Life history and sex allocation in the simultaneously hermaphroditic polychaete worm Ophryotrocha diadema: the role of sperm competition

Sex allocation theory predicts that, in hermaphroditic organisms,individuals allocate a fixed amount of resources divided amongmale and female functions to reproduction and that the proportiondevoted to each sex depends on the mating group size. As themating group size increases, hermaphrodites are predicted toallocate proportionally more resources to the male and lessresources to the female function (approaching equal allocationto both sexes) to face increased sperm competition. Up to nowlittle experimental evidence has been provided to support thetheory in hermaphroditic animals. Facultative shift betweenmale and female allocation in response to variation in localgroup size does occur in several taxa but not always in theexpected direction and not with similar patterns. In the protandricand then simultaneously hermaphroditic polychaete worm Ophryotrochadiadema reproductive resources are flexibly allocated in theprotandrous and the hermaphroditic phase. The cost of male reproductionduring adolescence is spread over the whole energy budget ofthe animal as shown by the shortening of lifespan and the loweringof growth rate in individuals with enhanced male expenditureduring the protandrous phase. Moreover, in this species, shortterm sex allocation adjustments differ from those describedin other taxa. Individuals regulate their reproductive outputso that where reproductive competitors are present, the numberof female gametes is strongly reduced but the number of malegametes (although it changes) is not significantly increased.Resources subtracted from the female function are not directlyallocated to sperm production, but to expensive male behaviorsthat are likely to enhance male reproductive success. Theseresults are discussed in the light of the relevance of sexualselection in large populations of hermaphrodites.     

Sex allocation in a simultaneously hermaphroditic marine shrimp

Two fundamental questions dealing with simultaneous hermaphrodites are how resources are optimally allocated to the male and female function and what conditions determine shifts in optimal sex allocation with age or size. In this study, I explored multiple factors that theoretically affect fitness gain curves (that depict the relationship between sex-specific investment and fitness gains) to predict and test the overall and size-dependent sex allocation in a simultaneously hermaphroditic brooding shrimp with an early male phase. In Lysmata wurdemanni, sperm competition is absent as hermaphrodites reproducing in the female role invariably mated only once with a single other shrimp. Shrimps acting as females preferred small over large shrimps as male mating partners, male mating ability was greater for small compared to large hermaphrodites, and adolescent males were predominant in the population during the breeding season. In addition, brooding constraints were not severe and varied linearly with body size whereas the ability to acquire resources increased markedly with body size. Using sex allocation theory as a framework, the findings above permitted to infer the shape of the male and female fitness gain curves for the hermaphrodites. The absence of sperm competition and the almost unconstrained brooding capacity imply that both curves saturate, however the male curve levels off much more quickly than the female curve with increasing level of investment. In turn, the predominance of adolescent males in the population implies that the absolute gain of the female curve is greater than that of the male curve. Last, the size-dependent female preference and male mating ability of hermaphrodites determines that the absolute gain of the male curve is greater for small than for large hermaphrodites. Taking into consideration the inferred shape of the fitness gain curves, two predictions with respect to the optimal sex allocation were formulated. First, overall sex allocation should be female biased; it permits hermaphrodites to profit from the female function that provides a greater fitness return than the male function. Second, sex allocation should be size-dependent with smaller hermaphrodites allocating more than proportionally resources to male reproduction than larger ones. This size-dependent sex allocation permits hermaphrodites to profit from male mating opportunities that are the greatest at small body sizes. Size-dependent sex allocation is also expected because the male fitness gain curve decelerates more quickly than the female gain curve and experiments indicated that resources are greater for large than small hermaphrodites. These two predictions were tested when determining the sex allocation of hermaphrodites by dissecting their gonad and quantifying ovaries versus testes mass. Supporting the predictions above, hermaphrodites allocated, on average, 118 times more to the female than to the male gonad and the proportion of resources devoted to male function was higher in small than in large hermaphrodites. A trade-off between male and female allocation is assumed by theory but no negative correlation between male and female reproductive investment was observed. In L. wurdemanni, the relationship between sex-specific investment and fitness changes during ontogeny in a way that is consistent with an adjustment of sex allocation to improve size-specific reproductive success.     

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.     

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.     

Experimental tests of sex allocation theory with two species of simultaneously hermaphroditic acorn barnacles

Sex allocation theory for simultaneous hermaphrodites predicts increases in relative allocation to male-specific function as competition for fertilizations increases. Theoretical models developed specifically for competing acorn barnacles predict that the proportional allocation to male function increases toward an asymptote of 50% as the number of competitors for fertilizations increases. Experimental manipulations were used to investigate how mate competition affected both relative and absolute allocation to the sex functions for two species of acorn barnacle: Semibalanus balanoides and Balanus glandula. The ratio of male to female allocation did not increase with the number of competitors for either species. However, both species showed increased allocation to male function (estimated as total mass of sex-specific tissues) with increased crowding. Allocation to female function seemed to be limited by other factors and did not vary with mating group size as predicted. Allocation to male and female function were both positively related to body size, but a trade-off between male and female function, a key assumption of prior models, was not observed.     

多年生龙胆属植物个体大小与花期资源分配研究

于各物种花中前期对青藏高原东部高寒草甸6种多年生龙胆属植物花期的繁殖分配和性分配进行分析,结果表明:(1)多年生龙胆属植物的植株个体越大,繁殖投入越高,繁殖分配越低;(2)随着植物个体的增大,对雌性、雄性和吸引结构的投入都在增加,这可保证资源的充分利用,不会因为单一部分的增加而造成资源的浪费;(3)6种龙胆属植物中,有4种其性分配结果与性别分配(SDS)的理论预测一致,即大个体更偏向雌性器官的资源投入,但麻花艽(Gentiana atraminea)和达乌里秦艽(Gentiana dahurica)的性分配与个体大小则没有表现出负相关,可能与其本身具有的雌雄异熟———雄性先熟特点有关;(4)资源在雌雄功能间的分配没有表现出权衡关系,可能是由于植物必须在许多不同生活史性状之间进行资源分配,而不是两两之间非此即彼.     

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.     

Sex allocation predicts mating rate in a simultaneous hermaphrodite

Sexual selection theory for separate-sexed animals predicts that the sexes differ in the benefit they can obtain from multiple mating. Conventional sex roles assume that the relationship between the number of mates and the fitness of an individual is steeper in males compared with females. Under these conditions, males are expected to be more eager to mate, whereas females are expected to be choosier. Here we hypothesize that the sex allocation, i.e. the reproductive investment devoted to the male versus female function, can be an important predictor of the mating strategy in simultaneous hermaphrodites. We argue that within-species variation in sex allocation can cause differences in the proportional fitness gain derived through each sex function. Individuals should therefore adjust their mating strategy in a way that is more beneficial to the sex function that is relatively more pronounced. To test this, we experimentally manipulated the sex allocation in a simultaneously hermaphroditic flatworm and investigated whether this affects the mating behaviour. The results demonstrate that individuals with a more male-biased sex allocation (i.e. relatively large testes and small ovaries) are more eager to mate compared with individuals with a more female-biased sex allocation (i.e. relatively small testes and large ovaries). We argue that this pattern is comparable to conventional gender roles in separate-sexed organisms.     

INTRASPECIFIC VARIATION IN SEX ALLOCATION IN A SIMULTANEOUS HERMAPHRODITE: THE EFFECT OF INDIVIDUAL SIZE

Within a population of simultaneous hermaphrodites, individuals may vary in both their current reproductive investment (biomass invested in gonads) and in how they allocate that investment between male and female function. In the chalk bass, Serranus tortugarum, estimates of both reproductive allocation and reproductive success as a male and a female can be made for individuals of different sizes. As individuals increase in size, their investment in gamete production increases, and there is a shift in allocation to a stronger female bias. Spawning frequency as a female in pair spawnings and as a male in both pair spawning and streaking (an alternative mating tactic) does not vary with individual size. As a result, larger individuals should release more sperm or eggs per spawn. Size-assortative pair spawning in this species leads to larger individuals having higher potential returns in total male reproductive success than smaller individuals, which should lead to increases in absolute levels of sperm production in larger individuals when individuals compete for fertilizations through sperm competition. However, smaller individuals contribute a smaller proportion of the sperm released in spawns with multiple spawners and thus are under more intense sperm competition than larger individuals, which should select for increases in male allocation in smaller individuals, all else equal. A local-mate-competition (LMC) model predicts that these factors select for increasing absolute male and female investment with individual size but a relative shift to more female-biased allocation as individual size increases. These predictions are supported by gonadal data. The predictions of average male allocation from the quantitative LMC model were 21.6% and 25.7%, whereas the collections averaged 21.3%. This close agreement of both the mean male allocation and its relative shift with individual size between model and data support the hypothesis that size-specific shifts in sex allocation in this species represent an adaptive response to patterns of mating success and sperm competition.     

Field evidence for bi-directional sex change in the polygynous gobiid fish Trimma okinawae

The social condition of bi-directional sex change in the gobiid fish Trimma okinawae was investigated at Akamizu Beach, Kagoshima, Japan. Social groups of T. okinawae usually consisted of a large male and one or more smaller females. The number of females in the group was positively correlated with male body size and groups were usually separated from each other by 1–3 m. In total, 22 instances of female-to-male sex change and three instances of male-to-female sex change were observed during the 16 months that social groups were monitored. Two individuals changed sex twice: female to male and back to female. Female-to-male sex change occurred when the male disappeared from a group. Either the largest remaining female changed sex to male or a large female from another group immigrated and changed sex to male. Larger individuals appear to benefit from becoming male because they can monopolize the breeding opportunities with several females, as reported in other protogynous fishes. Sex change from male-to-female only occurred when a solitary male joined another group as a subordinate. Mortality rates are high in these small fish, therefore joining another group and reproducing as a female is likely to increase the reproductive value of a solitary male.     

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.     

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.     

Bigger testes increase paternity in a simultaneous hermaphrodite,independently of the sperm competition level

Hermaphroditic animals face the fundamental evolutionary optimization problem of allocating their resources to their male vs. female reproductive function (e.g. testes and sperm vs. ovaries and eggs), and this optimal sex allocation can be affected by both pre‐ and post‐copulatory sexual selection. For example, local sperm competition (LSC) – the competition between related sperm for the fertilization of a partner's ova – occurs in small mating groups and can favour a female‐biased sex allocation, because, under LSC, investment into sperm production is predicted to show diminishing fitness returns. Here, we test whether higher testis investment increases an individual's paternity success under sperm competition, and whether the strength of this effect diminishes when LSC is stronger, as predicted by sex allocation theory. We created two subsets of individuals of the simultaneously hermaphroditic flatworm Macrostomum lignano – by sampling worms from either the highest or lowest quartile of the testis investment distribution – and estimated their paternity success in group sizes of either three (strong LSC) or eight individuals (weak LSC). Specifically, using transgenic focal individuals expressing a dominant green‐fluorescent protein marker, we showed that worms with high testis investment sired 22% more offspring relative to those with low investment, corroborating previous findings in M. lignano and other species. However, the strength of this effect was not significantly modulated by the experienced group size, contrasting theoretical expectations of more strongly diminishing fitness returns under strong LSC. We discuss the possible implications for the evolutionary maintenance of hermaphroditism in M. lignano.     

Phenotypically plastic adjustment of sex allocation in a simultaneous hermaphrodite

Sex allocation theory for simultaneous hermaphrodites predicts an influence of the mating group size on sex allocation. Mating group size may depend on the size of the group in which an individual lives, or on the density, but studies to date have not distinguished between the two factors. We performed an experiment in which we raised a transparent simultaneous hermaphrodite, the flatworm Macrostomum sp., in different group sizes (pairs, triplets, quartets and octets) and in different enclosure sizes (small and large). This design allows us to differentiate between the effects of group size and density. After worms reached maturity we determined their reproductive allocation patterns from microscopic images taken in vivo. The results suggest that the mating group size is a function of the group size, and not of the density. They support the shift to higher male allocation in larger mating groups predicted by sex allocation theory. To our knowledge, this is the first study that unambiguously shows phenotypically plastic sex allocation in response to mating group size in a simultaneous hermaphrodite.     

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.     

The Effects of Inflorescence Size and Flower Position on Biomass and Temporal Sex Allocation in <Emphasis Type="Italic">Lobelia sessiliflora</Emphasis>

To test the prediction of sex allocation theory that plants or flowers high in resource status emphasize the female function, we explored the variation in both biomass (the number of pollen grains and ovules) and temporal (male and female durations) sex allocation among and within plants of protandrous Lobelia sessilifolia in relation to plant size and flower position within plants. Among plants, the mean number of pollen grains and ovules per flower of a plant increased with plant size, whereas the mean P/O ratio (number of pollen grains/number of ovules ratio) decreased with plant size. The mean male duration, the mean female duration, and the mean ratio of male duration/flower longevity per flower of a plant were not correlated with plant size. Thus, large plants emphasized female function in terms of biomass sex allocation, which is consistent with the prediction of size-dependent sex allocation theory. The results for temporal sex allocation, however were inconsistent with the theory. Within plants, the mean number of pollen grains and ovules per flower at each position decreased from lower to upper flowers (early to late blooming flowers) and that of the P/O ratio increased from lower to upper flowers. The mean male duration and the mean female duration per flower decreased from lower to upper flowers, whereas the mean ratio of male duration/flower longevity increased from lower to upper flowers. The population sex ratio changed from male-biased to female-biased. Thus, later blooming flowers emphasized the male function in terms of both biomass and temporal sex allocation, consistent with the sex allocation theory, regarding the change in the population sex ratio.     

Floral sex allocation at individual and branch levels in Betula platyphylla var. japonica (Betulaceae), a tall, wind-pollinated monoecious tree species

Floral sex allocation at the individual and first-order branch levels and the relation between these levels were examined in Betula platyphylla var. japonica, a wind-pollinated monoecious tree. Floral sex allocation at the individual level varied with resource availability in a pattern similar to that predicted by the Masaka and Takada model (Journal of Theoretical Biology 240: 114-125). Thus, individual trees with few reproductive resources produced only female or male inflorescences, whereas individuals with many resources rarely had a high male ratio (i.e., number of male inflorescences/total number of inflorescences). Furthermore, the number of male inflorescences tended to reach an upper limit, whereas the number of female inflorescences increased monotonically with increasing reproductive investment. The patterns of floral sex allocation at the first-order branch level were analogous to those at the individual level. Thus, each first-order branch of B. platyphylla var. japonica behaves like an individual, and the floral sex allocation of a given branch does not necessarily represent the individual tree. The effect of the individual-level floral sex ratio on branch-level floral sex allocation indicates that branch behavior is controlled by the individual.     

No seasonal sex-ratio shift despite sex-specific fitness returns of hatching date in a lizard with genotypic sex determination

Sex allocation theory predicts that mothers should adjust their sex-specific reproductive investment in relation to the predicted fitness returns from sons versus daughters. Sex allocation theory has proved to be successful in some invertebrate taxa but data on vertebrates often fail to show the predicted shift in sex ratio or sex-specific resource investment. This is likely to be partly explained by simplistic assumptions of vertebrate life-history and mechanistic constraints, but also because the fundamental assumption of sex-specific fitness return on investment is rarely supported by empirical data. In short-lived species, the time of hatching or parturition can have a strong impact on the age and size at maturity. Thus, if selection favors adult sexual-size dimorphism, females can maximize their fitness by adjusting offspring sex over the reproductive season. We show that in mallee dragons, Ctenophorus fordi, date of hatching is positively related to female reproductive output but has little, if any, effect on male reproductive success, suggesting selection for a seasonal shift in offspring sex ratio. We used a combination of field and laboratory data collected over two years to test if female dragons adjust their sex allocation over the season to ensure an adaptive match between time of hatching and offspring sex. Contrary to our predictions, we found no effect of laying date on sex ratio, nor did we find any evidence for within-female between-clutch sex-ratio adjustment. Furthermore, there was no differential resource investment into male and female offspring within or between clutches and sex ratios did not correlate with female condition or any partner traits. Consequently, despite evidence for selection for a seasonal sex-ratio shift, female mallee dragons do not seem to exercise any control over sex determination. The results are discussed in relation to potential constraints on sex-ratio adjustment, alternative selection pressures, and the evolution of temperature-dependent sex determination.     

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.