Size‐dependent resource allocation and sex allocation in herbaceous perennial plants

Individuals within a population often differ considerably in size or resource status as a result of environmental variation. In these circumstances natural selection would favour organisms not with a single, genetically determined allocation, but with a genetically determined allocation rule specifying allocation in relation to size or environment. Based on a graphical analysis of a simple evolutionarily stable strategy (ESS) model for herbaceous perennial plants, we aim to determine how cosexual plants within a population should simultaneously adjust their reproductive allocation and sex allocation to their size. We find that if female fitness gain is a linear function of resource investment, then a fixed amount of resources should be allocated to male function, and to post‐breeding survival as well, for individuals above a certain size threshold. The ESS resource allocation to male function, female function, and post‐breeding survival positively correlate if both male and female fitness gains are a saturating function of resource investment. Plants smaller than the size threshold are expected to be either nonreproductive or functionally male only.     

How geitonogamous selfing affects sex allocation in hermaphrodite plants

Does the mode of self-pollination affect the evolutionarily stable allocation to male vs. female function? We distinguish the following scenarios. (1) An ‘autogamous’ species, in which selfing occurs within the flower prior to opening. The pollen used in selfing is a constant fraction of all pollen grains produced. (2) A species with ‘abiotic pollination’, in which selfing occurs when pollen dispersed in one flower lands on the stigma of a nearby flower on the same plant (geitonogamy). The selfing rate increases with male allocation but a higher selfing rate does not mean a reduced export of pollen. (3) An ‘animal-pollinated’ species with geitonogamous selfing. Here the selfing rate also increases with male allocation, but pollen export to other plants in the population is a decelerating function of the number of simultaneously open flowers. In all three models selfing selects for increased female allocation. For model 3 this contradicts the general opinion that geitonogamous selfing does not affect evolutionarily stable allocations. In all models, the parent benefits more from a female-biased allocation than any other individual in the population. In addition, in models 2 and 3, greater male allocation results in more local mate competition. In model 3 and in model 2 with low levels of inbreeding depression, hermaphroditism is evolutionarily stable. In model 2 with high inbreeding depression, the population converges to a fitness minimum for the relative allocation to male function. In this case the fitness set is bowed inwards, corresponding with accelerating fitness gain curves. If the selfing rate increases with plant size, this is a sufficient condition for size-dependent sex allocation (more allocation towards seeds in large plants) to evolve. We discuss our results in relation to size-dependent sex allocation in plants and in relation to the evolution of dioecy.     

Effects of phenological constraints on sex allocation in cosexual monocarpic plants

The effects of two phenological constraints in resource investment to reproduction – resource limitation at the flowering stage and unpredictability of resources gained after flowering – on the resource allocation between male and female functions in monocarpic plants are considered using the ESS (evolutionarily stable strategy) approach. The model predicts that the sex allocation including the seed maturation stage has a female bias, when the quantity of reproductive resources available at flowering is small compared with that which is obtained after flowering, or when the cost of seed maturation relative to ovule production is low. The fluctuation of the quantity of resources available for seed maturation favors overproduction of ovules. As a result, more resources are allocated to female function and less to male function at flowering. The ESS allocation depends on the variability of resources and the cost of seed maturation relative to ovule production. The probability that total resource allocation has a female bias becomes higher than 0.5, and it depends on the cost of seed maturation relative to ovule production rather than resource variability. On the other hand, the probability that resource allocation has a female bias decreases with resource variability if we assume that the floral sex ratio is fixed. Future studies of plant sex allocation would profit by taking account of the phenological process of reproduction such as ovule production or seed maturation.     

ATTRACTIVE STRUCTURES AND THE STABILITY OF HERMAPHRODITIC SEX EXPRESSION IN FLOWERING PLANTS

Plant attributes serving both male and female sex functions are thought to favor hermaphroditic breeding systems over systems where sex expression is separated between individuals. Morphological features used by plants to attract biotic pollinators can be considered in this light, because such “attractive” structures are prerequisite for both pollen donation and receipt. A model involving allocation of a limiting resource between attractive structures benefiting both sex functions, structures benefiting male function only, and structures benefiting female function only, is constructed and analyzed using the evolutionary stable strategy (ESS) approach. Contrary to expectation, conditions resulting in large allocation to attractive structures tend to destabilize hermaphroditism, although the effect is slight. Substantial asymmetry in the benefits accruing to male and female fertility for a given investment in pollinator attraction produces more pronounced effects, reducing the parameter space in which hermaphroditism is expected. Results of this model are used to assess the importance of “accelerating gain” curves previously suggested for the evolution of dioecy in flowering plants. Accelerating gains associated with fruit production and dispersal represent female function only, and can be sufficient to initiate evolution toward dioecy. Accelerating gains associated with pollinator attraction may contribute to the evolution of separate sexes only if male function benefits from increased investment in attractive structures at a disproportionate rate compared to female function. Even in this instance, though, hermaphroditism may be evolutionarily stable.     

Variation in lifetime male fitness in Ipomopsis aggregata: tests of sex allocation theory

Sex allocation theory assumes that a shift in allocation of resources to male function both increases male fitness and decreases female fitness. Moreover, the shapes of these fitness gain functions determine whether hermaphroditism or another breeding system is evolutionarily stable. In this article, I first outline information needed to measure these functions in flowering plants. I then use paternity analysis to describe the shapes of the fitness gain functions in natural populations of the hermaphroditic herb Ipomopsis aggregata. I also explore the relationships of male fitness (number of seeds sired) and female fitness (number of seeds produced) to the number of flowers produced by a plant. Plants with greater investment of biomass in the androecium, compared to the gynoecium and seeds, showed increased success at siring seeds, assumed by the models. That sex allocation trait, however, explained only 9% of the variance in estimates of male fitness. The shapes of the fitness gain functions were consistent with theoretical expectations for a hermaphroditic plant, but the model predicted a more female-biased evolutionarily stable strategy (ESS) allocation than was observed. These results lend only partial support the classical sex allocation model.     

Size-dependent sex allocation in hermaphroditic plants: the effects of resource pool and self-incompatibility

The effects of the resource pool and resource obtained during a season for seed maturation and self-incompatibility on the size-dependency of evolutionarily stable sex allocation were analysed theoretically. In hermaphroditic plants, reproductive resources allocated between male and female function may not be paid from a single resource pool, because plants can mature seeds using not only reserved resources but also newly gained resources after flowering. But the resource investment to male function is limited to the flowering stage. Under the assumption of constant reserve efficiency and diminishing resource return per investment to leaves, large plants should use both reserved and newly gained resources for seed maturation, while small plants should use only new resources. When both reserved and new resources are used, the optimal allocation for self-compatible species is to invest a constant amount of resources into male function irrespective of resource size, because the female fitness curve increases linearly and the male curve decelerates due to local mate competition. In self-incompatible species, on the other hand, fitness gain per investment through male function and the optimal amount of resources invested in male function decrease with size. Thus a decrease in maleness with size should be emphasized more in self-incompatible species than in self-compatible one. When only new resources are used for seed growth, the female fitness curve as well as male one decelerates with investment. Consequently, the investment in both male and female functions should increase with size, in both self-compatible and self-incompatible species. The magnitude of reserve efficiency relative to efficiency of resource gain after flowering affects size-dependent pattern of sex allocation, while the cost of seed maturation relative to ovule production has little effect on it. The plant size variation in a population emphasizes size-dependency of sex allocation. When size variation is large enough, it is possible that large plants become complete female in self-incompatible species, but it is not in self-compatible species.     

植物交酸系统的进化、资源分配对策与遗传多样性

影响植物自交率进化的选择力量主要体现在两个方面：当外来花粉量不足时,自交可以提高植物的结实率,即雌性适合度（繁殖保障）;而如果进行自交的花粉比异交花粉更易获得使胚珠受精的机会,那么自交也可以提高植物的雄性适合度（自动选择优势）。但是,鉴别什么时候是繁殖保障、什么时候是自动选择优势导致了自交的进化却是极其困难的。花粉贴现降低了自交植物通过异交花粉途径获得的适合度,即减弱了自动选择优势,而近交衰退既减少了自动选择优势也减少了繁残给自交者带来的利益。具有不同交配系统的植物种群将具有不同的资源分配对策。理论研究已经说明,自交率增加将减少植物对雄性功能的资源分配比例,但将使繁殖分配加大,而且在一定条件下交配系统在改变甚至可以导致植物生活史发生剧烈变化,即从多年生变为一年生。文献中支持自交减少植物雄性投入的证据有很多,但是对繁殖分配与自交率的关系目前还没有系统的研究,资源分配理论可以解释植物繁育系统的多样性,尤其是能够3说明为什么大多数植物都是雌雄同体的,自交对植物种群遗传结构的影响是减少种群内的遗传变异,增加种群间的遗传分化,长期以来人们一直猜测,自交者可能会丢掉一些长期进化的潜能,目前这个假说得到了一些支持。     

Sex allocation in clonal plants: might clonal expansion enhance fitness gains through male function?

The returns on investment in sexual reproduction are described by fitness gain curves and the shapes of these curves affect, among other things, the evolutionary stability of reproductive systems. The available evidence indicates that gain curves for male function decelerate, corresponding to diminishing fitness returns on investment in pollen. In contrast, the gain curve for female function is thought to decelerate less strongly than it does for male function (e.g., if seed fertility is limited by more by resources than by mating opportunities). Here we suggest that when the shapes of the female and male gain curves differ, clonality alters the rates of return on investment via the two sex functions. In particular, we propose that clonal expansion might increase fitness gains through male function because the subdivision of reproductive effort among ramets allows each ramet to take advantage of the steepest parts of the male gain curve. We examined the interaction between clonal expansion and fitness gains using numerical analysis of a model of sex allocation in which we assumed that there is no mating interference among ramets. We found that clonal expansion led to substantial increases in fitness through male function, but to decreases in fitness through female function. Under intermediate investment in clonal growth, marginal fertility gains through the two sex functions did not intersect over a broad range of sex allocation patterns, suggesting that clonality could favor the evolution of separate sexes. Finally, we suggest an alternative explanation for the common observation of male-biased sex ratios in clonal dioecious plants. If male function fitness is maximized under higher rates of clonal expansion than for female function, greater frequencies of male ramets might reflect the outcome of fertility selection, rather than constraints on clonal expansion imposed by greater costs of reproduction for females.     

植物交配系统的进化、资源分配对策与遗传多样性

影响植物自交率进化的选择力量主要体现在两个方面：当外来花粉量不足时,自交可以提高植物的结实率,即雌性适合度（繁殖保障）;而如果进行自交的花粉比异交花粉更易获得使胚珠受精的机会,那么自交也可以提高植物的雄性适合度（自动选择优势）。但是,鉴别什么时候是繁殖保障、什么时候是自动选择优势导致了自交的进化却是极其困难的。花粉贴现降低了自交植物通过异交花粉途径获得的适合度,即减弱了自动选择优势,而近交衰退既减少了自动选择优势也减少了繁殖保障给自交者带来的利益。具有不同交配系统的植物种群将具有不同的资源分配对策。理论研究已经说明,自交率增加将减少植物对雄性功能的资源分配比例,但将使繁殖分配加大,而且在一定条件下交配系统的改变甚至可以导致植物生活史发生剧烈变化,即从多年生变为一年生。文献中支持自交减少植物雄性投入的证据有很多,但是对繁殖分配与自交率的关系目前还没有系统的研究。资源分配理论可以解释植物繁育系统的多样性,尤其是能够说明为什么大多数植物都是雌雄同体的。自交对植物种群遗传结构的影响是减少种群内的遗传变异,增加种群间的遗传分化。长期以来人们一直猜测,自交者可能会丢掉一些长期进化的潜能,目前这个假说得到了一些支持。     

SEX ALLOCATION IN THE MONOECIOUS HERB BEGONIA SEMIOVATA

Sex-allocation models predict that the evolution of self-fertilization should result in a reduced allocation to male function and pollinator attraction in plants. The evolution of sex allocation may be constrained by both functional and genetic factors, however. We studied sex allocation and genetic variation for floral sex ratio and other reproductive traits in a Costa Rica population of the monoecious, highly selfing annual Begonia semiovata. Data on biomass of floral structures, flower sex ratios, and fruit set in the source population were used to calculate the average proportion of reproductive allocation invested in male function. Genetic variation and genetic correlations for floral sex ratio and for floral traits related to male and female function were estimated from the greenhouse-grown progeny of field-collected maternal families. The proportion of reproductive biomass invested in male function was low (0.34 at flowering, and 0.07 for total reproductive allocation). Significant among-family variation was detected in the size (mass) of individual male and female flowers, in the proportion of male flowers produced, and in the proportion of total flower mass invested in male flowers. Significant among-family variation was also found in flower number per inflorescence, petal length of male and female flowers, and petal number of female flowers. Except for female petal length, we found no difference in the mean value of these characters between selfed and outcrossed progeny, indicating that, with the possible exception of female petal length, the among-family variation detected was not the result of variation among families in the level of inbreeding. Significant positive phenotypic and broad-sense genetic correlations were detected between the mass of individual male and female flowers, between male and female petal length, and between number of male and number of female flowers per inflorescence. The ratio of stamen-to-pistil mass (0.33) was low compared to published data for autogamous species with hermaphroditic flowers, suggesting that highly efficient selfing mechanisms may evolve in monoecious species. Our results indicate that the study population harbors substantial genetic variation for reproductive characters. The positive genetic correlation between investment in male and female flowers may reflect selection for maximum pollination efficiency, because in this self-pollinating species, each female flower requires a neighboring male flower to provide pollen.     

Floral dimorphism, pollination, and self-fertilization in gynodioecious GERANIUM RICHARDSONII (Geraniaceae)

The selective maintenance of gynodioecy depends on the relative fitness of the male-sterile (female) and hermaphroditic morphs. Females may compensate for their loss of male fitness by reallocating resources from male function (pollen production and pollinator attraction) to female function (seeds and fruits), thus increasing seed production. Females may also benefit from their inability to self-fertilize if selfing and inbreeding depression reduce seed quality in hermaphrodites. We investigated how differences in floral resource allocation (flower size) between female and hermaphroditic plants affect two measures of female reproductive success, pollinator visitation and pollen receipt, in gynodioecious populations of Geranium richardsonii in Colorado. Using emasculation treatments in natural populations, we further examined whether selfing by autogamy and geitonogamy comprises a significant proportion of pollen receipt by hermaphrodites. Flowers of female plants are significantly smaller than those of hermaphrodites. The reduction in allocation to pollinator-attracting structures (petals) is correlated with a significant reduction in pollinator visitation to female flowers in artificial arrays. The reduction in attractiveness is further manifested in significantly less pollen being deposited on the stigmas of female flowers in natural populations. Autogamy is rare in these protandrous flowers, and geitonogamy accounts for most of the difference in pollen receipt between hermaphrodites and females. Female success at receiving pollen was negatively frequency dependent on the relative frequency of females in populations. Thus, two of the prerequisites for the maintenance of females in gynodioecious populations, differences in resource allocation between floral morphs and high selfing rates in hermaphrodites, occur in G. richardsonii.     

THE EFFECT OF INVESTMENT IN ATTRACTIVE STRUCTURES ON ALLOCATION TO MALE AND FEMALE FUNCTIONS IN PLANTS

Expressions for male and female fitnesses of partially self-fertilizing cosexual plants are derived, assuming that allocation to pollinator attraction at the time of flowering may decrease resources available for male and female primary structures. The total female fertility is assumed to be controlled by factors at two stages, flowering-time and fruiting-time, with resources for fruit maturation being limited so that maximum seed production may be limited by the availability of these resources. The fitness formulas are used to calculate ESS (evolutionarily stable strategy) allocations at flowering time to primary male and female sex functions and to attractive structures. These are compared with some data that are available for dry weights of different flower parts. The fitnesses of unisexual mutant forms are calculated, assuming that they are introduced into a population consisting mostly of the initial cosexual form and that they obey the same gain curves as that form. When compared with the fitness of the ESS cosexual form, this enables one to ask whether unisexual forms will be favored. We show that the spread of females is unlikely, unless there is high inbreeding depression and a rather high selfing rate, and that in some circumstances a linear relation between number of fertilized ovules and number of seeds matured can be less favorable for the invasion of females than is a highly concave relation. With a nearly linear relation between numbers of fertilized ovules and mature seeds, invasion by females is more likely when investment in attraction is low than when it is high. These effects are discussed in relation to the distribution of dioecy. The spread of male mutants is never likely in these models.     

FLORAL SEX ALLOCATION IN SEQUENTIALLY BLOOMING PLANTS

In plants whose flowers develop in a sequence, different flowers may exhibit temporal variation in pollen donation and receipt such that the fitness contributions through male and female functions can vary among flowers. Dichogamy, or directional pollinator movements within inflorescences, can create situations where flowers in different stages in the sequence may differ in the numbers of flowers in the female stage available as potential mates. We present an evolutionarily stable strategy (ESS) analysis of the resource allocations expected in different flowers in hermaphroditic plants when the mating environments vary among flowers. This introduces a modular element into sex-allocation models. Our analysis shows that such variation in the mating environments of flowers can select for differences in sex allocation between flowers. When male and female fertilities are nonlinear functions of the allocations, variation in resource availability can also select for variation in sex allocation among flowers. The influence of dichogamy and pollinator directionality on floral sex allocation is discussed, and the empirical evidence supporting the predictions derived from the model is briefly reviewed. The implications of our results for the evolution of andromonoecy and monoecy are discussed.     

Allocation of resources to male and female functions in hermaphrodites

The question of"how a self-fertile hermaphrodite will distribute the resources that it allocates to reproduction is studied by means of the ESS approach. Different models of the relations between allocation to male function, the male and female fertilities, and the selfing rate, yield different conclusions about how much resource should be allocated to male function. Values below a half are obtained with one model, while another can give values greater than a half. Even with no selfing, values other than a half are usually obtained; with both models studied, the values decrease with increasing selling. If the selfing rate is assumed to be independent of the fraction of resources allocated to male function, it can be shown that the ESS allocation to male function always decreases as selling increases. The types of relations that might be expected in species with different types of breeding biology, and some data on allocation to male function, are reviewed.
The implications for the fitness of male- and female-sterility mutations are discussed. It is argued that the concavity or convexity of the curve relating female fertility to male fertility is not a good guide to when hermaphroditism should exist when there is some selfing. Even with a concave relation, male-sterility mutants can have a higher fitness than hermaphrodites, if there is some selling and inbreeding depression. Also, when the selfing rate depends on allocation to male I unction, an hermaphrodite ESS does not always exist when the function is concave (as it does when there is no selfing), and such an ESS may exist when the relation is convex. The fitness of male- or female-sterility mutants may also depend on the existence of 'fixed costs'. It is shown that these do not ailed the ESS allocation of resources.     

繁殖保障和延迟自交的研究进展

尽管植物在进化过程中面临不利自花授粉的选择,但许多植物仍维持混合的授粉机制。繁殖保障假说是解释自交进化的最重要因子之一,一直是植物生殖生态学和进化生物学关注的焦点之一。概述了近年来的主要研究热点及其进展,包括自交进化的遗传和生态机制及理论模型探讨、繁殖保障假说的提出、验证自交能否提供繁殖保障的例证、延迟自交的类型及延迟自交能否提供繁殖保障的例证等方面。介绍了我国在繁殖保障和延迟自交方面研究的现状和不足之处,结合国际上研究繁殖保障假说的发展趋势已由单季节、单种群、单因子的研究阶段过渡到多季节、多种群、多因子（自交方式及其所占比例、花粉折损、种子折损、自交率和近交衰退）的综合研究阶段,及由传统的、经典的研究方法过渡到应用现代实验手段（如SSR、SNP等分子标记）和先进仪器设备的研究阶段,提出今后研究中应注意的问题。有必要借用多学科（植物学、生态学和分子生物学）的方法及手段进行不同物种的对比和综合细致的研究。     

Size-dependent sex allocation within flowers of the annual herb Clarkia unguiculata (Onagraceae): ontogenetic and among-plant variation

The relative allocation of resources to male and female functions may vary among flowers within and among individual plants for many reasons. Several theoretical models of sex allocation in plants predict a positive correlation between the resource status of a flower or individual and the proportion of reproductive resources allocated to female function. These models assume that, independent of resource status, a negative correlation exists between male and female investment. Focusing on the allocation of resources within flowers, we tested these theoretical predictions and this assumption using the annual Clarkia unguiculata (Onagraceae). We also sought preliminary evidence for a genetic component to these relationships. From 116 greenhouse-cultivated plants representing 30 field-collected maternal families, multiple flowers and fruits per plant were sampled for gamete production, pollen?:?ovule ratio, seed number, ovule abortion, seed biomass/fruit, mean individual seed mass, and petal area. If sex allocation changes as predicted, then (1) assuming that flowers produced early have access to more resources than those produced later, basal flowers should exhibit a higher absolute and proportional investment in female function than distal flowers and (2) plants of high resource status (large plants) should produce flowers with a higher proportional investment in female function than those of low resource status. Within plants, variation in floral traits conformed to the first prediction. Among plants and families, no significant effects of plant size (dry stem biomass) on intrafloral proportional sex allocation were observed. We detected no evidence for a negative genetic correlation between male and female investment per flower, even when controlling for plant size.     

Main role of self-pollination rate on reproductive allocations in pseudogamous apomicts

 Most apomicts are hermaphroditic and pseudogamous (pollination is necessary to trigger parthenogenesis). In these plants, fitness depends on the number of progeny obtained by maternal reproduction. We determined the evolutionary stable strategy for male and female sex allocation. We show that the efficiency of pollination determines male and female resource allocations. Predictions are made of these allocations, of pollen/ovule ratio and of seed-set. We show that self-compatibility in apomicts is necessary for the maintenance of an apomictic population, and thus can account for the association between the loss of self-incompatibility and pseudogamous apomixis. In contrast to sexuals, male investment in pseudogamous apomicts increases with the rate of self-pollination. Received: 15 June 1996 / Accepted: 20 September 1996     

Size-dependent resource allocation among vegetative propagules and male and female functions in the forest herb Laportea bulbifera

Reproductive resource investment among vegetative propagules and male and female sexual function and their size-dependence were investigated in a perennial forest herb, Laportea bulbifera . A theoretical model based on fitness gain curves predicts that optimal investments in three reproductive modes will increase with plant size if fitness returns in all three modes increase but become saturated with investment. In a field population, large plants of L. bulbifera produced both male and female inflorescences with propagules, while small plants produced only vegetative propagules. Biomass of propagules, male inflorescences, and infructescences with achenes were all positively correlated with plant size. The increase in investment with plant size was larger for propagule production than for sexual reproduction. The relationship between propagule biomass and plant size was constant irrespective of year, while the relationship between the biomass of sexual reproductive organs and plant size differed between two successive years. Annual change of individual sex expression was investigated for 25 transplanted plants. Although each plant changed its sex expression variously among male, female and bisexual from year to year, 23 out of 25 plants produced both male and female inflorescences in at least one year. The number of viable (germinated and survived) offspring from seeds was not significantly different from the number from propagules. The production cost of a propagule was higher than that of a seed. Resource allocation theory does not seem to be applicable to size-dependent resource allocation, especially the allocation between seeds and propagules in this species.     

Spatial seed and pollen games: dispersal,sex allocation,and the evolution of dioecy

The evolutionary forces shaping within‐ and across‐species variation in the investment in male and female sex function are still incompletely understood. Despite earlier suggestions that in plants the evolution or cosexuality vs. dioecy, as well as sex allocation among cosexuals, is affected by seed and pollen dispersal, no formal model has explicitly used dispersal distances to address this problem. Here, we present a game‐theory model as well as a simulation study that fills in this gap. Our model predicts that dioecy should evolve if seeds and pollen disperse widely and that sex allocation among cosexuals should be biased towards whichever sex function produces more widely dispersing units. Dispersal limitations stabilize cosexuality by reinforcing competition between spatially clumped dispersal units from the same source, leading to saturating fitness returns that render sexual specialization unprofitable. However, limited pollen dispersal can also increase the risk of selfing, thus potentially selecting for dioecy as an outbreeding mechanism. Finally, we refute a recent claim that cosexuals should always invest equally in both sex functions.     

Intraspecific variation in sex allocation in hermaphroditic Plantago coronopus (L.)

Models for sex allocation assume that increased expenditure of resources on male function decreases the resources available for female function. Under some circumstances, a negative genetic correlation between investment in stamens and investment in ovules or seeds is expected. Moreover, if fitness returns for investment in male and female function are different with respect to size, sex allocation theory predicts size‐specific gender changes. We studied sex allocation and genetic variation for investment in stamens, ovules and seeds at both the flower and the plant level in a Dutch population of the wind‐pollinated and predominantly outcrossing Plantago coronopus. Data on biomass of floral structures, stamens, ovules, seedset and seedweight were used to calculate the average proportion of reproductive allocation invested in male function. Genetic variation and (genetic) correlations were estimated from the greenhouse‐grown progeny of maternal families, raised at two nutrient levels. The proportion of reproductive biomass invested in male function was high at flowering (0.86 at both nutrient levels) and much lower at fruiting (0.30 and 0.40 for the high and low nutrient treatment, respectively). Androecium and gynoecium mass exhibited moderately high levels of genetic variance, with broad‐sense heritabilities varying from 0.35 to 0.56. For seedweight no genetic variation was detected. Significant among‐family variation was also detected for the proportion of resources invested in male function at flowering, but not at fruiting. Phenotypic and broad‐sense genetic correlations between androecium and gynoecium mass were positive. Even after adjusting for plant size, as a measure of resource acquisition, maternal families that invested more biomass in the androecium also invested more in the gynoecium. This is consistent with the hypothesis that genetic variation for resource acquisition may in part be responsible for the overall lack of a negative correlation between male and female function. Larger plants had a more female‐biased allocation pattern, brought about by an increase in seedset and seedweight, whereas stamen biomass did not differ between small and large plants. These results are discussed in relation to size‐dependent sex allocation theory (SDS). Our results indicate that the studied population harboured substantial genetic variation for reproductive characters.