SEXUAL DIMORPHISM IN BIOMASS ALLOCATION AND CLONAL GROWTH OF XANTHOXYLUM AMERICANUM

Reproduction can have a high resource cost. It has been suggested that greater investments in sexual reproduction by female dioecious plants leads to a lower rate of vegetative growth in females than in males. In this study, we investigated sexual dimorphism in biomass allocation and genet growth of the dioecious clonal shrub, northern prickly ash (Xanthoxylum americanum). The allocation of biomass over the course of one growing season to reproductive tissue, leaves, and growth of aboveground first-year wood, was compared in 18 clones growing in fields and six clones in woods in southeastern Wisconsin during 1985 and 1986. In addition, the number of shoots per clone, and weight of nonfirst-year wood (accumulated biomass) above- and below-ground were estimated. In open field sites, male clones allocated more biomass to new wood and less to reproduction than females, although males allocated more to flowers alone. Accordingly, male clones had significantly more shoots and more accumulated biomass both above- and below-ground than female clones. In the woods, where fruit set was near zero, there were few significant differences between male and female clones in either biomass allocation or accumulated biomass. These results support the hypothesis that the high resource investment in fruit production by females reduces their vegetative growth relative to males.     

Reproductive allocation and costs in gynodioecious Leucopogon melaleucoides (Ericaceae): implications for the evolution of gender dimorphism

In dioecious species, females typically allocate more resources to reproduction and incur greater costs of reproduction than males. In gynodioecious species, sex-based differences in reproductive allocation (RA) and costs have been less studied. Such knowledge, however, is relevant to address how females establish and increase in frequency in populations. We examine RA and reproductive costs by comparing fruit set, the proportion of biomass allocated to reproduction, and the responses of fruit set and vegetative growth to shoot defoliation in females and hermaphrodites in gynodioecious Leucopogon melaleucoides. Relative to hermaphrodites, females exhibited a two-fold fruit set advantage. Female fruit set increased proportionately with flower number, but hermaphrodite fruit set was reduced on plants with more flowers. Sex-based differences in allocation to other traits were small. Thus, female RA at flowering was similar to hermaphrodite RA, but was 1.4-fold greater at fruiting. Relative to controls, defoliation reduced fruit set and the percentage of shoots that produced new vegetative growth similarly in both sexes. However, females had a lower proportion of shoots with new growth overall. Further, defoliation on females reduced the dry mass of new growth by 44% compared with controls, whereas hermaphrodites were not affected. These results indicate a trade-off between reproduction and vegetative growth, and greater female costs of reproduction, particularly under resource-limiting conditions. In the absence of compensatory traits to offset higher female reproductive costs, such trade-offs have the potential to retard the spread of females in gynodioecious populations.     

DIFFERENCES IN BIOMASS ALLOCATION TO REPRODUCTIVE AND VEGETATIVE STRUCTURES OF MALE AND FEMALE PLANTS OF A DIOECIOUS,PERENNIAL HERB,SILENE ALBA (MILLER) KRAUSE

The pattern of biomass allocation of males and females and the sex ratio and growth characteristics of plants from three seed-size classes in Silene alba were investigated in a greenhouse study. Seed size significantly affected adult plant size and flower production of both male and female plants, but there was no significant difference in the proportion of males and females emerging in three seed-size categories. Male and female plants differed in the proportion of total biomass allocated to vegetative and reproductive structures and these differences were consistent across all seed-size categories. Males allocated a greater proportion of their biomass to flowers than did females. Female reproductive effort was dependent upon the percentage of flowers producing mature capsules. Only females with greater than 20% fruit set have a higher reproductive expenditure than males. Consequently, female expenditure is potentially greater than males, but is spread out over a longer portion of the growing season. This difference in the timing of reproductive expenditures by males and females allows females to allocate more biomass to growth during the early flowering period and may therefore account for the common pattern in herbaceous perennial dioecious species in which adult females are larger than adult males.     

The effect of flight on reproduction in an outbreaking forest lepidopteran

Post‐flight reproductive investment by female insects may be limited as a result of a trade‐off in resource allocation between flight and reproduction. Outbreaking forest pests reduce their habitat quality as a result of severe defoliation when population densities are high. Female relocation to better‐quality habitats can increase offspring survival but reduce their reproductive fitness through flight. In the present study, the effect of flight on the capacity of female Choristoneura conflictana (Walker) (Lepidoptera: Tortricidae) to mate and produce eggs is examined. Females are flown on flight mills, and the subsequent reproductive capacity of each moth is assessed through measures of mating success and egg production. There is no effect of flight on commencement or the duration of mating. Although flight does not affect egg production directly, energy expenditure as a result of flight (as measured by weight loss) shows a negative correlation to potential fecundity, possibly indicating the resorption of eggs in some females. The effect of female size on fecundity is dependent on mating status, suggesting that energy allocated to reproduction is not dependent on flight treatment. Female moth longevity also has a significant effect on egg production but is dependent on flight and mating treatments. There is a relationship between energy expenditure to flight and reproduction in C. conflictana. Females that fly away from dense populations may produce fewer offspring, although this cost may be mitigated by improved offspring survivorship in less defoliated habitats.     

Reproductive correlates of mating system variation in Eichhornia paniculata (Spreng.) Solms (Pontederiaceae)

Comparative studies of related plant species indicate that evolutionary shifts in mating systems are accompanied by changes in reproductive attributes such as flower size, floral morphology, and pollen/ovule ratio. Recent theoretical work suggests that patterns of investment in reproduction should also change with the mating system. In a glasshouse study, we investigated the extent to which mating system differences among populations of Eichhornia paniculata (Pontederiaceae) were correlated with changes in allocation to male and female function, floral display, and the regulation of investment in reproduction through fruit and ovule abortion. Significant differences in the amount of biomass allocated to reproductive structures were evident among six populations of E. paniculata. As predicted by sex allocation theory, the proportion of dry weight allocated to male function decreased with the outcrossing rate of populations. Six of the eight attributes used to characterize floral display also differed significantly among populations. However, with the exception of two attributes describing the number of flowers produced by inflorescences, these were not correlated with outcrossing rate. Levels of fruit and ovule abortion were determined in two populations with contrasting mating systems under different nutrient and pollination treatments. Virtually all fruits initiated by plants from a self-fertilizing population were matured, while the amount of fruit abortion in an outcrossing population increased with flower production. Ovule abortion was low in both populations. Our results demonstrate that the evolution of self-fertilization in E. paniculata is associated with changes in investment to reproduction that normally distinguish selfing and outcrossing species.     

Soil water content and patterns of allocation to below- and above-ground biomass in the sexes of the subdioecious plant Honckenya peploides

Background and aims Dioecious plants often show sex-specific differences in growth and biomass allocation. These differences have been explained as a consequence of the different reproductive functions performed by the sexes. Empirical evidence strongly supports a greater reproductive investment in females. Sex differences in allocation may determine the performance of each sex in different habitats and therefore might explain the spatial segregation of the sexes described in many dimorphic plants. Here, an investigation was made of the sexual dimorphism in seasonal patterns of biomass allocation in the subdioecious perennial herb Honckenya peploides, a species that grows in embryo dunes (i.e. the youngest coastal dune formation) and displays spatial segregation of the sexes at the studied site. The water content in the soil of the male- and female-plant habitats at different times throughout the season was also examined. Methods The seasonal patterns of soil-water availability and biomass allocation were compared in two consecutive years in male and female H. peploides plants by collecting soil and plant samples in natural populations. Vertical profiles of below-ground biomass and water content were studied by sampling soil in male- and female-plant habitats at different soil depths. Key Results The sexes of H. peploides differed in their seasonal patterns of biomass allocation to reproduction. Males invested twice as much in reproduction than females early in the season, but sexual differences became reversed as the season progressed. No differences were found in above-ground biomass between the sexes, but the allocation of biomass to below-ground structures varied differently in depth for males and females, with females usually having greater below-ground biomass than males. In addition, male and female plants of H. peploides had different water-content profiles in the soil where they were growing and, when differences existed (usually in the upper layers of the soil), the water content of the soil was higher for the female plants had than for the male plants. Conclusions Sex-differential timing of investment in reproduction and differential availability and use of resources from the soil (particularly water) are factors that probably offset the costs of reproduction in the above-ground growth in males and females of H. peploides. The results suggest that the patterns of spatial segregation of the sexes observed in H. peploides may contribute to maximize each sex's growth and reproduction.     

Sexual dimorphism in resource acquisition and deployment: both size and timing matter

Background and Aims

The males and females of many dioecious plant species differ from one another in important life-history traits, such as their size. If male and female reproductive functions draw on different resources, for example, one should expect males and females to display different allocation strategies as they grow. Importantly, these strategies may differ not only between the two sexes, but also between plants of different age and therefore size. Results are presented from an experiment that asks whether males and females of Mercurialis annua, an annual plant with indeterminate growth, differ over time in their allocation of two potentially limiting resources (carbon and nitrogen) to vegetative (below- and above-ground) and reproductive tissues.

Methods

Comparisons were made of the temporal patterns of biomass allocation to shoots, roots and reproduction and the nitrogen content in the leaves between the sexes of M. annua by harvesting plants of each sex after growth over different periods of time.

Key Results and Conclusions

Males and females differed in their temporal patterns of allocation. Males allocated more to reproduction than females at early stages, but this trend was reversed at later stages. Importantly, males allocated proportionally more of their biomass towards roots at later stages, but the roots of females were larger in absolute terms. The study points to the important role played by both the timing of resource deployment and the relative versus absolute sizes of the sinks and sources in sexual dimorphism of an annual plant.     

Reproductive allocation in hermaphrodite and female plants of Sidalcea oregana SSP. spicata (Malvaceae) using four currencies

Reproductive allocation was investigated in female and hermaphrodite plants of gynodioecious Sidalcea oregana ssp. spicata. Total reproductive investment and partitioning of that investment was documented at the level of whole plants in terms of four ecologically relevant currencies: biomass, nitrogen, phosphorus, and potassium. Nutrient augmentations in the field confirmed that nutrients were limiting plant vegetative growth and propensity to flower; thus the use of these nutrients as currency was appropriate. Once the effects of plant size were removed, the sex morphs allocated similar total amounts of biomass, nitrogen, phosphorus, and potassium to reproduction, but partitioned those differentially. For any given individual size, females allocated larger proportions of their reproductive resource budgets to seeds. Hermaphrodites' reproductive investment in pollen and flowers was allocated at the expense of allocation to seeds. These data are relevant to the evolution of gynodioecy from hermaphroditism and support the hypothesis that females reallocate resources not spent on pollen to seeds.     

Costs of reproduction as related to the timing of phenological phases in the dioecious Shrub Pistacia lentiscus L

Females of woody dioecious species usually devote more resources to reproduction than males. This may lead to a decrease in female survival and growth. The costs of reproduction, however, can be lightened through a number of mechanisms, as for example avoiding the temporal coincidence of reproduction and vegetative growth. The aim of this study was to evaluate whether males and females of P. lentiscus differ in the timing of their vegetative growth, and to assess whether the sequencing of vegetative growth and reproduction reduces reproductive costs. We monitored phenology in males and females. We also compared male and female allocation of nutrients and biomass in the branch, and the developmental stability of the growing shoots. We did this both prior to and at the end of the fruiting period. Males and females showed similar vegetative and flowering phenologies. Males invested more biomass in flowering, but the sexes showed equal vegetative biomass and nutrient content prior to the fruiting period. In female branches, no trade-off was found between fruit load and current-year vegetative growth. In P. lentiscus, avoiding the overlap of flowering, vegetative growth and fruiting probably contributes to reduce the immediate costs of reproductive efforts, both in males and females.     

Energy allocation for reproduction in a marsupial arboreal folivore,the common ringtail possum (Pseudocheirus peregrinus)

This study examines the annual energetics of a small folivorous marsupial, Pseudocheirus peregrinus. Particular attention was given to the energy and time allocated to reproduction by the females. Daily energy expenditure was measured directly using the doubly labelled water technique. Energy transferred to the young via the milk was estimated from information on milk composition and production. There was no significant seasonal variation in the energy expenditure or water influx of males or females. The mean daily energy expenditure of a 1-kg non-lactating adult ringtail possum was 615 kJ day^–1 or 2.2 times standard metabolic rate. Females showed significant changes in daily energy expenditure according to their reproductive status. Without the burden of lactation the total annual energy expenditure of an adult female was estimated as 212.4 MJ kg^–1 year^–1. The total annual energy expenditure of a female rearing two young was 247.5 MJ kg^–1 year^–1, with the late stage of lactation constituting the most energetically expensive period accounting for 30% of the total yearly energy expenditure during 24% of the time. Total metabolisable energy allocation during reproduction (22 MJ kg) was similar to estimates available for other herbivores, although, the peak metabolisable energy allocation during lactation (759 kJ day^–1) was lower than values available for other herbivores. The total energy requirement for reproduction (metabolisable energy plus potential energy exported to young via milk) suggests that the ringtail possum also has a relatively low overall energy investment in reproduction. It is suggested that the lactational strategy of the ringtail possum has been selected in order to spread the energy demands of reproduction over time due to constraints on the rate of energy intake imposed by a leaf diet and/or to prolong the mother-young bond. The strategies a female ringtail possum may employ to achieve energy balance when faced with the energy demands of reproduction are discussed.     

Differences in response to defoliation between males and females of Silene dioica

Summary The response by male and female plants to herbivory was studied by experimental defoliation of the dioecious perennial herb Silene dioica in a green-house. Male and female plants were defoliated prior to and during the early flowering phase at two intensities (50% and 100% of leaf-area removed) in two consecutive years. Defoliation resulted in a decrease in the number of flowers initiated in both sexes, while a larger delay of peak flowering and a higher mortality was observed in males compared to females. In female plants, severe defoliation resulted in a reduction in seed number per capsule and in seed size compared to control. Females showed a negative correlation between the production of flowers in the first and second season in all treatments, while flowering in males the first season was not correlated with flowering in the second season. Females also showed a lower frequency of flowering than males during the two seasons studied. However, during the flowering period, males allocated significantly more biomass to flowers than did females. This outcome supports the idea that females may have a higher total reproductive expenditure than males, but males have a higher reproductive effort during flowering. Male rosette leaves were significantly preferred by the generalist herbivore Arianta arbustorum in experiments. This preference was most pronounced in trials with leaves from fertilized plants compared to nonfertilized plants. A greater storage of resources in aboveground leaves during winter by males compared to females may explain the higher preference for male leaves and the higher male mortality following early defoliation. Furthermore, males are smaller than females and may have a lower ability than females to replace lost resources needed for reproduction when defoliated early in the season.     

Sexual dimorphism in the energetics of reproduction and growth of North Sea plaice, Pleuronectes platessa L.

The chemical composition and energy content of North Sea plaice during the spawning period were examined in mature males and females and in immature fish, to study differences in the allocation of energy over reproduction and somatic growth between the sexes. At the beginning of the spawning period mature males and females had equal dry weights of lipid that were 70% higher than in immatures. Protein content in mature males was equal to that in immatures but was 23 % higher in mature females. Immature males and females did not differ in chemical composition. At the end of the spawning period, spent and immature fish had equal lipid contents, but protein content in spent females was 10% lower than in spent males, and 17% lower than in immatures. Gross energy content of the body decreased by 44% (65·2 to 36·3 J cm^-3) in mature females, 27% (55·0 to 40·OJ cm^-3) in mature males, and 9% (48·7 to 44·2J cm^-3) in immatures. Energy content of plaice eggs was estimated at 6·60 kJ per 1000 eggs. Reproductive investment was estimated from the energy loss during the spawning period and included the energy of sex products and spawning metabolism. Somatic growth comprised the annual increase in energy content of fish. The pattern of energy allocation over reproduction and somatic growth differed between males and females. Males started their reproduction at a smaller length and a younger age and allocated a higher proportion of the available energy into reproduction than females. Available energy resources for somatic growth and reproduction (surplus production) were equal between the sexes up to a length of about 30 cm. Beyond this length male surplus production levelled off whereas female surplus production continued to increase. The differences in surplus production and the allocation patterns are discussed. For female plaice the energy allocated into egg production was estimated as between 48 and 64% of the total amount of energy lost during spawning. The remaining energy is used for metabolism during the spawning period, yielding an estimate of the metabolic rate of mature females of between 6·4 and 9·1 kJ day^-1. A maximum estimate of the metabolic rate of mature males was 7·4 kJ day^-1.     

Sexual and vegetative reproduction in the aboveground part of a dioecious clonal plant, <Emphasis Type="Italic">Dioscorea japonica</Emphasis> (Dioscoreaceae)

In dioecious clonal plants, the reproductive effort required to set seeds will be responsible for the larger investment in sexual reproduction by females. If there will be a trade-off in resource allocation between sexual and clonal reproduction, this differential sexual reproduction will lead to sexual differentiation in the relative amount of clonal reproduction. To test this prediction, we studied differences between the sexes in their phenologies and investments in sexual and vegetative reproduction (clonal reproduction by means of bulbils) with respect to ramet size in a dioecious clonal plant, Dioscorea japonica Thunb. The period of bulbil production overlapped the period during which infructescences developed. Females flowered later, produced heavier inflorescences, and fewer flowers per inflorescence than did males. Regression analysis using the size of the individual plants demonstrated that large females made smaller investments in inflorescences and larger investments in sexual reproduction than did large males. In contrast, females invested fewer resources in vegetative reproduction than did males. However, the total investments in sexual and vegetative reproduction did not differ between the sexes. These results supported our hypothesis on the sexual differentiation in sexual and clonal reproduction.     

Response of the sexes of the subdioecious plant Honckenya peploides to nutrients under different salt spray conditions

Sex dimorphic plants often show sex-specific differences in growth and biomass allocation. These differences have been explained as a consequence of the different reproductive functions performed by the sexes. Such differences may determine the performance of each sex in different habitats and therefore might explain the spatial segregation of the sexes described in many dimorphic plants. We examined the growth, reproductive and physiological responses of the sexes of the subdioecious plant Honckenya peploides to two levels each of salt spray and nutrients, which are assumed to be important selective forces in coastal environments. We found sex-related differences in H. peploides. In particular, females allocated proportionally more dry mass to reproduction and grew less and more slowly than males regardless of salt spray and nutrient conditions, which is interpreted as a trade-off between reproductive and vegetative growth. Regarding physiological response, nutrients significantly increased values of photochemical reflectance index (PRI) in females but not in males, suggesting that photochemical efficiency is more limited by nutrients in females than in males. PRI values also suggest intersexual differences in protection requirements against photoinhibition. The study did not find sex-differential responses to salt spray, which caused a decrease in reproductive effort in both sexes. The sex-related differences in relative growth rates, reproductive allocation and photosynthetic features found here may contribute to explaining habitat-related between-sex differences in performance and, therefore, the spatial segregation of the sexes observed in H. peploides.     

Bud banks of perennial savanna grasses in Botswana

Three semi‐arid savanna grasses in Botswana (Stipagrostis uniplumis, Eragrostis lehmanniana, and Aristida stipitata) were sampled to quantify their belowground bud banks during the dormant season and to estimate their relative allocation to vegetative and sexual reproduction. Bud banks of these African perennial caespitose grasses were also compared with four perennial caespitose grasses of semi‐arid North American grasslands. The three African grasses each maintained approximately two buds per tiller and showed a high percentage (88–99%) of tillers producing seed. Only E. lehmanniana produced new aerial tillers from axillary buds at elevated nodes on the stem as well as from the belowground bud bank. Compared with species of North American grasslands, these African grasses produced fewer belowground buds but showed a much higher percentage of tillers producing seed. These patterns indicate relatively greater belowground meristem limitation, lower allocation to vegetative reproduction (tillering) and higher allocation to seed reproduction in these African grasses, although studies of more species are needed to assess the generality of this pattern. The management of savannas in ways that favour the maintenance of a reserve population of belowground buds may increase the ability of grasses to respond to pulses of resource availability, increase their compensatory growth capacity following grazing or drought, and decrease the invasibility of these plant communities by exotic species, whereas maintaining allocation to sexual reproduction may be important for conserving genetic variation and enhancing their capacity to adapt to environmental change.     

Growth and Investment in Hominin Life History Evolution: Patterns, Processes, and Outcomes

The transitions from apes to lineages allied to humans are marked by shifts in the allocation of parental effort, associated with discontinuous changes in rates of infant and juvenile growth both prenatally and postnatally. Here, I assess growth and life history characteristics of apes within a general mammalian / primate paradigm, using time and energy expenditure as 2 fundamentals that covary with infant survival and success probabilities. I suggest that these survival probabilities depend on the quality, amount, and timing of parental care allocated to infants. Growth to birth, growth to weaning, and growth to reproductive onset are partitioned as separate periods within a life history on the basis of comparative mammalian data. Growth problems such as sexual dimorphism can be incorporated into an investment perspective by assessing when and how sex-specific parental care affects growth rates and the onset of reproduction. I compare features of the hominoid life history with developmental rates for hominin lineages as seen in dentition, and the fossil record of body and brain size changes over time. The links between parental effort and allocation of care to infant growth and survival generate speculative scenarios of sex-specific parental care allocation; I then explore hominin social evolution?—mating system and childhood—?for the lineages thought to lead to modern Homo, and for those that coexisted with ancestors of Homo.     

No differential consequences of reproduction according to sex in Juniperus communis var. depressa (Cupressaceae)

In dioecious plant species, males and females are thought to have dissimilar allocation patterns. Females are believed to invest more in reproduction and less in growth and maintenance than males. This differential investment between sexes could result in distinct growth patterns and contrasting survival rates, thereby affecting the sex ratio of a population and the age and size distribution of males and females, possibly leading to habitat segregation according to sex. These effects might become more apparent under particularly limiting conditions, such as in nutrient-deficient soils or in climatically stressed environments. To verify these predictions, growth patterns, microsite characteristics, and age and size distribution of male and female individuals were compared, and population sex ratio was determined in three populations of the dioecious shrub Juniperus communis var. depressa (Cupressaceae, Pinophyta) along a short latitudinal gradient on the eastern coast of Hudson Bay (Northern Québec, Canada). We found that the northernmost population had a male-biased sex ratio, but that the southernmost one had a higher proportion of females. Our results failed to reveal any significant differences in radial growth patterns, mean sensitivity, annual elongation of the main axis, and size and age frequency distribution between males and females in any population. Furthermore, there was no evidence of microhabitat segregation according to sex as indicated by the lack of differences in the physicochemical characteristics of the substrate under males and females. Clearly, the expected ecological consequences of a presumed greater investment of females in reproduction were not apparent even under the very stressful conditions prevailing on subarctic dunes. Many factors could reduce differences in the cost of reproduction between males and females, such as the number and quality of reproductive structures produced annually by individuals of each sex, the possible photosynthetic activity of the immature female cones, and the complexity of the source/sink relationship within individuals. Alternatively, there may be no differences between sexes in their reproductive investment.     

Sexual differences in biomass and nutrient allocation of first-year Silene dioica plants

Reproductive and somatic biomass, nitrogen (N), and phosphorus (P) pools were compared between females and males in 1st-year plants of Silene dioica. We estimated irretrievable resources allocated to seeds, pollen, flowers, and unrecovered summer leaf investment by collecting plant parts at abscission throughout the season. At the end of the season, we determined resources lost through senescent stems and autumn leaf turnover and resources stored in perennial roots and overwintering buds. Sexual differences in allocation patterns depended on the resource used for comparison, and whether absolute or proportional resource pools were assessed. Total resource pools in terms of biomass and N were similar for females and males. However, male plants acquired relatively more P. The proportional reproductive investment, i.e., reproductive effort, was similar for males and females in terms of biomass and N. In terms of P, male reproductive effort was higher. There was no difference between sexes in the proportional and relative biomass allocated to perennial roots and overwintering buds. However, in terms of absolute and relative N allocation to below-ground parts, females had larger reserves than males. Females, moreover, had a larger proportion of their P in below-ground parts. However, as male total P pools were larger, absolute P reserves did not differ between sexes. The high reproductive effort and N depletion of below-ground parts in males resulted largely from higher flower production compared to females. In females, seeds were the major component of reproductive effort. These results show that if biomass and nutrient allocation are assessed in parallel for dioecious plants, we obtain a more complete view of their sexual differences. Received: 07 May 1998 / Accepted: 30 October 1998     

Sex‐specific reproductive investment of summer spawners of Illex argentinus in the southwest Atlantic

Energy investment in reproduction and somatic growth was investigated for summer spawners of the Argentinean shortfin squid Illex argentinus in the southwest Atlantic Ocean. Sampled squids were examined for morphometry and intensity of feeding behavior associated with reproductive maturation. Residuals generated from length‐weight relationships were analyzed to determine patterns of energy allocation between somatic and reproductive growth. Both females and males showed similar rates of increase for eviscerated body mass and digestive gland mass relative to mantle length, but the rate of increase for total reproductive organ weight relative to mantle length in females was three times that of males. For females, condition of somatic tissues deteriorated until the mature stage, but somatic condition improved after the onset of maturity. In males, there was no correlation between somatic condition and phases of reproductive maturity. Reproductive investment decreased as sexual maturation progressed for both females and males, with the lowest investment occurring at the functionally mature stage. Residual analysis indicated that female reproductive development was at the expense of body muscle growth during the immature and maturing stages, but energy invested in reproduction after onset of maturity was probably met by food intake. However, in males both reproductive maturation and somatic growth proceeded concurrently so that energy allocated to reproduction was related to food intake throughout the process of maturation. For both males and females, there was little evidence of trade‐offs between the digestive gland and reproductive growth, as no significant correlation was found between dorsal mantle length‐digestive gland weight residuals. The role of the digestive gland as an energy reserve for gonadal growth should be reconsidered. Additionally, feeding intensity by both males and females decreased after the onset of sexual maturity, but feeding never stopped completely, even during spawning.     

DOES THE TRADE-OFF BETWEEN GROWTH AND REPRODUCTION SELECT FOR FEMALE-BIASED SEXUAL ALLOCATION IN COSEXUAL PLANTS?

We analyzed sexual allocation in cosexual plants while taking the trade-off between growth and reproduction into consideration and showed that this trade-off does not select for female-biased sexual allocation. There are two problems in sexual allocation: optimizing the amount of resources allocated to reproduction in a growing season and equalizing the resources allocated to the male and the female functions. If these two are possible at the same time, equal resource allocation to the male and the female functions is the evolutionarily stable strategy (ESS; given that the fitness gains through the male and the female functions are proportional to the amount of the resources allocated to these functions). Biased sexual allocation only occurs when constraints make it impossible to simultaneously optimize allocation to reproduction and allocation to male and female functions. However, even if female-biased sexual allocation occurs due to the addition of other constraints, the trade-off between growth and reproduction itself is not an important factor that selects for female-biased sexual allocation.