PATTERNS OF BIOMASS ALLOCATION IN SPINACIA OLERACEA (CHENOPODIACEAE) ACROSS A SALINITY GRADIENT: EVIDENCE FOR A NICHE SEPARATION

Spinacia oleracea var. Americana was hydroponically grown across an experimental salinity gradient ranging from 0.0 to 6.0 grams NaCl/liter nutrient medium. Twenty male, 20 female, and 20 monoecious plants were randomly harvested from each treatment where possible. Root biomass (RTA), biomass allocated to leaves and stems (VA), and reproductive biomass (RA) were determined. Male, female, and monoecious plants were shown to have significantly different patterns of RTA, VA, and RA. Moreover, the treatments influenced allocation patterns of the sexes differently, i.e., a partial niche separation was observed among the sexes along the gradient with respect to RTA, VA, and RA. Additionaly, the sexes expressed different trade-offs between allocation of resources to the different tissues; most notably, under high salinity, RTA, and RA were strongly negatively correlated in females, but positively correlated in male and monoecious plants.     

Trade-offs between clonal and sexual reproduction in Sagittaria latifolia (Alismataceae) scale up to affect the fitness of entire clones

? Many plants combine sexual reproduction with vegetative propagation, but how trade-offs between these reproductive modes affect fitness is poorly understood. Although such trade-offs have been demonstrated at the level of individual shoots (ramets), there is little evidence that they scale up to affect genet fitness. For hermaphrodites, reproductive investment is further divided between female and male sexual functions. Female function should generally incur greater carbon costs than male function, which might involve greater nitrogen (N) costs. ? Using a common garden experiment with diclinous, clonal Sagittaria latifolia we manipulated investment in reproduction through female and male sex functions of 412 plants from monoecious and dioecious populations. ? We detected a 1?:?1 trade-off between biomass investment in female function and clonal reproduction. For male function, there was no apparent trade-off between clonal and sexual reproduction in terms of biomass investment. Instead, male function incurred a substantially higher N cost. ? Our results indicate that: trade-offs between investment in clonal propagation and sexual reproduction occur at the genet level in S.?latifolia; and sexual reproduction interferes with clonal expansion, with investment in female function limiting the quantity of clonal propagules produced, and investment in male function limiting the nutrient content of clonal propagules.     

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.     

SIZE-DEPENDENT GENDER CHANGE IN GREEN DRAGON (ARISAEMA DRACONTIUM; ARACEAE)

Green dragon (Arisaema dracontium; Araceae) is a perennial woodland herb capable of switching gender from year to year. Small flowering plants produce only male flowers but when larger they produce male and female flowers simultaneously. Distinct male and monoecious phenotypes (referred to hereafter as plants) share a single underlying cosexual genotype. Four populations in southern Louisiana were sampled to determine frequencies and size distributions of male and monoecious plants, and to determine the relationship of plant size with male and female flower production in monoecious plants. Male plants were significantly smaller than monoecious plants and made up 34%–78% of flowering plants within populations. Flower number (average = 120) was weakly positively correlated with size. Monoecious plants produced an average of 169 flowers (90 female) and had 100% fruit set, with individual berries containing an average of 2.5 ovules and 1.3 filled seeds. Male flower number was negatively correlated, and female flower number positively correlated, with basal stem diameter. Extrapolation of regression slopes suggested that green dragon should become completely female at a size 20% larger than the largest plant observed in this study. A simple model of inflorescence development is presented to illustrate how the reproductive system of green dragon is related to that of jack-in-the-pulpit (A. tnphyllum), which exhibits a more distinct switch between male and female phenotypes.     

Flowering phenology and sexual allocation in single-mutation lineages of Arabidopsis thaliana

We investigated the relationship between flowering time and sexual allocation in wild-type Arabidopsis thaliana and in genetically similar lineages with single-locus mutations of floral induction genes. We examined whether the mechanisms of growth and development that govern resource investment would permit the independent evolution of reproductive phenology and sexual allocation, or whether constraints, manifested as pleiotropic effects of the single mutations, would link these two life-history traits. Flowering times differed significantly among genotypes, and, as expected, later flowering times were associated with larger vegetative size. Later flowering genotypes produced heavier floral parts (larger petals, in particular), and allocated a significantly lower proportion of biomass to androecia, especially in final allocations that included fruit biomass. At least part of this pleiotropic covariation of flowering time and sexual allocation is likely to be mediated by vegetative size and the rate of resource supply to growing reproductive tissues, because the larger fruits of late-flowering genotypes required the same time, or proportionately less time than the difference in biomass, to mature. Because fruit mass is considered an investment in female function, sexual allocation measured at the end of a growing season tends to be highly female biased in angiosperms. We consider the implications of the pleiotropic association of flowering time, vegetative size, and sexual investment for the theory of sex allocation, and suggest that the idiosyncratic phenology of sexual investment in flowering plants creates a departure from a central assumption of Fisher's seminal sex allocation argument.     

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.     

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.     

Complex sex determination in the stinging nettle Urtica dioica

The dioecious species Urtica dioica harbours wide variation in sex ratio of seeds. We conducted a series of crosses to analyse the genetic basis of sex determination in this species. Dutch populations of U. dioica contain low proportions of monoecious individuals beside male and female plants. Self-pollination of monoecious plants always yielded female, male and monoecious plants, generally in a ratio of one female to three male/monoecious individuals. This motivated us to write down a simple model in which gender is determined by one major sex-determination locus with four alleles. In the model males and monoecious plants have distinct genotypes but are both heterozygous at the sex-determination locus. We first made crosses among progeny obtained after self-pollination of monoecious plants. These crosses showed that the monoecious trait generally showed Mendelian inheritance and was passed on to the next generation via both pollen and seeds. Further crosses between monoecious plants and plants from dioecious system indicated that alleles from the dioecious system are often dominant. However, many exceptions to our genetic model are observed which suggest that dominance is incomplete and/or that more genes are involved in sex determination. We discuss to what extent sex determination genes explain the strongly biased seed sex ratios and argue that additional genes, for instance genes for female choice, must also be involved.     

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

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

Size advantage for male function and size‐dependent sex allocation in Ambrosia artemisiifolia,a wind‐pollinated plant

In wind‐pollinated plants, male‐biased sex allocation is often positively associated with plant size and height. However, effects of size (biomass or reproductive investment) and height were not separated in most previous studies. Here, using experimental populations of monoecious plants, Ambrosia altemisiifolia, we examined (1) how male and female reproductive investments (MRI and FRI) change with biomass and height, (2) how MRI and height affect male reproductive success (MRS) and pollen dispersal, and (3) how height affects seed production. Pollen dispersal kernel and selection gradients on MRS were estimated by 2,102 seeds using six microsatellite markers. First, MRI increased with height, but FRI did not, suggesting that sex allocation is more male‐biased with increasing plant height. On the other hand, both MRI and FRI increased with biomass but often more greatly for FRI, and consequently, sex allocation was often female‐biased with biomass. Second, MRS increased with both height and MRI, the latter having the same or larger effect on MRS. Estimated pollen dispersal kernel was fat‐tailed, with the maximum distance between mates tending to increase with MRI but not with height. Third, the number of seeds did not increase with height. Those findings showed that the male‐biased sex allocation in taller plants of A. artemisiifolia is explained by a direct effect of height on MRS.     

Sex Allocation Ratio in a Wind-Pollinated Self-Incompatible Monoecious Tree, Alnus firma Sieb. et Zucc. (Betulaceae)

Abstract Biomass sex allocation of Alnus firma , a wind-pollinated self-incompatible monoecious tree species, was directly measured at the inflorescence (prezygotic) stage. Sex allocation ratio at the fruit stage was estimated from the number of female inflorescences, percent fruit production, and the mean dry weight of a female inflorescence and a fruit catkin. Investment in male function far exceeds that in female function at the prezygotic stage. When the costs of fruit catkins and seeds were included in the estimation of female function, then the sex allocation bias was reversed towards female. Intrapopulational variation of the sex allocation ratio with respect to tree height and size was also investigated. A strong negative correlation was observed in both cases, although a positive relationship has been predicted by theorists and some previous empirical data have supported this prediction.     

Tests of a mechanistic model of one hormone regulating both sexes in Buchloe dactyloides (Poaceae)

A mechanistic model of one hormone regulating both sexes in flowering plants was tested in buffalograss (Buchloe dactyloides). This model assumes that one hormone has male and female cell receptors to inhibit one sex and induce the other independently. Three components—the normal range of hormone level in the plant and the sensitivity levels of the two receptors—interact to regulate sex expression. The study organism, buffalograss, is usually considered dioecious, but natural populations consist of varying proportions of male, female, and monoecious plants. Prior research with growth regulators had shown that only gibberellin (GA) had consistent and significant effects on sex expression in this species. To test the model assumption of a hormone with a dual function, GA and a GA inhibitor (paclobutrazol, PAC) were applied to three monoecious genotypes; in two of the genotypes the GA treatment yielded a significantly higher proportion of male inflorescences, and this transition involved both inducing male and inhibiting female. PAC treatment produced exclusively female inflorescences, illustrating the dual effects of GA. To test the predictability of the model, GA was applied to two dwarf female genotypes. These plants were transformed into neuter and near-neuter plants with normal height and vegetative growth, as predicted by our model for genotypes with a physiologically wide overlapping of male and female sterile regions. The model also predicts that male or female plants would be induced to produce inflorescences of the other sex if the hormone level could be shifted from one side of the overlapping sterile regions to the other. This was verified by applying high levels of GA to a normal female genotype that resulted in the production of male inflorescences. However, this is the only normal female that has responded to GA application by producing male inflorescences, and males lose vigor and/or die without producing female inflorescences at high levels of PAC. The model suggests that the constancy of these males and females is due to the relative location of the sensitivity levels in relation to each other and to the hormone range. We conclude that the one-hormone model can facilitate both applied and basic research.     

Gender plasticity in Sagittaria sagittifolia (Alismataceae), a monoecious aquatic species

 Aquatic plants are well known for their high degree of phenotypic plasticity in vegetative structures, particularly leaves. Less well understood is the extent to which their sexuality can be modified by environmental conditions. Here we investigate gender plasticity in the European clonal monoecious aquatic Sagittaria sagittifolia (Alismataceae) to determine how floral sex ratios may vary with plant size and inflorescence order. We sampled two populations from aquatic habitats in East Anglia, U.K. and measured a range of plant attributes including ramet size and the number of female and male flowers per inflorescence. The two populations exhibited similar patterns of phenotypic gender, despite contrasting patterns of total allocation to female and male flower number. Plants produced male-biased floral sex ratios but female flower number increased from the first to the second inflorescence whereas male flower number decreased. Size-dependent gender modification occurred in both populations, but the patterns of allocation to female flower production differed between the two populations. Our results are consistent with the view that monoecy is a sexual strategy that enables plants to adjust female and male allocation in response to changing environmental conditions. Received September 16, 2002; accepted October 23, 2002 Published online: March 20, 2003     

FECUNDITY AND SIZE RELATIONSHIPS IN JACK-IN-THE-PULPIT,ARISAEMA TRIPHYLLUM (ARACEAE)

Arisaema triphyllum is a gender-labile woodland herb in which sex expression is correlated with the abundance of stored resources. Larger plants are female or monoecious, smaller ones are male. Among females larger plants produce more flowers, fruits and seeds, and the rate of successful fruit and seed formation is greater for plants of greater ht and corm diam. Average seed wt is greater in larger plants. Seed number per fruit and average seed wt per fruit taper towards the top of the infructescence. Pollinator limitation and resource supply may both contribute to the regulation of yield; their effects can be interpreted sequentially.     

Changes in Sexual Expression as Result of Defoliation and Environment in a Monoecious Shrub in Mexico: Implications for Pollination

Folivory may indirectly impact plant reproduction through changes in sexual expression ( i.e. , number or proportion of male and female flowers produced), which influence plant–pollinator interactions via changes in pollinator preference or efficiency. This study is an experimental evaluation of the effect that defoliation has on sex expression in the monoecious shrub Cnidoscolus aconitifolius , how such effect varies across sites, as well as how such changes indirectly affect pollinator visitation rates. The present study used three populations of C. aconitifolius , each one located in a different site in Yucatán (México): pasture, deciduous forest and subdeciduous medium height forest and three levels of defoliation: 50 percent, 100 percent, and a control (no damage). Results showed that defoliation reduced significantly the total number of male flowers produced in two of the sites. Defoliation did not impact female flower production or the proportion of female flowers produced. Finally, floral visit rates were not affected by defoliation via changes in sexual expression and neither by site or by the interaction site × defoliation. Findings showed that defoliation had an effect on sex expression in C. aconitifolius , although apparently this change did not affect the plant–pollinator interactions.     

LIFE-HISTORY STRATEGIES AND SEX RATIOS FOR A CULTIVAR AND A WILD POPULATION OF BUCHLOE DACTYLOIDES (GRAMINEAE)

Buchloe dactyloides (Nutt.) Engelm. is a perennial, sod-forming shortgrass of the grasslands of central North America from Montana to northern Mexico. The objectives of this research were to compare a cultivar (Sharps Improved) and a wild population from the Oklahoma panhandle, both in the field and in the greenhouse over several growing seasons, in regard to life-history strategies, sex ratios, and constancy of sex expression. Sex ratios of offspring from individual burs (usually 1–5 seeds per bur) and for all surviving plants germinated from seeds in the greenhouse were markedly different for the two populations. The wild population exhibited a 1: 1 male/female ratio with no monoecious plants and complete stability in sex expression over 3 yr of study in the greenhouse. In contrast, 13% of the cultivar plants were monoecious, and its unisexual plants showed a female bias. Burs of the two populations, in flats exposed to three separated month-long periods of favorable moisture with intervening “drought” periods, showed differential germinability of seeds within burs. The cultivar had less dormancy and produced a greater number of seedlings in the first and second periods; wild burs significantly exceeded cultivar burs in the number of seedlings during the third period and in the number of burs giving rise to seedlings in all three periods. Field and greenhouse measurements of vegetative (VRE) and sexual reproductive effort (SRE) per plant indicated that SRE was minimal (ca. 2% of aboveground biomass) in relation to asexual ramet propagation by rapidly growing stolons (VRE = ca. 80%). Mean SRE and VRE were not only similar for the populations, but also for the males and females within a population, although the values for individual plants were highly variable. It is concluded that Buchloe dactyloides is largely dioecious, with at least some wild populations showing a 1: 1 male/female ratio and constancy of sex expression, and that the bur as a dispersal unit containing 1–5 seeds is particularly appropriate for the breeding system and habitat ecology of Buchloe dactyloides.     

EFFECT OF DENSITY ON SECONDARY SEX CHARACTERISTICS AND SEX RATIO IN SILENE ALBA (CARYOPHYLLACEAE)

A field survey of plant and flower sex ratio and secondary sex characteristics was made in Silene alba. Female-biased plant sex ratios were found, as seems typical for the species. Sex ratio distribution correlated with a gradient of soil moisture (with the more moist area having a more female-biased ratio) and with changes in the density of Silene (intermediate and higher density areas having greater female bias). The floral sex ratio was significantly female-biased only at the site that was most female-biased in terms of plant sex ratio. Otherwise the population of flowers was significantly male-biased. Male and female plants harvested from the field differed in secondary sexual characteristics. Males had more flowers and invested proportionately more biomass in leaf, but less in root, stem and reproductive tissue than did females. Although both males and females were larger in terms of total dry weight at the moist site, males produced more flowers at the driest (high density) site. Here the female bias in plant sex ratio was intermediate, but the floral sex ratio was significantly male-biased. A glasshouse experiment was performed in which plants were grown at four densities. Density significantly influenced plant survivorship and the probability of flowering, and increased female bias in the pots, but it did not affect patterns of biomass allocation in flowering plants. Patterns of male and female biomass allocation did not differ in the experiment, except in terms of reproductive allocation (greater in females) and allocation to leaf, greater in males, but only at the lowest density. This work urges caution in interpreting differences between males and females in the field as secondary sex characteristics, since we find such properties to be overlapping under experimental conditions. It supports the idea that males and females of a species may sustain different reproductive output under differing conditions.     

Sex-ratio variation among Arisaema species with different patterns of gender diphasy

Arisaema species exhibit gender diphasy, or sex change, where individual plants produce either male, monoecious or female inflorescences depending on their size. Three basic sex-change patterns have been described in Arisaema. Type I species change between male and monoecious phases, type II species change between male, monoecious and female phases, while type III species change between male and female phases. Theoretical models suggest that sex ratios should be biased toward males, the sex with the lowest cost of reproduction. The goal of this study was to examine sex-ratio variation among Arisaema species that differ in sex-change patterns. Data from an extensive literature review, consisting of all available studies reporting Arisaema sex ratios, were combined with data from extensive field surveys of Arisaema dracontium and Arisaema triphyllum in southern Indiana, USA. This data set contains nearly 30 000 plants from 12 species. All species conformed to either the type I or type III pattern of sex change. There was little evidence for a distinct type II pattern of sex change, given that plants with monoecious inflorescences were rare relative to plants with pistillate inflorescences. The mean sex ratio in type I species (79.9% male) was significantly greater than in type III species (63.7% male). The data are consistent with the prediction that type I species are likely to have greater costs associated with female reproduction. We suggest that all Arisaema species have similar patterns of floral development, but differ in their ontogenetic patterns for male and female flowering.     

Life‐history trade‐offs promote the evolution of dioecy

Most dioecious plants are perennial and subject to trade‐offs between sexual reproduction and vegetative performance. However, these broader life‐history trade‐offs have not usually been incorporated into theoretical analyses of the evolution of separate sexes. One such analysis has indicated that hermaphroditism is favoured over unisexuality when female and male sex functions involve the allocation of nonoverlapping types of resources to each sex function (e.g. allocations of carbon to female function vs. allocations of nitrogen to male function). However, some dioecious plants appear to conform to this pattern of resource allocation, with different resource types allocated to female vs. male sex functions. Using an evolutionarily stable strategy approach, we show that life‐history trade‐offs between sexual reproduction and vegetative performance enable the evolution of unisexual phenotypes even when there are no direct resource‐based trade‐offs between female and male sex functions. This result might help explain the preponderance of perennial life histories among dioecious plants and why many dioecious plants with annual life histories have indeterminate growth with ongoing trade‐offs between sexual reproduction and vegetative growth.     

Production of male flowers does not decrease with plant size in insect-pollinated Sagittaria trifolia, contrary to predictions of size-dependent sex allocation

It has been proposed that relative allocation to female function increases with plant size in animalpollinated species.Previous investigations in several monoecious Sagittaria species seem to run contrary to the prediction of size-dependent sex allocation (SDS),throwing doubt on the generalization of SDS.Plant size,phenotypic gender,and flower production were measured in experimental populations of an aquatic,insect-pollinated herb Sagittaria trifolia (Alismataceae) under highly different densities.The comparison of ramets produced clonally can reduce confounding effects from genetic and environmental factors.In the high-density population,48％ of ramets were male without female flowers,but in the low-density population all ramets were monoecious.We observed allometric growth in reproductive allocation with ramet size,as evident in biomass of reproductive structures and number of flowers.However,within both populations female and male flower production were isometric with ramet size,in contrast to an allometric growth in femaleness as predicted by SDS.Phenotypic gender was not related to ramet size in either population.The results indicated that large plants may increase both female and male function even in animal-pollinated plants,pointing towards further studies to test the hypothesis of size-dependent sex allocation using different allocation currencies.