Plasticity in sex allocation in the plant Mercurialis annua is greater for hermaphrodites sampled from dimorphic than from monomorphic populations

Plants are notoriously variable in gender, ranging in sex allocation from purely male through hermaphrodite to purely female. This variation can have both a genetic and an adaptive plastic component. In gynodioecious species, where females co‐occur with hermaphrodites, hermaphrodites tend to shift their allocation towards greater maleness when growing under low‐resource conditions, either as a result of hermaphrodites shifting away from an expensive female function, or because of enhanced siring advantages in the presence of females. Similarly, in the androdioecious plant Mercurialis annua, where hermaphrodites co‐exist with males, hermaphrodites also tend to enhance their relative male allocation under low‐resource conditions. Here, we ask whether this response differs between hermaphrodites that have been evolving in the presence of males, in a situation analogous to that supposed for gynodioecious populations, vs. those that have been evolving in their absence. We grew hermaphrodites of M. annua from populations in which males were either present or absent under different levels of nutrient availability and compared their reaction norms. We found that, overall, hermaphrodites from populations with males tended to be more female than those from populations lacking males. Importantly, hermaphrodites' investment in pollen and seed production was more plastic when they came from populations with males than without them, reducing their pollen production at low resource availability and increasing their seed production at high resource availability. These results are consistent with the hypothesis that plasticity in sex allocation is enhanced in hermaphrodites that have likely been exposed to variation in mating opportunities due to fluctuations in the frequency of co‐occurring males.     

Genetic and environmental control of temporal and size-dependent sex allocation in a wind-pollinated plant

Sex allocation in hermaphrodites can be affected by spatial and temporal variation in resources, especially in plants where size-dependent gender modification is commonplace. The evolution of sex allocation will depend on the relative importance of genetic and environmental factors governing patterns of investment in female and male function. In wind-pollinated plants, theoretical models predict a positive relation between size and male investment because of the fitness advantages associated with more effective pollen dispersal. Theory also predicts that the timing and allocation to each sex function should depend on available resources. We grew maternal half-sibling families of annual, wind-pollinated, Ambrosia artemisiifolia in sun and shade treatments to investigate these predictions. There was significant genetic variation for female and male flower production in both sun and shade treatments. Size-dependent sex allocation occurred in the direction predicted by theory, with male flower production increasing more rapidly in larger plants. The timing of sex function also varied, with significant genetic variation for dichogamy within environments and plasticity of this trait between environments. Protandry was expressed more commonly in the sun and protogyny in the shade. The occurrence of dynamic sex allocation with changing size and experimental treatment indicates the potential for adaptive responses under different ecological conditions.     

Mating frequency and resource allocation to male and female function in the simultaneous hermaphrodite land snail Arianta arbustorum

Sex allocation theory predicts that mating frequency and long‐term sperm storage affect the relative allocation to male and female function in simultaneous hermaphrodites. We examined the effect of mating frequency on male and female reproductive output (number of sperm delivered and eggs deposited) and on the resources allocated to the male and female function (dry mass, nitrogen and carbon contents of spermatophores and eggs) in individuals of the simultaneous hermaphrodite land snail Arianta arbustorum. Similar numbers of sperm were delivered in successive copulations. Consequently, the total number of sperm transferred increased with increasing number of copulations. In contrast, the total number of eggs produced was not influenced by the number of copulations. Energy allocation to gamete production expressed as dry mass, nitrogen or carbon content was highly female‐biased (>95% in all estimates). With increasing number of copulations the relative nitrogen allocation to the male function increased from 1.7% (one copulation) to 4.7% (three copulations), but the overall reproductive allocation remained highly female‐biased. At the individual level, we did not find any trade‐off between male and female reproductive function. In contrast, there was a significant positive correlation between the resources allocated to the male and female function. Snails that delivered many sperm also produced a large number of eggs. This finding contradicts current theory of sex allocation in simultaneous hermaphrodites.     

Effective heritability of targets of sex-ratio selection under environmental sex determination

Selection is expected to maintain primary sex ratios at an evolutionary equilibrium. In organisms with temperature-dependent sex determination (TSD), targets of sex-ratio selection include the thermal sensitivity of the sex-determining pathway (hereafter, sex determination threshold) and nest-site choice. However, offspring sex may be canalized for nests located in thermally extreme environments; thus, genetic variance for the sex determination threshold is not expressed and is invisible to direct selection. The concept of 'effective heritability' accounts for this dependence and provides a more realistic prediction of the expected evolutionary response to selection in the wild. Past estimates of effective heritability of the sex determination threshold, which were derived from laboratory data, suggested that the potential for the sex determination threshold to evolve in the wild was extremely low. We re-evaluated original estimates of this parameter by analysing field-collected measures of nest temperatures, vegetation cover and clutch sex ratios from nests in a population of painted turtles (Chrysemys picta). We coupled these data with measurements of broad-sense heritability of the sex determination threshold in C. picta, using an experiment that splits clutches of eggs between a constant temperature (i.e. typical laboratory incubation) and a daily fluctuating temperature (i.e. similar to natural nests) with the same mean. We found that (i) the effective heritability of the sex determination threshold appears to have been historically underestimated and the effective heritability of nest-site choice has been overestimated and (ii) significant family-by-incubation treatment interaction exists for sex for C. picta between constant- and fluctuating-temperature regimes. Our results suggest that the thermal sensitivity of the sex-determining pathway may play a larger, more complex role in the microevolution of TSD than traditionally thought.     

The roles of female and hermaphroditic flowers in the gynodioecious–gynomonoecious Silene littorea: insights into the phenology of sex expression

Some gynodioecious species have intermediate individuals that bear both female and hermaphroditic flowers. This phenomenon is known as a gynodioecious–gynomonoecious sexual system. Gender expression in such species has received little attention in the past, and the phenologies of male and female functions have also yet to be explored. In this study, we examined variations in gender patterns, their effects on female reproductive success and sex expression in depth throughout the flowering period in two populations. The studied populations of Silene littorea contained mostly gynomonoecious plants and the number of pure females was very low. The gynomonoecious plants showed high variability in the total proportion of female flowers. In addition, the proportion of female flowers in each plant varied widely across the flowering season. Although there was a trend towards maleness, our measures of functional gender suggested that most plants transmit their genes via both pollen and ovules. Fruit set and seed set were not significantly different among populations; in contrast, flower production significantly varied between the two populations – and among plants – with consequent variation in total seed production. Conversely, gender and sex expression were similar in both populations. Plants with higher phenotypic femaleness did not have higher fruit set, seed set or total female fecundity. The mating environment fluctuated little across the flowering period, but fluctuations were higher in the population with low flower production. We therefore conclude that the high proportion of gynomonoecious individuals in our studied populations of S. littorea may be advantageous for the species, providing the benefits of both hermaphroditic and female flowers.     

Plasticity in resource allocation based life history traits in the Pacific oyster, Crassostrea gigas. I. Spatial variation in food abundance

We investigated the quantitative genetics of plasticity in resource allocation between survival, growth and reproductive effort in Crassostrea gigas when food abundance varies spatially. Resource allocation shifted from survival to growth and reproductive effort as food abundance increased. An optimality model suggests that this plastic shift may be adaptive. Reproductive effort plasticity and mean survival were highly heritable, whereas for growth, both mean and plasticity had low heritability. The genetic correlations between reproductive effort and both survival and growth were negative in poor treatments, suggesting trade-offs, but positive in rich ones. These sign reversals may reflect genetic variability in resource acquisition, which would only be expressed when food is abundant. Finally, we found positive genetic correlations between reproductive effort plasticity and both growth and survival means. The latter may reflect adaptation of C. gigas to differential sensitivity of fitness to survival, such that genetic variability in survival mean might support genetic variability in reproductive effort plasticity.     

The effects of fire on ant trophic assemblage and sex allocation

Fire plays a key role in ecosystem dynamics worldwide, altering energy flows and species community structure and composition. However, the functional mechanisms underlying these effects are not well understood. Many ground‐dwelling animal species can shelter themselves from exposure to heat and therefore rarely suffer direct mortality. However, fire‐induced alterations to the environment may change a species' relative trophic level within a food web and its mode of foraging. We assessed how fire could affect ant resource utilization at different scales in a Mediterranean forest. First, we conducted isotopic analyses on entire ant species assemblages and their potential food resources, which included plants and other arthropods, in burned and unburned plots 1 year postfire. Second, we measured the production of males and females by nests of a fire‐resilient species, Aphaenogaster gibbosa, and analyzed the differences in isotopic values among workers, males, and females to test whether fire constrained resource allocation. We found that, in spite of major modifications in biotic and abiotic conditions, fire had little impact on the relative trophic position of ant species. The studied assemblage was composed of species with a wide array of diets. They ranged from being mostly herbivorous to completely omnivorous, and a given species' trophic level was the same in burned and unburned plots. In A. gibbosa nests, sexuals had greater δ¹⁵N values than workers in both burned and unburned plots, which suggests that the former had a more protein‐rich diet than the latter. Fire also appeared to have a major effect on A. gibbosa sex allocation: The proportion of nests that produced male brood was greater on burned zones, as was the mean number of males produced per nest with the same reproductive investment . Our results show that generalist ants with relatively broad diets maintained a constant trophic position, even following a major disturbance like fire. However, the dramatically reduced production of females on burned zones compared to unburned zones 1 year postfire may result in considerably reduced recruitment of new colonies in the mid to long term, which could yield genetic bottlenecks and founder effects. Our study paves the way for future functional analyses of fire‐induced modifications in ant populations and communities.     

The evolution of dioecy, heterodichogamy, and labile sex expression in Acer

The northern hemisphere tree genus Acer comprises 124 species, most of them monoecious, but 13 dioecious. The monoecious species flower dichogamously, duodichogamously (male, female, male), or in some species heterodichogamously (two morphs that each produce male and female flowers but at reciprocal times). Dioecious species cannot engage in these temporal strategies. Using a phylogeny for 66 species and subspecies obtained from 6600 nucleotides of chloroplast introns, spacers, and a protein-coding gene, we address the hypothesis (Pannell and Verdú, Evolution 60: 660-673. 2006) that dioecy evolved from heterodichogamy. This hypothesis was based on phylogenetic analyses (Gleiser and Verdú, New Phytol. 165: 633-640. 2005) that included 29-39 species of Acer coded for five sexual strategies (duodichogamous monoecy, heterodichogamous androdioecy, heterodichogamous trioecy, dichogamous subdioecy, and dioecy) treated as ordered states or as a single continuous variable. When reviewing the basis for these scorings, we found errors that together with the small taxon sample, cast doubt on the earlier inferences. Based on published studies, we coded 56 species of Acer for four sexual strategies, dioecy, monoecy with dichogamous or duodichogamous flowering, monoecy with heterodichogamous flowering, or labile sex expression, in which individuals reverse their sex allocation depending on environment-phenotype interactions. Using Bayesian character mapping, we infer an average of 15 transformations, a third of them involving changes from monoecy-cum-duodichogamy to dioecy; less frequent were changes from this strategy to heterodichogamy; dioecy rarely reverts to other sexual systems. Contra the earlier inferences, we found no switches between heterodichogamy and dioecy. Unexpectedly, most of the species with labile sex expression are grouped together, suggesting that phenotypic plasticity in Acer may be a heritable sexual strategy. Because of the complex flowering phenologies, however, a concern remains that monoecy in Acer might not always be distinguishable from labile sex expression, which needs to be addressed by long-term monitoring of monoecious trees. The 13 dioecious species occur in phylogenetically disparate clades that date back to the Late Eocene and Oligocene, judging from a fossil-calibrated relaxed molecular clock.     

Expression of a life-history trade-off in a seed beetle depends on environmental context

Within populations of the seed beetle Callosobruchus maculatus (F.) (Coleoptera: Bruchidae), female longevity is positively correlated with fecundity, both phenotypically and genetically. Yet manipulations of egg-laying rate (through deprivation of seeds or mates) consistently suggest a trade-off between these traits. In this study, females were exposed to four levels of seed availability and the relationship between fecundity and longevity was examined both across and within environments. The expected trade-off was observed across environments, as females with few or no seeds laid fewer eggs and lived longer than females with many seeds. Within environments, however, the relationship was inconsistent; individual longevity and fecundity were positively correlated when seeds were abundant, uncorrelated when seeds were few and negatively correlated when seeds were absent. Body mass at adult emergence was positively correlated with fecundity, but its effect was stronger when seeds were present than when they were absent. After the effects of mass were statistically controlled, longevity and fecundity remained positively correlated among females with many seeds and negatively correlated among those without seeds. Thus, even a single technique (phenotypic correlation) can produce divergent conclusions about the ‘cost of reproduction’ in insects. The reversal in the correlation appears to depend on variation in reproductive allocation; when seeds are scarce or absent, differences in reproductive effort among individuals may be large enough to overwhelm differences in resource acquisition.     

Environmentally induced changes in carotenoid-based coloration of female lizards: a comment on Vercken et al

Colouration may either reflect a discrete polymorphism potentially related to life-history strategies, a continuous signal related to individual quality or a combination of both. Recently, Vercken et al. [J. Evol. Biol. (2007) 221] proposed three discrete ventral colour morphs in female common lizards, Lacerta vivipara, and suggested that they reflect alternative reproductive strategies. Here, we provide a quantitative assessment of the phenotypic distribution and determinants of the proposed colour polymorphism. Based on reflectance spectra, we found no evidence for three distinct visual colour classes, but observed continuous variation in colour from pale yellow to orange. Based on a 2-year experiment, we also provide evidence for reversible colour plasticity in response to a manipulation of the adult population sex ratio; yet, a significant portion of the colour variation was invariant throughout an adult female's life. Our results are thus in agreement with continuous colour variation in adults determined by environmental factors and potentially also by genetic factors.     

Adaptation and constraint in the evolution of environmental sex determination

When environments differentially influence male and female performance, environmental sex determination (ESD) might evolve. The conclusion from several previous theoretical models was that reaction norms for sex determination should have a single, sharp threshold, with only females being produced in some environments and only males in others. These reaction norms can be disadvantageous in fluctuating environments, however, because they lead to sex-ratio fluctuations. We analysed the evolution of ESD, looking for equilibrium strategies in unconstrained as well as constrained strategy spaces. We identified situations where a single-threshold reaction norm is not evolutionarily stable. In these cases, we found stable strategies in the form of complex reaction norms, showing an oscillatory pattern of sex determination with respect to variation in an environmental variable. Considering that constraints could prevent such phenotypes from being realized, we found that certain randomized reaction norms, with probabilistic sex determination for a range of environments, would achieve nearly the same fitness. We also investigated reaction norms constrained to have a single threshold and found that genetic polymorphism in the environmental threshold value could evolve, producing a similar effect as a randomized reaction norm. We argue that the appearance of genetic variation can be regarded as an alternative outcome when constraints prevent the evolution of a more complex or a randomized strategy.     

Variation in female life-history traits among Alaskan populations of the threespine stickleback, Gasterosteus aculeatus L. (Pisces: Gasterosteidae)

Life-history characteristics of female threespine stickleback (Gasterosteus aculeatus) were examined in 12 populations, 11 freshwater and one anadromous, within the Cook Inlet region of Alaska. Because this area has been deglaciated during the last 20 000 years, the freshwater populations are recently derived, probably independendy, from the local marine or anadromous stickleback. Freshwater threespine stickleback have undergone considerable morphological evolution within this region, apparently in response to environmental factors including predatory regimes and environmental productivity. Our freshwater study populations were selected to sample this range of morphological variation in order to determine whether life-history traits and morphologies have followed similar evolutionary trajectories. Freshwater populations could be categorized generally into one of three ecomorphotypes: those inhabiting relatively productive lakes having one or more piscivorous fishes present, and in which the stickleback exhibit a fully developed pelvic girdle; those inhabiting low-calcium lakes that lack piscivorous fishes, and in which the pelvic structures are incomplete; those living in streams with piscivorous fishes, in which the stickleback have fully developed pelvic girdles. The anadromous population constituted a fourth ecomorphotype that lives in marine waters, and is robusdy armored. The freshwater populations showed considerable variation in all life-history traits assessed, and this variation generally corresponded to our ecomorphological classifications. Nevertheless, within each ecomorphotype there was sufficient variation to suggest that morphological and life-history traits may not always respond in the same manner in response to the same selective regime.     

Male-biased sex allocation in late-blooming flowers driven by resource limitation in the clonal perennial Aconitum kusnezoffii (Ranunculaceae)

Flowering phenology and clonal growth are known to affect resource and pollen availability, and therefore select for adaptive or constrained sex allocation strategies to some degree. However, the consequences of temporal sex allocation patterns for reproductive fitness across the flower, inflorescence, and genet levels have rarely been examined. Moreover, experimental tests of the underlying regulatory mechanisms are scarce. We examined the association of flowering phenology and inflorescence position with temporal sex allocation and reproductive success in the protandrous perennial clonal herb, Aconitum kusnezoffii, over four consecutive growing seasons by examining more than 39 000 flowers. We also conducted controlled experiments to test the effects of resource and pollen limitation on the female reproductive success of lateral inflorescences. We found that some male functions were positively correlated with flowering phenology, whereas female reproductive success was negatively correlated with flowering phenology and inflorescence position. Lateral inflorescences invested more in male function than terminal inflorescences and therefore yielded fewer and smaller seeds. Resource limitation may serve as the key mechanism underlying this differentiated pattern. Decreased female reproductive success was consistently observed at the flower and inflorescence levels as flowering occurred later in the growth season. Late-blooming lateral inflorescences specialized in the male function, and their female reproductive success was constrained by early-blooming terminal inflorescences. This might be the first attempt to systematically demonstrate sex allocation strategy differentiation in a protandrous plant species at the inflorescence level. In addition, our study provides empirical evidence of dichogamy selecting for specialized sex allocation strategies among inflorescences.     

The effect of cryptic female choice on sex allocation in simultaneous hermaphrodites

Sex allocation theory for simultaneous hermaphrodites has focused primarily on the effects of sperm competition, but the role of mate choice has so far been neglected. We present a model to study the coevolution of cryptic female choice and sex allocation in simultaneous hermaphrodites. We show that the mechanism of cryptic female choice has a strong effect on the evolutionary outcome: if individuals remove a fixed proportion of less-preferred sperm, the optimal sex allocation is more female biased (i.e. more biased towards egg production) than without cryptic female choice; conversely, if a fixed amount of sperm is removed, sex allocation is less female-biased than without cryptic female choice, and can easily become male biased (i.e. biased towards sperm production). Under male-biased sex allocation, hermaphroditism can become unstable and the population can split into pure males and hermaphrodites with a female-biased allocation. We discuss the idea that the evolution of sex allocation may depend on the outcome of sexual conflict over the fate of received sperm: the sperm donor may attempt to manipulate or by-pass cryptic female choice and the sperm recipient is expected to resist such manipulation. We conclude that cryptic female choice can have a strong influence on sex allocation in simultaneous hermaphrodites and strongly encourage empirical work on this question.     

Exploring the evolution of environmental sex determination, especially in reptiles

Environmental sex determination has been documented in a variety of organisms for many decades and the adaptive significance of this unusual sex-determining mechanism has been clarified empirically in most cases. In contrast, temperature-dependent sex determination (TSD) in amniote vertebrates, first noted 40 years ago in a lizard, has defied a general satisfactory evolutionary explanation despite considerable research effort. After briefly reviewing relevant theory and prior empirical work, we draw attention to recent comparative analyses that illuminate the evolutionary history of TSD in amniote vertebrates and point to clear avenues for future research on this challenging topic. To that end, we then highlight the latest empirical findings in lizards and turtles, as well as promising experimental results from a model organism, that portend an exciting future of progress in finally elucidating the evolutionary cause(s) and significance of TSD.     

密度对尖头叶藜生物量分配格局及异速生长的影响

植物器官指示植物不同的功能,而植物器官生物量分配比例的变化表征了植物对资源获取能力的调整。在植物生长发育过程中,植物各器官呈一种明显的异速生长规律。利用异速生长分析方法,通过模拟不同密度(16、44.4、100、400株/m~2)下尖头叶藜(Chenopodium acuminatum)的生长特性,研究密度对尖头叶藜器官生物量分配格局及异速生长的影响。结果表明,随密度增加,尖头叶藜地上和地下器官都存在不同程度的竞争:其中,根和主茎生物量分配增加,茎和地上生物量分配减少,而叶和繁殖生物量分配不随密度变化而变化。研究发现,尖头叶藜各器官间具有显著的异速生长关系:其中叶∶主茎、根∶地上部分、根∶茎、根∶主茎、繁殖器官∶地上部分及繁殖器官∶根生物量间的异速生长不随密度变化而变化,属于表观可塑性;而叶∶地上部分、叶∶根、叶∶茎、茎∶地上部分、主茎∶地上部分、繁殖器官∶茎、繁殖器官∶主茎生物量间具有极显著的异速生长关系,异速指数和个体大小显著受密度变化影响,属于真正可塑性,这表明密度能够影响尖头叶藜各器官的生长变化。尖头叶藜叶∶主茎、叶∶根及主茎∶地上部分生物量间的异速指数在D4-密度时与3/4差异不显著(P0.05),符合生态代谢理论,而在D1—D3密度时与3/4差异显著(P0.05),表明充分竞争的植株更符合代谢理论,而竞争不激烈的植株对资源的投入具有物种特异性。     

An experimental study of the reproductive biology and hybridization in the European and North American species of Equisetum

Sporophytc production was investigated in the eleven North American and European species of Equisetum by means of fertilization experiments on both intact and subcultured gametophytes. Comparison of intra- and inter-gametophytic selfing with intergametophytic crossing failed to reveal any evidence of recessive sporophytic lethals, leaky lethals or deleterious recessive genes. Fertilization in Equisetum depends solely on the availability of ripe archegonia. Despite the absence of detectable genetic load in Equisetum , intergametophytic mating is ensured by gametophyte dioecism in the wild. All populations tested were freely interfcrtile indicating a high degree oi genetic con-specihcity, even between North America and Europe. Low sporophyte frequencies in European E. scirpoides are attributed to defective spermatozoids. Polyembryony occurs. Cessation in gametophyte development is mediated by allelopathic compounds liberated from the young sporophytes.
Reciprocal crosses resulted in the synthesis of 12 hybrids (six in subgenus Equisetum and six in subgenus Hippochaete plus abortive zygotes in the cross E. ramosissimum female x E. scirpoides male), 11 of which had previously been reported in nature. No intersubgeneric hybrids were produced, indicating that both subgenera are natural groups only distantly related to one another. The putative wild combinations E. pratense x E. atvense and E. variegatum x E. scirpoides were not synthesized, but a new hybrid from the allopatric E. ramosissimum and E. laevigatum arose. Reciprocal crosses within the subgenus Hippochaete usually yielded equal frequencies of hybrids, but in the subgenus Equisetum the majority of successful crosses were unidirectional. All taxa oi Hippochaete are linked by hybrids whereas in subgenus Equisetum, E. pratense and E. sylvatkum form an isolated pair, Hybridization studies, and evidence from gametophyte morphology support the traditional sectional classification of Equisetum.     

Patterns of resource allocation in the dioecious alpine herb Aciphylla simplicifolia (Apiaceae)

Abstract Patterns of reproductive and vegetative biomass allocation were compared in male and female plants of the alpine herb Aciphylla simplicifolia. Male and female plants had similar vegetative biomass but differed in the pattern of resource allocation. Inflorescences of males and females were similar in weight at the time of flowering, but differed in biomass allocation to some structures within the inflorescences, particularly those associated with ovule production and pollinator attraction (number and size of flowers). At the time of fruit production, female inflorescences were 2.6 times heavier than at flowering with developing fruit six times heavier than flowers. In addition to the increase in biomass allocated to structures associated with the provisioning and dissemination of seed, support structures (main and side stalks) were also heavier. As a result of this additional investment of resources at the time of fruit production, the reproductive effort (RE) of female plants was much higher than that of males: 37% of above ground biomass compared with 21% for males. Differences in RE did not change with plant size; however, allocation to reproduction appeared to be a constant proportion of biomass over nearly all plant sizes sampled. These results show that sex‐specific resource allocation can be a complex of temporal and morphological patterns.