SECONDARY SEX CHARACTERISTICS IN SPINACIA OLERACEA L.: QUANTITATIVE EVIDENCE FOR THE EXISTENCE OF AT LEAST THREE SEXUAL MORPHS

Spinacia oleracea L. was grown hydroponically across an experimental salinity gradient with treatments of 0.0, 1.5, 3.0, 4.5, and 6.0 grams of NaCl/liter of nutrient medium. Plant height, root biomass, vegetative biomass, fruit morphology, average fruit weight, and fruit mass: volume ratio were examined in each treatment to determine if differences other than primary sex characteristics exist between male, female, and monoecious plants. Average fruit weight was the only parameter examined that was not useful as a secondary sex characteristic. Orthogonal contrasts examining plant height, vegetative biomass, and root biomass of plants were distinct from one another across the salinity gradient. Similarly, female and monoecious plants differed significantly from one another in terms of root biomass and the average mass: volume ratio of the fruit each sexual morph produced, as well as the proportion of smooth to wrinkled fruit produced. The data indicate that male, female, and monoecious plants are three distinct sexual morphs.     

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.     

Responses to moisture stress in male and female plants of Rumex acetosella L. (Polygonaceae)

Summary Male and female plants of Rumex acetosella were grown on a moisture gradient to measure possible differences in the drought tolerance of the sexes. The growth of both sexes declined under water stress but males were significantly more drought tolerant. This could not be explained by greater water use efficiency in the male plants; measured rates of both photosynthesis and leaf conductance did not differ significantly between the sexes. Multiple discriminant analysis showed that the sexes differed at all moisture regimes in their overall patterns of biomass allocation. Males had proportionately greater investment in root and leaf tissue which could explain their growth advantage over females under water stress. Despite essentially equal water use efficiencies, on a per plant basis males, with more leaf and root biomass, could fix more carbon and more rapidly exploit the local water resource than females. Thus the pattern of biomass allocation rather than intrinsic physiological differences appears to explain the greater drought tolerance of male plants of Rumex acetosella.     

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.     

HYDRILLA INVADES WASHINGTON,D.C. AND THE POTOMAC

The aquatic weed hydrilla (Hydrilla verticillata Royle) was discovered growing in the Potomac River, south of Alexandria, VA, in Kenilworth Aquatic Gardens, Washington, D.C., and in the Chesapeake and Ohio (C&O) Canal near Seneca, MD. Cultures in Florida of the Kenilworth clone produced male flowers. This is the first report of the occurrence of the male in the U.S. Two distinct isoenzyme patterns have been identified for plants from various locations in the U.S., corresponding to a monoecious strain and a dioecious female. The occurrence of the wild colonies of the monoecious Hydrilla greatly increase the potential for physiological diversity through sexual reproduction, which may have serious consequences for the management of this weed.     

Sex-specific physiological, allocation and growth responses to water availability in the subdioecious plant Honckenya peploides

The gender of dimorphic plant species is often affected by ecophysiological variables. Differences have been interpreted as a response of the sexes to meet specific resource demands associated with reproduction. This study investigated whether sex‐specific variations in ecophysiological traits in response to water availability determine the performance of each sex in different habitats, and therefore promote extreme spatial segregation of the sexes in the subdioecious plant, Honckenya peploides. Twenty‐seven plants of each sex were individually potted in dune sand and assigned randomly to one of three water treatments. Well‐watered plants were watered daily to field capacity, whereas plants in the moderate and high‐water stress treatments received 40% and 20%, respectively, of the water given to well‐watered plants. Photochemical efficiency, leaf spectral properties and components of relative growth rate (leaf area ratio and net assimilation rate) were measured. Photochemical efficiencies integrated over time were higher in male than in female plants. Water deficit decreased maximum quantum yield in female plants more rapidly than in male plants, but female plants (unlike male plants) had recovered to initial values by the end of the experiment. Maximum quantum yield in male plants was more affected by water stress than in female plants, indicating that male plants were more susceptible to photoinhibition. The two sexes did not differ in growth rate, but male plants invested a higher proportion of their biomass in leaves, had a higher leaf area per unit biomass and lower net assimilation rate relative to female plants. Female plants had a higher water content and succulence than male plants. Differences in stomatal density between the sexes depended on water availability. The results suggest that the two sexes of H. peploides have different strategies for coping with water stress. The study also provides evidence of sex differences in allocation traits. We conclude that between‐sex differences in ecophysiological and allocation traits may contribute to explain habitat‐related between‐sex differences in performance and, therefore, the spatial segregation of the sexes.     

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

Abstract It has been proposed that relative allocation to female function increases with plant size in animal‐pollinated 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.     

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.     

Laboratory growth rates of six species of freshwater Gymnamoebia

Summary Amaranthus and several other wind-pollinated species of plants are used to test some of the theoretical models of relative reproductive effort towards the male and female sexes. Consistent with these models, in self-compatible, monoecious Amaranthus, Chenopodium, Digitaria, Setaria, and Lepidium, female effort represented over 90% of the total reproductive effort. Also consistent with predictions, Lolium, a self-incompatible wind-pollinated species, was found to have about equal male and female effort. A method is described here that should prove useful in quantifying male and female effort in both wind and insect-pollinated species of plants.     

Low siring success of females with an acquired male function illustrates the legacy of sexual dimorphism in constraining the breakdown of dioecy

Dioecy has often broken down in flowering plants, yielding functional hermaphroditism. We reasoned that evolutionary transitions from dioecy to functional hermaphroditism must overcome an inertia of sexual dimorphism, because modified males or females will express the opposite sexual function for which their phenotypes have been optimised. We tested this prediction by assessing the siring success of monoecious individuals of the plant Mercurialis annua with an acquired male function but that are phenotypically still female‐like. We found that pollen dispersed by female‐like monoecious individuals was ~ 1/3 poorer at siring outcrossed offspring than pollen from monoecious individuals with an alternative male‐like inflorescence. We conclude that whereas dioecy might evolve from functional hermaphroditism by conferring upon individuals certain benefits of sexual specialisation, reversion from a strategy of separate sexes to one of combined sexes must overcome constraints imposed by the advantages of sexual dimorphism. The breakdown of dioecy must therefore often be limited to situations in which outcrossing cannot be maintained and where selection favours a capacity for inbreeding by functional hermaphrodites.     

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.     

YEAR-TO-YEAR CHANGES IN PHENOTYPIC GENDER IN A MONOECIOUS CUCURBIT,APODANTHERA UNDULATA

Year-to-year variation in phenotypic gender in the monoecious cucurbit, Apodanthera undulata Gray was investigated. Small plants produce no flowers. Larger plants produce only staminate flowers (“male” plants), while a somewhat greater threshold size is necessary for pistillate flower production (cosexual plants). Approximately 85% of the plants that bloomed did not change gender group between years. Two measures of phenotypic gender were used: prospective femaleness, a measure standardized to the population floral ratio, and morphological femaleness, an unstandardized measure. Femaleness of cosexes between years was positively correlated; r values were somewhat greater when using morphological femaleness values. Plants that opened only staminate flowers one year were likely to open only staminate flowers the next year. Similarly, cosexes were likely to be cosexes again the following year, with similar femaleness values. Beyond the threshold size for pistillate flower production, plant size was not correlated with femaleness. These patterns suggest that all plants are male until they reach a certain size and that plants in their cosexual phase may have an intrinsic femaleness tendency due to either genotype or microsite effects.     

Size-dependent variation of gender in high density stands of the monoecious annual,Ambrosia artemisiifolia (Asteraceae)

Summary We examined the relationship between size variability and the distribution of functional gender in stands of the monoecious, wind-pollinated annual Ambrosia artemisiifolia. Populations of 60 individuals were grown in the greenhouse at a density of 372 m^–2 and at two nutrient levels. Among the surviving plants, after self-thinning, variability in above-ground biomass and gender was higher in the high nutrient treatment. Among individuals there was a significant positive correlation between maleness and both height and biomass. Fecundity was also positively correlated with both measures of size. Based on the pattern of distribution of male and female flowers within the plant, it appears that the increase in maleness in larger plants is due to increased branching and axis elongation. These results demonstrate that competitive interactions, which lead to increased variability in biomass and fecundity, can also generate variability in gender within populations.     

VARIATION IN SEX EXPRESSION IN CANADA YEW (TAXUS CANADENSIS)

Sex expression was measured in several Canada yew (Taxus canadensis Marsh.) populations of the Apostle Islands of Wisconsin and southeastern Minnesota to determine the extent of variation within and among populations. Sex expression was recorded qualitatively (monoecious, male, or female) and quantitatively (by male to female strobilus ratios or standardized phenotypic gender). No discernible trends in differences in sex expression among populations or habitats were recorded. Trends in sex expression of individuals within populations were complex. Small yews tended to be male or, if monoecious, had female-biased strobilus ratios. Large yews were monoecious but had male-biased strobilus ratios. Phenotypic gender, recorded as relative maleness, however, was negatively, but weakly, correlated with plant size. Gender distribution in four of five populations was bimodal, suggesting that cosexual populations consist of male and female morphs. Strobilus ratios of individuals in Apostle Island populations showed significant annual variation, but gender for these same plants was significantly correlated from year to year. Annual adjustments in gender were most pronounced in small yews. The results indicate that relative investment in male and female reproductive structures by Canada yew individuals is responsive to environmental variation, but sex expression also has a proximate genetic component.     

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.     

Environmental conditions affect sex expression in monoecious, but not in male and female plants of Urtica dioica

Urtica dioica is a sub-dioecious plant species, i.e. males and females coexist with monoecious individuals. Under standard conditions, seed sex ratio (SSR, fraction of males) was found to vary significantly among seed samples collected from female plants originating from the same population (0.05–0.76). As a first step, we investigated the extent to which SSR and sex expression of male, female, and monoecious individuals is influenced by external factors. We performed experiments to analyse: (1) whether the environment of a parental plant affects the sex ratio (SR) of its offspring, (2) whether SSR can be affected by environmental conditions before flowering, and (3) whether sex expression of male, female and monoecious plants that have already flowered can be modified by environmental conditions or by application of phyto-hormones. Within the range of our experimental design, SSR was not influenced by external factors, and gender in male and female plants was stable. However, sex expression in monoecious plants was found to be labile: flower sex ratio (FSR, fraction of male flowers) differed considerably between clones from the same individual within treatments, and increased toward 100% maleness under benign conditions. These results provide strong evidence that monoecious individuals are inconstant males, which alter FSR according to environmental circumstances. In contrast, we consider sex expression in male and female individuals to be solely genetically based. The observed variation in SSR between maternal parents cannot be explained by sex-by-environment interactions.     

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.     

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.     

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     

浑善达克沙地无芒雀麦（Bromus inermis）空间分布格局

通过西北-东南和东北-西南方向两条宽5m×长3 km的样带和一块4m×4m的样方调查,研究了浑善达克沙地无芒雀麦、植被、土壤水分及土壤盐分的空间变异特点及其相互关系。结果表明,样带内植被盖度、无芒雀麦盖度、土壤水分和土壤盐分的空间变化格局相似,然而,在不同的空间方向上,它们具有不同的空间变化;样方内无芒雀麦地上部生物量、分株数和土壤盐分具有相似的空间变化格局,但却不同于植被地上部总生物量和土壤水分的空间变化格局。相关分析表明,样带内植被盖度、无芒雀麦盖度与土壤水分和土壤盐分之间均具有显著的正相关关系。土壤水分和土壤盐分也具有显著的正相关。样方内无芒雀麦地上部生物量、分株数及植被地上部总生物量与土壤水分均具有显著的正相关关系,但与土壤盐分没有显著的相关关系。无芒雀麦地上部生物量、分株数和植被地上部总生物量三者之间,以及土壤水分和土壤盐分之间也均具有显著的正相关关系。研究结果表明了植被与土壤之间的相互关系依赖于空间尺度。