pollination and dimorphism has occurred in Schiedea, where species with different female frequencies provide an opportunity to observe the effect of wind pollination on sex allocation and floral morphology. Differences among species in the ratio of anther to ovary volume were not the result of sex allocation trade-offs, but instead resulted from production of vestigial stamens in females; there were no changes in ovary volume in males and hermaphrodites (MH) of dimorphic species. Relative to hermaphroditic species, dimorphic species had more condensed inflorescences, a pattern often associated with wind pollination. Within dimorphic species, MH had longer filament lengths than females, and females had longer stigmas than MHs. These traits are characteristic of wind pollination, but there was no relationship between the degree of sexual dimorphism and female frequency. Ovary volume and ovule number and size had positive phenotypic correlations between females and MH of dimorphic species, making sex specialization more difficult. In dimorphic Schiedea species, selection for wind pollination may have a greater effect on floral traits than trade-offs in allocation between male and female function.

Adaptive sex allocation in relation to life‐history in the common brushtail possum, Trichosurus vulpecula

Summary Maternal control of offspring sex remains a contentious topic in the evolutionary and behavioural ecology of vertebrates. Two models of adaptive sex allocation − the Trivers–Willard (TW) and local resource competition (LRC) hypotheses − have been supported in some studies, but in many other cases the forces shaping offspring sex ratios are unclear. One reason that past studies may not have provided clear support for either model is that the two processes interact, producing complex patterns of sex allocation in relation to the life histories of individual mothers. The TW hypothesis predicts that in dimorphic, polygynous mammals, females in good condition should be more likely to produce male offspring. The LRC hypothesis predicts an independent effect of age on the offspring sex ratio; females should be more likely to produce sons early in their reproductive lives and daughters later, in order to minimize the duration and reproductive cost of mother–daughter competition. We tested these two predictions in a study of sex allocation in the common brushtail possum, Trichosurus vulpecula. Offspring sex was strongly dependent on maternal age: females breeding for the first time were likely to produce sons, and at subsequent breeding attempts the sex ratio of offspring was slightly biased towards females. After the first breeding season, sex of offspring was influenced more strongly by body condition than by age, and females in good condition were more likely to have sons. Individual females that showed a between‐year gain in condition were also more likely to produce a male in the subsequent year. Body mass of male offspring at 5 months was positively correlated with maternal condition. No relationship was found between maternal condition and mass of female offspring. We conclude that patterns of change in sex allocation in relation to individual life histories of female common brushtail possums can be explained by the interaction of TW and LRC effects.

Adaptive production of fighter males: queens of the ant Cardiocondyla adjust the sex ratio under local mate competition.

Hamilton's concept of local mate competition (LMC) is the standard model to explain female-biased sex ratios in solitary Hymenoptera. In social Hymenoptera, however, LMC has remained controversial, mainly because manipulation of sex allocation by workers in response to relatedness asymmetries is an additional powerful mechanism of female bias. Furthermore, the predominant mating systems in the social insects are thought to make LMC unlikely. Nevertheless, several species exist in which dispersal of males is limited and mating occurs in the nest. Some of these species, such as the ant Cardiocondyla obscurior, have evolved dimorphic males, with one morph being specialized for dispersal and the other for fighting with nest-mate males over access to females. Such life history, combining sociality and alternative reproductive tactics in males, provides a unique opportunity to test the power of LMC as a selective force leading to female-biased sex ratios in social Hymenoptera. We show that, in concordance with LMC predictions, an experimental increase in queen number leads to a shift in sex allocation in favour of non-dispersing males, but does not influence the proportion of disperser males. Furthermore, we can assign this change in sex allocation at the colony level to the queens and rule out worker manipulation.