CLEISTOGAMY IN ANDROPOGON L. (GRAMINEAE)

Eleven species of Andropogon produce single-stamened flowers, and all but one of these species bear some cleistogamous flowers. Nine of the species, all diploid perennials, make up the Andropogon virginicus complex. Cleistogamy in the virginicus complex results from precocious maturation of the flowers, which leaves them confined within their spikelets at anthesis. Variation in flowering mode (expressed as mean % chasmogamy) is correlated significantly (P < 0.01) with variation in peduncle length (r_s = 0.67), spikelet length (r_s = 0.49), and anther length (r_s = 0.69). The strength of the correlation (r = 0.94) between pollen grain number per anther and anther length makes the latter a good measure of pollen-ovule ratios in the single-stamened flowers of this group. Taxa with the highest frequency of cleistogamous flowering are also the most successful colonizers of disturbed habitats. It is suggested that cleistogamy is important in the reproductive isolation of the numerous, closely related subspecific taxa of the virginicus complex.     

Heterocarpy in Salvia roemeriana (Lamiaceae)

Although Salvia roemeriana has long been known to produce both chasmogamous and cleistogamous flowers, the mericarps resulting from those flowers have received little attention. We germinated seeds from chasmogamous and cleistogamous flowers, recorded germination times, and fit time‐to‐germination, three‐parameter log‐logistic regressions to analyze differences in germination progress. Additionally, we compared the mass and size of mericarps from both kinds of flowers. Our results show that the mericarps produced from chasmogamous flowers are larger and heavier than those from cleistogamous flowers. In addition, seeds from chasmogamous flowers had a longer dormancy than those from cleistogamous flowers. This is the first report of heterocarpy in Salvia and in the family Lamiaceae. Together, cleistogamy and heterocarpy are a multiple strategy that may be advantageous in heterogeneous environments.     

Reproductive ecology of a cleistogamous annual,Impatiens noli-tangere L., occurring under different environmental conditions

Demographic and reproductive schedules were compared among five populations of a cleistogamous annual,Impatients noli-tangere L., occurring in habitats with contrasting moisture and/or light environments. In all populations, flowering extended for 2–3 months during which light environment and mortality changed. Seasonal patterns of growth and mortality were significantly different among the five populations studied. The beginning and duration of flowering and the ratio of chasmogamous flowers to cleistogamous ones were also significantly different among populations. An experiment was conducted under different light conditions (open and closed) to separate the genetic and environmental components of the variation in reproductive traits observed among populations ofI. noli-tangere. Transplanted plants showed significant among-population variation in flowering time, as is observed in natural populations, suggesting genetic differentiation among populations of this species. On the other hand, the ratio of chasmogamous flowers did not differ among plants transplanted from three populations. Based on these results, the authors suggest that facultative cleistogamy is a conditional strategy under seasonally changing environments.     

Cleistogamy in the rare high Andean perennial herb Cryptantha capituliflora (Boraginaceae)

 Axial and apical flowers of Cryptantha capituliflora were analyzed with regard to morphology and pollen tube growth to assess the occurrence of cleistogamy. Although intermediate floral forms do occur, cleistogamous flowers were significantly smaller than chasmogamous flowers, had fewer anthers, and showed a distinctive stigmatic surface. Chasmogamous flowers can be cross-pollinated. Nevertheless, the growth of self-pollen tubes in few chasmogamous buds jointly with flower characters suggests that these flowers can probably produce fruits through autonomous selfing. The mean seed number per fruit did not differ between fruits from chasmogamous and cleistogamous flowers. Cleistogamous flowers were only observed in axial inflorescences, which are completely covered by the leaf. Other species of section Cryptantha also show the same trend, with cleistogamous flowers located in the lower half of the stems. This pattern is discussed in relation to dissimilarities in the outcrossing opportunities between flower types within the plant. Received May 22, 2002; accepted November 14, 2002 Published online: March 20, 2003     

The maintenance of mixed mating by cleistogamy in the perennial violet Viola septemloba (Violaceae)

The production of both potentially outcrossed (chasmogamous) and obligately self-fertilized (cleistogamous) flowers presents a clear exception to the prediction that the only evolutionarily stable mating systems are complete selfing and complete outcrossing. Although cleistogamy has evolved repeatedly, the reason for its stability is not known for any species. We tested the hypothesis that the production of cleistogamous and chasmogamous flowers by a perennial violet constitutes adaptive phenotypic plasticity. We manipulated the season of flowering for each flower type and determined fruit set and the germination percentage of seeds produced by cleistogamous and chasmogamous flowers to test the hypothesis that adaptive plastic response to seasonal environmental variation makes mixed mating stable. Cleistogamous flowers had greater fruit set in all seasons and produced seeds with germination percentages as great as or greater than those from chasmogamous flowers. The consistent advantage of cleistogamous flowers is clearly not consistent with a role of adaptive plastic response to seasonal variation. The biomass cost of seed production by chasmogamous flowers was nearly three times that for cleistogamous flowers. Explaining why chasmogamous flower have not been eliminated by natural selection requires that this difference be balanced by an advantage to chasmogamous flowers that has not yet been identified.     

DISPERSAL OF HETEROMORPHIC SEEDS IN AMPHICARPAEA BRACTEATA (FABACEAE)

The dispersal of the chasmogamous, aerial cleistogamous, and subterranean cleistogamous seeds of Amphicarpaea bracteata was examined. The chasmogamous and aerial cleistogamous seeds are ballistically dispersed. Chasmogamous seeds were dispersed farther than the aerial cleistogamous seeds due to the height advantage of the chasmogamous pods. There was no difference in the firing angle or the initial velocity of the seeds discharged from the two aerial pod types. The subterranean cleistogamous seeds are “dispersed” the shortest distance by the elongation of runners. Differential dispersal of the three types of seeds may be a factor involved in the evolution of cleistogamy in A. bracteata.     

MITOCHONDRIAL DNA VARIATION IN THE FUNGUS ATKINSONELLA HYPOXYLON INFECTING SYMPATRIC DANTHONIA GRASSES

The purpose of this study was to determine how host identity and geographic proximity were related to genetic variation in the fungus Atkinsonella hypoxylon infecting four sympatric Danthonia grasses. We analyzed 192 A. hypoxylon isolates from 48 geographic sites for mitochondrial DNA (mtDNA) variation by restriction digestion of total DNA with EcoRI, BamHI, and PstI, and subsequent hybridization with purified A. hypoxylon mtDNA. Thirtynine haplotypes were identified. UPGMA cluster analysis showed that mtDNA type is highly correlated with host-species group; isolates from Danthonia compressa and D. spicata formed one branch of the phenogram, and isolates from D. epilis and D. sericea formed the other. Restriction maps of the most common mtDNA haplotypes infecting each host-species pair revealed a 30-kb size difference and a minimum of eight length changes and one restrictionsite change between them. Mapping of the mutation differences among all haplotypes occurring at North Carolina sites resulted in two distinct gene trees corresponding to the two Danthonia species groups, corroborating the phenetic analysis. The results indicate a high degree of host-dependent isolation and establish the existence of host races in A. hypoxylon. Possible mechanisms responsible for this isolation are discussed. Little differentiation existed between isolates from within a Danthonia species pair, and some variation was explained by geographic origin. Analysis of progeny from a natural sexual cross revealed that mitochondria are maternally inherited in A. hypoxylon.     

THE HETEROCHRONIC ORIGIN OF THE CLEISTOGAMOUS FLOWER IN ASTRAGALUS CYMBICARPOS (FABACEAE)

Members of the species Astragalus cymbicarpos form chasmogamous and cleistogamous flowers, as well as a large variety of intermediate floral types. Bivariate allometry and Gould's clock models were used to investigate the possible heterochronic evolution of the cleistogamous flower from the chasmogamous flower. In three of the whorls analyzed (pistil, stamens, and corolla) comparison of chasmogamous, pseudocleistogamous, and cleistogamous flowers revealed a progressive “juvenilization” of the adult form. This paedomorphic morphology proved to be partly the result of a process of progenesis, also evident in acceleration of sexual maturity. The retardation of shape with respect to size in these three whorls suggests, however, the existence of a heterochronic process other than progenesis.     

Genetic and linkage analysis of cleistogamy in soybean

Early maturing cultivars of soybean [Glycine max (L.) Merr.] native to the shores of the Sea of Okhotsk (Sakhalin and Kuril Islands) and eastern Hokkaido (northern Japan) have been used in breeding for chilling tolerance. These cultivars have a strong tendency to produce cleistogamous flowers throughout their blooming period. This study was conducted to determine the genetic basis of cleistogamy in an early maturing cultivar, Karafuto-1, introduced from Sakhalin. Genetic analysis was performed using F1 plants, the F2 population, and 50 F3 families produced by crossing between Karafuto-1 and a chasmogamous cultivar, Toyosuzu. F2 plants had chasmogamous flowers, indicating that chasmogamy was dominant to cleistogamy. Analysis of F2 populations and F3 families generated segregation data that was close to a two-gene model with epistatic interactions, although a portion of the pooled F3 data on the frequency of chasmogamous segregants from cleistogamous families significantly deviated from the model. The results suggested that a minimum of two genes with epistatic effects were involved in the genetic control of cleistogamy. Furthermore, cleistogamy was associated with early flowering in the F2 and F3 populations. A gene for cleistogamy was linked to one of the recessive genes responsible for insensitivity to incandescent long daylength.     

Reproductive investment in a cleistogamous morph of <Emphasis Type="Italic">Polygonum jucundum</Emphasis> (Polygonaceae)

Cleistogamy, a breeding system with permanently closed and self-pollinated flowers, is expected to assure reproductive success at a lower cost. Previous studies have inferred the occurrence of cleistogamous flowers in Polygonum, but there are no detailed studies on their reproductive investment compared with that of the chasmogamous flowers in this genus. Here, we studied a cleistogamous morph of P. jucundum to investigate the investment in pollen number, tepal and nectary size. The number of pollen grains per flower was counted with a light microscope. Nectaries and perianths were observed via scanning electron microscope and light microscopy, photographed and measured via ImageJ. The perianths of the cleistogamous flowers, as well as the pollen numbers and nectary sizes, were significantly smaller than those of the chasmogamous flowers. The pollen numbers of the CL flowers were seven times lower than those in the CH flowers. The tepal areas of the CL flowers were, on average, approximately 38% those of the CH flowers. The nectary areas of the CH flowers were almost twice those of the CL flowers. In addition, the nectaries of the cleistogamous flowers were degenerated and inconspicuous, in distinct contrast with the well-developed and conspicuous nectaries of the chasmogamous flowers. Self-fertilization was completed and produced seeds. The cleistogamous P. jucundum, compared with chasmogamous individuals, exhibited lower costs in male function, pollinator attraction and reward structure investment. This cleistogamy appears to be favourable for the plant reproduction under suboptimal conditions.     

Isozyme evidence for host races of the fungus Atkinsonella hypoxylon (Clavicipitaceae) infecting the Danthonia (Poaceae) complex in the southern Appalachians

On granite outcrops around Highlands, North Carolina four sympatric Danthonia grass species are infected by the fungus Atkinsonella hypoxylon (Clavicipitaceae). Danthonia epilis and D. sericea (rare hosts) are infected only in this small region, although they range more widely, while D. compressa and D. spicata (widespread hosts) are infected throughout their ranges in eastern North America. To test the hypothesis that infection of the rare hosts has recently spread from the widespread hosts, the genetic structure of fungal populations on different Danthonia hosts was investigated by isozyme electrophoresis. Two hundred isolates were collected from 46 sites and analyzed for ten polymorphic isozyme loci to examine levels of variation, genetic differentiation, and gene flow among fungal populations infecting different hosts. Genetic identities (I) indicated that isolates from within each of the widespread and rare host species pairs were homogeneous (I > 0.99), but the pairs were highly differentiated (0.66 < I < 0.69). With one exception there was no overlap in fungal genotypes isolated from the widespread hosts (12 genotypes) vs. the rare hosts (seven genotypes). Two plants of D. compressa growing sympatrically with infected D. epilis at one site were infected with a genotype characteristic of the latter species. Estimates of gene flow suggest a high degree of host-mediated reproductive isolation in A. hypoxylon, indicating that the fungus forms distinct, long-standing host races or sibling species in the southern Appalachians.     

SPATIAL AND TEMPORAL VARIATION IN CHASMOGAMY AND CLEISTOGAMY IN OXALIS MONTANA (OXALIDACEAE)

Chasmogamous (CH) and cleistogamous (CL) flower production was investigated in natural populations of the perennial herb Oxalis montana in southern Quebec, Canada. Every 10–12 days during two flowering seasons, we recorded the reproductive output of approximately 2,000 plants distributed among five forest sites. The percentage of plants flowering, proportion of flowering plants producing CH and CL flowers, CH and CL flower number per flowering plant, and the proportion of flowers that are CL differed significantly among sites and corresponded to site differences in forest type and habitat quality. Unlike patterns in most CL species, however, reproduction by cleistogamy increased in sites with habitat conditions favorable to plant growth and sexual reproduction, and decreased in less productive sites. Chasmogamous and CL flower production increased with increasing plant size but plant size explained a significantly greater proportion of the variation in CL flower numbers. The percentage of flowering plants producing CH flowers decreased between years while the proportion of CL flowers increased at all sites during the second flowering season. The somewhat unusual aspects of reproduction in Oxalis may stem from its perennial habit which allows use of stored resources in adjusting the balance of cleistogamy versus chasmogamy in different environmental regimes.     

The selection of cleistogamy and heteromorphic diaspores

Models for the evolution of a mixture of cleistogamous (closed, autogamous) flowers and chasmogamous (open) flowers are described. The 'basic' model takes into account features associated with cleistogamous self-pollination, including the greater economy and certainty of cleistogamous fertilization and the inability of cleistogamous flowers to contribute pollen to the outcrossed pollen pool. Complete cleistogamous selfing is favoured when allocation to maternal function, fertilization rate, and viability of progeny are sufficiently greater for the cleistogamous component, and when the resources spent on ancillary structures in cleistogamous flowers, cleistogamous seed costs, and inbreeding depression are low. The result is discussed with respect to the cost of sex argument and relevant ecological data. Suggestions for the apparent rarity of cleistogamy are presented. The 'complex habitat' model extends the basic model to situations in which the success of reproduction by cleistogamy or chasmogamy varies according to the environment of the parent. In this situation, reproduction by both cleistogamy and chasomogamy is sometimes selected. A 'near and far dispersal' model addresses the question of the evolution of dual modes of dispersal, which occur in some cleistogamous and non-cleistogamous plants. A dual mode of dispersal may evolve if a narrowly dispersed seed type is more successful in establishing at the sites located within its dispersal range compared with a second, more widely dispersed seed type which experiences less sib competition. The prediction is discussed with respect to data from amphicarpic plants.     

Consequences of a mixed reproductive system in the hog peanut,Amphicarpaea bracteata, (Fabaceae)

Summary The forest annual, Amphicarpaea bracteata L. can reproduce via aerial chasmogamous, aerial cleistogamous, and subterranean cleistogamous flowers. Both plant size and light intensity influenced the utilization of the three modes of reproduction. chasmogamous and aerial cleistogamous flower number and the ratio of chasmogamous flowers to the total number of aerial flowers increased with plant size. The latter demonstrated a shift to xenogamy and outbreeding in larger plants. Light intensity indirectly influenced reproductive modes through its infuence on plant size. Seed set by both types of aerial flowers was low and unrelated to plant size. Subterranean seed number and the total dry weight of subterranean seeds per plant increased with size. The subterranean seeds of Amphicarpaea bracteata are thirty-four times larger than the aerial seeds (fresh weight). Under field conditions, subterranean seeds had greater germination after one year than acrial seeds. The plants arising from subterranean seeds were significantly larger and more fecund than those from aerial seeds. Seeds produced by aerial cleistogamous, hand selfpollinated chasmogamous, and naturally pollinated chasmogamous flowers had equivalent germination rates and produced plants of equal size and fecundity. This suggests that the outbred progeny from chasmogamous flowers have no advantage over the inbred progeny from aerial cleistogamous flowers.     

Level ofFusarium infection in wheat spikelets related to location and number of inoculated spores

The flowering time is the most susceptible period for primary infection of wheat heads byFusarium spp. During this period spores can be deposited into the opened wheat florets where they may later cause infections. We quantitatively explored the relationship between variables related to the flowering process and the infection level byFusarium graminearum in single spikelets. We imitated open (chasmogamous) and closed (cleistogamous) flowering by injecting well-defined amounts of spores into and between wheat florets. Applying the spores between the florets resulted in weaker disease symptoms and significantly lower amounts ofFusarium mycotoxins. With larger numbers of spores, the disease symptoms became more pronounced and the mycotoxin amounts per spikelet increased significantly. Our results indicate that the probability of primary infection is approximately proportional to the number of spores reaching the open florets during the flowering process. The breeding of wheat lines which flower partially or completely cleistogamously might reduce theFusarium susceptibility in wheat.     

A COMPARISON OF VARIABILITY IN UTRICULARIA CORNUTA AND UTRICULARIA JUNCEA

Although Utricularia cornuta Michx. and U. juncea Vahl, sympatric in the southeastern United States, have been considered conspecific by various authors, the present biosystematic approach shows them to be separate species. The taxa are seasonally isolated. While both have the same chromosome number (n = 9), strong internal isolation is apparent since artificial hybrids cannot be produced by standard methods. In Utricularia cornuta the mean values of characters studied quantitatively are much higher than those of U. juncea though the extremes of the ranges may overlap. Utricularia juncea has both cleistogamous flowers and chasmogamous flowers while U. cornuta has only chasmogamous flowers. The flowers are self-fertile and apparently are usually, if not always, self-pollinated, even though they are highly adapted to specialized insect pollinators.     

Effects of light and nutrient availability on chasmogamy and cleistogamy in an understory tropical herb,Calathea micans (Marantaceae)

The effects of light and nutrient availability on chasmogamous and cleistogamous flower and fruit production were investigated in an understory tropical herb, Calathea micans (Marantaceae). I censused chasmogamous and cleistogamous flower and fruit production at five permanently marked demographic study sites in Costa Rica, characterized by different successional stages. Using a transplant experiment, I examined whether an increase in light and/or nutrients would increase chasmogamous and/or cleistogamous flower production. In natural populations as well as in the experiment, chasmogamous reproduction increased with light; nutrient availability significantly increased chasmogamous reproduction in the transplant experiment. Field observations indicated that very few plants reproduced and low investment in reproduction might have been due to poor environmental conditions. Larger plants had a higher probability of reproducing, but plant size did not have an effect on the mode of reproduction, chasmogamy vs. cleistogamy. However, the production of new vegetative shoots increased the probability of producing chasmogamous inflorescences. Cleistogamy may allow the plant to reproduce even in conditions of poor habitat quality and assures some seed set even in conditions unfavorable to plant growth.     

EFFECTS OF WATER STRESS,ABSCISIC ACID,AND GIBBERELLIC ACID ON FLOWER PRODUCTION AND DIFFERENTIATION IN THE CLEISTOGAMOUS SPECIES COLLOMIA GRANDIFLORA DOUGL. EX LINDL. (POLEMONIACEAE)

The effects of water stress, abscisic acid (ABA), and gibberellic acid (GA₃) on flower production and differentiation by Collomia grandiflora were investigated. An untreated plant typically produced both small, closed cleistogamous (CL) and large, open chasmogamous (CH) flowers. The larger corolla of CH flowers was due to a greater cell number and size. When plants were water-stressed or sprayed with ABA, both the percentage of CH flowers and the total number of flowers were reduced significantly. The corolla dimensions and epidermal cell numbers and sizes of CL flowers produced by water-stressed and ABA-sprayed plants did not differ from those of CL flowers produced by control plants. Application of GA₃ to both well-watered and water-stressed plants significantly increased the percentage of CH flowers formed compared to well-watered controls. In the absence of GA₃, water-stressed plants produced almost entirely CL flowers. GA₃-sprayed plants produced CH flowers whose corolla dimensions were intermediate between those of CL and CH flowers formed by control plants. Epidermal cells of these intermediate corollas were reduced only in number and not in size when compared to control CH flowers. Endogenous levels of ABA and gibberellins may control the type of flower produced by C. grandiflora and may mediate some of the observable effects of water stress on flowering.     

A unique growth pattern associated with cleistogamy in a tropical legume, Vigna minima (Roxb.) Ohwi & Ohashi (Leguminosae)

Obligate subterranean cleistogamy, observed in a natural population of Vigna minima , is associated with the production of one or two negatively geotropic, leafy shoots and several positively geotropic, highly pigmented, leafless shoots. The latter branch profusely after penetrating the soil and produce much reduced, cleistogamous flowers. The undehisced anthers contain germinated pollen grains. The seeds developing from the cleistogamous flowers differ in size, weight and surface features from those produced by the chasmogamous flowers of other natural populations within the species.     

Why is cleistogamy a selected reproductive strategy in Impatiens capensis (Balsaminaceae)?

Cleistogamy (self-fertilization in closed flowers) differs from chasmogamy (open-pollinated fertilization) mainly in sustaining selfing. Why numerous species develop both of the reproductive modes on the same individuals has long puzzled biologists. In a novel hypothesis presented here, I propose that cleistogamy could be a means by which inbred lines are created and maintained in natural populations; these lines would continuously experience self-improvement via natural selection and via crosses among lines at the chasmogamous flowers to benefit the populations. Supporting evidence for the hypothesis was found in Impatiens capensis where cleistogamous ovules were fertilized proportionately less (56%) than chasmogamous ovules (67%) in natural populations, but crosses among cleistogamous progeny in the greenhouse led to a nearly 10% increase of fertilized chasmogamous ovules. I established a novel fitness model specific to the cleistogamous species to further examine how various aspects of the mating system affect plant performance. A low inbreeding depression (0.07) was consequently found for the surveyed natural populations of I. capensis , suggesting that the individual-level percentage of cleistogamy and the population-level selfing rate may have evolved in the direction of reducing the overall inbreeding depression. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 543–553.