RELATIONSHIP BETWEEN SYNAPTOSPERMY AND DIOECY IN THE LIFE HISTORY STRATEGIES OF BUCHLOE DACTYLOIDES (GRAMINEAE)

Buchloe dactyloides (Nutt.) Engelm. is a perennial dioecious grass, exhibiting synaptospermy in that the 1–5 seeds in the dispersal unit (bur) are not released prior to germination. A major positive effect is that at least 50% of the burs contain both a male and female, making sexual reproduction possible and reducing founder effects upon colonization by one or a few burs. The objectives of this research were to examine the relationships between synaptospermy and dioecy in the life history strategies of Buchloe dactyloides, and to investigate whether the potential costs of synaptospermy would exceed its benefits if the species were not dioecious. An experiment that tested the effects of tightly clumping three seedlings vs. a uniform spacing of single seedlings at two densities and two moisture levels indicated only slightly negative effects of this aspect of synaptospermy. Although number of tillers and shoot biomass per plant were lower for clumped seedlings at all density and moisture combinations, only 50% of the possible uniform vs. clumped comparisons were significantly different. In addition, these effects, even at low moisture and/or high density, were not great enough to affect survivorship during the 16-week growth period. Experiments under both optimal and low moisture conditions showed that seeds within burs germinated at a significantly higher percentage than isolated or “hulled” seeds; in contrast to the differential timing of germination from burs, almost all of the seedlings from hulled seeds were produced during the first of three germination periods. Although survivorship for these seedlings was quite high under optimal conditions, it was greatly reduced under low moisture, being only one-fourth of that for seedlings arising from burs. The possible reduction in the wind dispersal of propagules due to their teeth and to their greater weight and size appears offset by more effective ungulate and water dispersal. Considering the evidence that the bur anchors the seedling in the soil, protects the seeds from fire, reduces precocious germination by requiring considerable moisture for saturation, and enhances seed longevity, the benefits of synaptospermy appear to outweigh the costs, even if those benefits related to dioecy are not considered.     

A MACROMUTATION IN TRIPSACUM DACTYLOIDES (POACEAE): CONSEQUENCES FOR SEED SIZE,GERMINATION, AND SEEDLING ESTABLISHMENT

A recessive allele of a gene in Tripsacum dactyloides L. (eastern gamagrass) changes staminate florets to pistillate or hermaphrodite, and restores fertility to suppressed florets. There were ten to 25 times more seeds in the mutant pistillate form, and these were 0.32 to 0.59 times smaller than seeds from the normal form. Seeds from pistillate plants had significantly lower germination rates (22% vs. 50%), and seedlings grew 20% slower than those of normal plants in a greenhouse experiment. Pistillate seedling survival rates were lower in both high- (18.8% vs. 62.6%) and low- (52.8% vs. 72.6%) competition environments in a field experiment, and surviving seedlings were smaller. The maternal parent of volunteer seedlings next to a plantation of normal and pistillate plants was determined by dissecting the attached fruitcases of 1,313 seedlings. Pistillate plants in the plantation produced 90% of all seeds falling on the site but only 29% of the volunteer seedlings. The pistillate macromutation is not likely to spread in the wild due to morphological constraints on seed size and packaging.     

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.     

POLLEN-MEDIATED GENE FLOW IN CUCURBITA FOETIDISSIMA (CUCURBITACEAE)

Pollen-mediated gene flow along a linear series of patches of the functionally gynodioecious, bee-pollinated Cucurbita foetidissima was assessed using electrophoretic analysis of seed allozymes. Gene flow was documented at distances up to 0.7 km. For the 13 patches examined, interpatch pollen must have sired from 0% to 48.3% (X̄ = 8.5%) of the seeds of monoecious plants (hermaphrodites). Rates of interpatch siring of seeds of pistillate plants (females) averaged 20.4% (range 8.6%-40%) for the three patches examined. Heterogeneity among fruits in seeds sired by interpatch pollen indicates that the arrival of interpatch pollen is clumped with respect to stigmas. Within patches, plants of the same sex type usually shared identical five-locus genotypes, suggesting that clonal propagation predominates. Since approximately 90% of seeds are sired by intrapatch pollen, seeds of monoecious plants appear to result primarily from geitonogamous (self-) fertilization. This may help explain the existence of female plants in natural populations, since self-fertilization has been shown to severely reduce the survival of seedlings in this species.     

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.     

MORPHOLOGY,INHERITANCE, AND EVOLUTIONARY SIGNIFICANCE OF SEX REVERSAL IN TRIPSACUM DACTYLOIDES (POACEAE)

Inflorescences of Tripsacum dactyloides (L.) L. (Andropogoneae) are characterized by single female spikelets at one to several basal nodes and paired male spikelets at several nodes above them on each raceme. Female spikelets are one-flowered and male spikelets are two-flowered. A sex form variant was found in a wild population in north central Kansas and classified as T. dactyloides (L.) L. forma prolificum Dayton et Dewald. The variant of this native distant relative of maize (Zea mays L. spp. mays) differs from the normal form by having both pistillate and perfect rather than staminate spikelets in the terminal (tassel) portion of the inflorescence and by having two functional pistillate florets in the basal spikelets instead of one. A recessive major gene at a single locus regulates the change of the inflorescence from monoecious to gynomonoecious.     

EFFECTS OF LIGHT AND DENSITY ON RESOURCE ALLOCATION IN A FOREST HERB,ASTER ACUMINATUS (COMPOSITAE)

Recent studies show that forest herbs have low sexual reproductive efforts (SRE) and high vegetative reproductive efforts (VRE) and that a variety of factors may affect the levels of reproduction and the distribution patterns of forest species. Aster acuminatus is a micro-successional forest herb common in northern New England. Resource allocation patterns were determined for seven aster patches varying in density and light level. Plant density affects neither plant size nor any aspect of resource allocation, and VRE remains constant over all density and light levels. Light level is, however, significantly correlated with patch location, average plant size and average patch SRE. Only large plants produce flowers suggesting that light affects sexual reproduction indirectly through effects on plant size. The high rhizome and low seed survivorship of this species may partly explain the constant VRE and varying SRE levels, although the resource allocation patterns observed do not correspond well with the predictions of certain theoretical models.     

THE RELATIVE IMPORTANCE OF INBREEDING AND MATERNAL SEX IN DETERMINING PROGENY FITNESS IN SIDALCEA OREGANA SSP. SPICATA,A GYNODIOECIOUS PLANT

Gynodioecious plant populations contain both hermaphrodite and female individuals. For females to be maintained they must compensate for their loss of reproductive fitness through pollen. Females may achieve compensation by producing more and/or higher quality seeds than hermaphrodites. In this study, I investigated the independent and interactive effects of maternal sexual identity and inbreeding level on fitness of the progeny of hermaphrodites and females of Sidalcea oregana ssp.spicata. Seeds produced by selling hermaphrodites and by outcrossing or sib-crossing hermaphrodites and females, were planted in the field and greenhouse. Maternal-sex effects were substantial at the juvenile stages of the life cycle; seeds of females germinated in higher proportions and produced seedlings that grew significantly faster. Inbreeding effects were manifested primarily at the adult stage of the life cycle. Outcrossed plants were significantly larger and produced more flowers per plant than sib-crossed and selfed plants growing in the greenhouse. Progeny of hermaphrodites and females appeared to respond similarly to sib-matings. The maternal-sex effects observed in Sidalcea may have been related to cytoplasmically inherited factors and could be a driving force in the maintenance of females. Inbreeding depression could play a role in determining the fitness of both sex morphs, if females experience biparental inbreeding in the field. Frequent inbreeding of hermaphrodites may not be necessary to explain the maintenance of gynodioecy in this species.     

EVOLUTION OF DIOECY IN BUCHLOE DACTYLOIDES (GRAMINEAE): TESTS FOR SEX-SPECIFIC VEGETATIVE CHARACTERS,ECOLOGICAL DIFFERENCES,AND SEXUAL NICHE-PARTITIONING

Buchloe dactyloides is a perennial dioecious grass in which male and female inflorescences are so strikingly dimorphic that they were originally assigned to different genera. The objective of this paper is to present the results of tests for sex-specific vegetative characters, ecological differences, and sexual niche-partitioning, combining them with prior information on the reproductive biology of Buchloe for an evaluation of the key factors leading to the evolution of dioecy and sexual dimorphism. Field and greenhouse data were collected from Oklahoma and Kansas populations on vegetative characters, allocation to reproduction, and relative growth and competitive success along resource gradients. Except for greater susceptibility to leaf rust by males, there were no significant differences between males and females in vegetative characters, total biomass, or reproductive effort. Field studies of spatial distributions of males and females failed to show any relation to soil, topography, or soil moisture. In a 45-month greenhouse experiment starting at the seedling stage, the relative growth and competitive success of randomly paired individuals showed no evidence for differential competitive success or for niche-partitioning of males and females. The “outcrossing advantage” and subsequent sexual specialization of the female inflorescence appear to be the major factors underlying this dimorphic system.     

Testing the cultivar vigor hypothesis: comparisons of the competitive ability of wild and cultivated populations of Pascopyrum smithii along a restoration chronosequence

Restoration practitioners often rely on seeds of widely available cultivars representing native species but nonlocal germplasm. Cultivation improves the supply of plant materials and minimizes revegetation costs, but can also favor agronomic traits, and resulting vigor may affect the competitive ability and long‐term persistence of cultivated genotypes at restoration sites. We compared cultivated, restored, and wild populations of Pascopyrum smithii (western wheatgrass) in a greenhouse study to test the extent to which cultivars outcompete local plants in biomass production, and to determine if morphological differences (including height and number of leaves) among cultivated and wild populations persist at restoration sites over time. We found evidence of vigor and greater competitive ability of cultivars in seed mass, growth rate, plant height, and biomass and this advantage occurred when plants were grown alone or in competition with other seed sources. Cultivar vigor persisted at restoration sites over 30 years, but restored populations more closely resembled wild, local populations when cultivars were planted in closer proximity to nearby undisturbed sites. This study supports the cultivar vigor hypothesis and provides evidence for the long‐term persistence of cultivated traits in the environment.     

The influence of deer browsing on the reproductive biology of Canada yew (Taxus canadensis marsh.)

Summary Browsed Canada yew (Taxus canadensis) populations have a higher proportion of males and a lower proportion of monoecious plants than unbrowsed yew populations. The proportion of monoecious plants increases with time following protection from browsing suggesting that deer browsing causes male-biased sex expression in Canada yew. In contrast, results from comparing browsed and unbrowsed populations, exclosure studies, and browse simulation experiments indicate that strobilus ratios and phenotypic gender of browsed yews may be female-biased. In part, these results correspond to the influence of size on sex expression in Canada yew; small yews tend to be male, but if monoecious, have female-biased strobilus ratios. Large yews are monoecious, but have male-biased strobilus ratios. There is, however, no consistent relationship between size and gender in Canada yew, suggesting that in some circumstances, yews shift allocation to female function in response to browsing.     

INFLUENCE OF ENVIRONMENT AND POPULATION ORIGIN ON SURVIVORSHIP AND REPRODUCTION IN RECIPROCAL TRANSPLANTS OF AMPHICARPIC PEANUTGRASS (AMPHICARPUM PURSHII)

Reciprocal transplants of both seeds and seedlings were utilized to determine whether populations of the annual grass Amphicarpum purshii have become locally adapted to specific habitats due to the consistent production of cleistogamous subterranean seeds from year to year. The hypothesis was that subterranean seeds placed in the same habitat as the parents will produce seedlings of greater vigor and adults of higher reproductive capacity than plants from seeds transplanted to a different habitat far removed from the parents. For both seed and seedling transplant experiments involving three sites in the Pine Barrens of New Jersey, the effects of site on shoot dry weight and production of aerial spikelets, subterranean spikelets, and seeds were generally much more significant than the effects of population origin. With one exception, there was no tendency for seedlings (or plants from seeds) replanted into their home sites to outperform alien seedlings (or plants from seeds) transplanted into these same sites. The overriding importance of environmental factors (relative to genetic differences among populations) in determining the phenotypic expression of life history characters, and selection occurring during succession at a site may retard the evolution of genetic adaptation to local habitat conditions in this species.     

Genetic adaptations to grazing and mowing in the unpalatable grass Cenchrus incertus

Summary To identify morphological and life history adaptations to grazing, mowing, and cultivation, seeds of the grass Cenchrus incertus were collected from two populations in each of three types of sites: cemeteries (mown occasionally), pastures (grazed continuously), and orchards (plowed twice a year). Seeds from each population were germinated and grown in a common greenhouse.Plants originating from the two cemetery populations had, on average, the most leaves and the most tillers per plant at each census, and they were on average the shortest in stature. Cemetery plants had on average the greatest number of panicles and of burs per plant, but the fewest burs per panicle. The occasionally-mown but ungrazed cemetery populations in this study were therefore more similar to grazed populations described in other studies; the pasture and orchard populations in this study were more similar to ungrazed populations described in other studies. We suggest that this may be due to the low acceptability of Cenchrus incertus, which makes its defoliation relatively infrequent in unmown sites.Some of the traits that distinguished the cemetery populations from the orchard and pasture populations, such as shorter stature, are probably direct adaptations to defoliation. Others may be secondary effects of these, or the result of allocation trade-offs.     

Effect of Tylenchorhynchus robustoides on Growth of Buffalo Grass and Western Wheatgrass

Tylenchorhynchus robustoides reduced (P = 0.05) growth of Agropyron smithii (western wheatgrass) at soil temperatures of 20, 25, 30, and 35 C. Growth reduction increased with increasing soil temperatures. Highest populations of T. robustoides were recovered at 25 and 30 C. Clipping weights of Buchloe dactyloides (buffalo grass) were reduced at 25 and 30 C; however, root/crown weights were reduced at 15, 20, 30, and 35 C in nematode infested vs. noninfested soil. Reproduction of T. robustoides was greater at 25, 30, and 35 C than at 20 C on B. dactyloides. In a greenhouse study, T. robustoides reduced clipping and root/crown weights of both grasses 24-64%.     

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.     

Rock-degrading endophytic bacteria in cacti

A plant–bacterium association of the cardon cactus (Pachycereus pringlei) and endophytic bacteria promotes establishment of seedlings and growth on igneous rocks without soil. These bacteria weather several rock types and minerals, unbind significant amounts of useful minerals for plants from the rocks, fix in vitro N₂, produce volatile and non-volatile organic acids, and reduce rock particle size to form mineral soil. This study revealed the presence of large populations of culturable endophytic bacteria inside the seeds extracted from wild plants, from seeds extracted from the guano of bats feeding on cactus fruit, in seedlings growing from these seeds, in the pulp of fruit, and in small, mature wild plants, and are comparable in size to populations of endophytic populations in some agricultural crops. The dominant culturable endophytes were isolates of the genera Bacillus spp., Klebsiella spp., Staphylococcus spp., and Pseudomonas spp. Based on partial sequencing of the 16s rRNA gene, the isolated strains had low similarity to known strains in these genera. However, these strains have higher molecular similarity among endophytes obtained from seeds, endophytes from roots, and some bacterial strains from the rhizoplane. Seedlings developed from seeds with endophytes contain the similar species of endophytes in their shoots, possibly derived from the seeds. This study shows the involvement of endophytic bacteria in rock weathering by cacti in a hot, subtropical desert and their possible contribution to primary colonization of barren rock. This study proposes that cacti capable of acquiring diverse populations of endophytes may give them an evolutionary advantage to gain a foothold on highly uncompromising terrain.     

ENHANCEMENT OF INBREEDING DEPRESSION BY DOMINANCE AND SUPPRESSION IN IMPATIENS CAPENSIS

We investigated the effect of intraspecific competition on the magnitude of inbreeding depression in Impatiens capensis by planting seeds from chasmogamous (CH) and cleistogamous (CL) flowers in three experimental greenhouse treatments: in individual pots, in flats in dense pure stands according to seed type, and in flats with the two seed types intermixed in a checkerboard array. The size distributions of plants grown in flats were significantly more hierarchical than those of plants grown individually, indicating that larger plants competitively suppressed smaller plants in the high-density treatments. The magnitude of inbreeding depression at high density depended upon the planting arrangement of CL and CH seeds. CH advantage was greatest when CH and CL seedlings were grown in competition with one another, suggesting that fitness differences between outcrossed and inbred individuals were intensified by dominance and suppression. For plants grown individually, the effects of maternal parent, seed weight, and emergence date on seedling size disappeared with plant age, whereas at high density these effects remained at the final harvest. Thus, plant density may influence patterns of natural selection both on mating system and on juvenile traits in natural Impatiens populations.     

Monoecious Plants of Myriophyllum ussuriense (Regel) Maxim. in Japan

Myriophyllum ussuriense has been described as dioecious but monoecious plants were newly found from some populations in south-western Japan. Sex expression of monoecious plants proved labile and they sometimes bore male or female flowers alone. On the other hand, sex expression of dioecious plants was stable and seemed to be fixed genetically. M. ussuriense may be still in the course of differentiation from monoecy to dioecy. Received 13 April 2001/ Accepted in revised form 22 May 2001     

SEED ABORTION AND SEED SIZE VARIATION WITHIN FRUITS OF PHASEOLUS VULGARIS: POLLEN DONOR AND RESOURCE LIMITATION EFFECTS

A new plant cohort starts with the formation of zygotes at ovule fertilization. This paper presents an ecological study of the effects of resource limitation and pollen donor on the survivorship and size of plant embryos in maturing fruits of the common bean, Phaseolus vulgaris L. For naturally selfed flowers of domesticated and wild beans, embryos in basal ovular positions were more likely to abort or, if they survived, become lighter seeds than stylar embryos. Embryo survivorship was higher for the cultivar when maternal plants grew in larger pots. Early seed abortion in the cultivar was random with respect to ovular position. In late abortion basal embryos had a higher mortality than stylar ones. The basal embryos lacked gross genetic defects since they developed adult organs in embryo culture. In more outbred crosses with the cultivar the position effect in embryo survivorship and size disappeared. This difference among paternal parents implicates a genetic component to the observed position effects. The variation in mature seed size may affect adult characteristics since large seeds produced larger juvenile plants than small seeds.     

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.