REPRODUCTIVE BEHAVIOR OF AMPHICARPAEA BRACTEATA (LEGUMINOSAE), AN AMPHICARPIC ANNUAL

Amphicarpaea bracteata, an annual legume common in woodland communities in the eastern United States, produces three distinct types of flower: subterranean cleistogamous (SCL), aerial cleistogamous (ACL), and aerial chasmogamous (ACH). We sought to quantitatively describe the growth and reproduction of four diverse A. bracteata populations in an effort to explain the adaptive significance of this species’ variable reproductive modes. Virtually all plants develop 1 to 3 SCL seeds on subterranean cotyledonary shoots, starting in late July. Larger plants produce additional SCL seeds later from the tips of axillary shoots which bury. Seeds produced by SCL flowers are large (39 to 134 mg), have restricted dispersal, lack dormancy, have high germination , and produce relatively vigorous seedlings. ACL flowers develop from early August through plant senescence in early October, while ACH flowers appear in a pulse in late August. The number of each aerial flower type was positively correlated with plant size. The ACL flowers take significantly less time to develop mature fruits than do ACH flowers. Both aerial seeds are smaller than the subterranean (mean weights: ACL 12.1, ACH 8.7 mg), have lower germination (ACL 20.2%, ACH 15.3%), appear relatively resistant to environmental extremes, and have the potential for more widespread dispersal. Plants derived from subterranean seeds are much larger than plants derived from aerial seeds, which makes them more likely to produce axillary SCL and aerial seeds. Only larger plants derived from subterranean seeds and growing in favored sites produce ACH flowers. This imposes an alternation of selfing with outcrossing generations. This pattern of reproductive behavior in A. bracteata is similar to that found in several other amphicarpic species.     

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.     

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.     

Why is there variation in mean seed size among plants within single populations? test of the fertilization efficiency hypothesis

We tested the fertilization efficiency hypothesis, which attempts to explain mean seed size variation among plants within single populations, by comparing the patterns of seed size variation between chasmogamous (CH) flowers and cleistogamous (CL) flowers in Impatiens noli-tangere and Viola grypoceras, respectively. The fertilization efficiency hypothesis predicts that larger plants produce larger seeds if the number of pollen grains captured by a plant increases with increased allocation of resources to its attractive structures (e.g., corolla and nectar), but with diminishing gains. Thus, seed size should depend on plant size in seeds from CH flowers because of the diminishing gains of capturing pollen in these flowers, whereas seed size should not depend on plant size in seeds from CL flowers because CL flowers need not capture outcross pollen. We found significant positive correlations between mean seed size per plant and plant size for seeds from CH flowers in both species. However, there was no significant positive correlation between these two factors for seeds from CL flowers of both species. The results of the present investigations were thus consistent with the fertilization efficiency hypothesis.     

Reproductive Biology of an Amphicarpic Annual, Polygonum thunbergii (Polygonaceae): Spatio-temporal Changes in Growth, Structure and Reproductive Components of a Population over an Environmental Gradient

Abstract The process and mechanisms of spatio-temporal changes in growth, population structure, as well as various yield and reproductive components of a population of an amphicarpic annual, Polygonum thunbergii (Polygonaceae), with two reproductive systems (aerial chasmogamous flowers and subterranean cleistogamous flowers) along an environmental gradient (light, moisture and soil nitrogen levels) were investigated in the field.
The results clearly demonstrate that growth and allocation patterns, population structure, and reproductive output of individuals changed sharply along the environmental gradient in response to seasonal and spatial changes in resource availability. Models predicted that light conditions bring about one-sided competition, whereas nutrient conditions in the soil engender two-sided competition. As expected, the degree of one-sided competition was prominent in the case of a planophile, Polygonum thunbergii. Allocation patterns, seed outputs, individual seed size as well as relative energy costs of chasmogamous and cleistogamous seed as affected by light and nutrient levels were also critically analyzed. The most noteworthy finding was that the size of chasmogamous seeds sharply decreased in response to a decrease in the light regime, while cleistogamous seed size remained constant along the gradient. However, relative cost of both chasmogamous and cleistogamous seeds sharply increased with decrease in the light level, reflecting different degrees of environmental stress, biotic interference, or both.     

Genetic diversity of an amphicarpic species, Amphicarpaea edgeworthii Benth. (Leguminosae) based on RAPD markers

Amphicarpaea edgeworthii Benth. is an amphicarpic legume widespread in China. Amphicarpy describes the phenomenon that a plant produces aerial as well as subterranean fruits. A. edgeworthii can reproduce via three kinds of flowers: aerial chasmogamous flowers, aerial cleistogamous flowers, and subterranean cleistogamous flowers. Although there are some studies on the population genetic structure of species with both chasmogamous and cleistogamous flowers, none has been done for that of an amphicarpic species so far. The present study uses random amplified polymorphic DNA (RAPD) to assess level and pattern of genetic variation in 15 natural populations of A. edgeworthii. A total of 131 stable and clearly scored RAPD bands were achieved from 13 primers. The average genetic diversity within populations estimated by Shannon's information index was 0.218 at the population level, but ranged from 0.119 to 0.302, which was significantly different (P < 0.01). Different statistical analyses revealed a high level of genetic differentiation among populations (G_ST = 0.473–0.527). Thus, the pattern of genetic structure of A. edgeworthii is consistent with that of an inbreeding species.     

Herbivory alters the expression of a mixed-mating system

The direct and indirect effects of vegetative herbivory on the mating system of Impatiens capensis were analyzed through a survey of herbivory in natural I. capensis populations and manipulation of leaf damage in the field. Across 10 wild populations of I. capensis proportion of cleistogamous flowers had a significant positive exponential relationship with natural levels of herbivory. Similarly, experimental leaf damage increased the proportion of flowers and seeds that were cleistogamous. Leaf damage also reduced the biomass of cleistogamous progeny more severely relative to that of chasmogamous progeny. The cumulative effect of leaf damage was to increase plant reliance on fitness derived from cleistogamous progeny. Leaf damage indirectly affected mating system traits by reducing chasmogamous flower size, leading to a reduction in pollinator visitation. Under these experimental conditions, herbivory did not significantly reduce the number of simultaneously open flowers and potential for geitonogamy, nor did it result in significant changes in the composition of the pollinator fauna. These findings are among the first to demonstrate that herbivory has consequences for mating system and should be considered a factor shaping mating system evolution.     

CLEISTOGAMY IN MICROLAENA POLYNODA (GRAMINEAE): AN EXAMINATION OF SOME MODEL PREDICTIONS

A population of the cleistogamous grass, Microlaena polynoda, was investigated to determine some of the factors responsible for adjusting the balance between reproduction by chasmogamous (CH) and cleistogamous (CL) flowers, the tissue costs associated with the two reproductive modes, the fate of progeny produced by each mode, and the genetic diversity of the progeny. Cleistogamous flower production begins earlier in the season than CH flower production. There is a distinct threshold of low light intensity below which mostly CL flowers are produced. Paternal expenditure per plant is an order of magnitude larger for the CH than the CL component. The opposite relation holds for maternal expenditure. Increased maternal expenditure in the CL component may be due to greater fertilization success and retrieval of paternal costs. Cleistogamous seeds are dispersed later than CH seeds. Following dispersal, the spikelet encloses the CL seed but not the CH seed, and is responsible for inducing dormancy. The ratio of seedlings arising from CH seeds to that from CL seeds in a natural habitat is significantly lower than the ratio of estimated numbers of CH to CL seeds produced. There were no detectable polymorphisms among ten presumptive enzyme loci assayed. Many of the features associated with CH and CL reproduction in M. polynoda are in accord with the theoretical requirements for the evolution of closed flower self-pollination and the maintenance of two distinct methods of reproduction.     

Divergence and isolation of cryptic sympatric taxa within the annual legume Amphicarpaea bracteata

The amphicarpic annual legume Amphicarpaea bracteata is unusual in producing aerial and subterranean cleistogamous flowers that always self‐fertilize and, less commonly, aerial chasmogamous flowers that outcross. Although both morphologic and genetic variants are known in this highly selfing species, debate continues over whether this variation is continuous, reflecting the segregation of standing genetic variation, or discontinuous, reflecting distinct taxa that rarely intercross. We characterized SNP variation in 128 individuals in southern Wisconsin to assess within‐ and among‐population variation at 3928 SNPs. We also assessed genotype and leaf morphology in an additional 76 individuals to connect phenotypic variation with genetic variation. Genetic variation maps onto three strongly divergent and highly inbred genetic groups showing little relation to site location. Each group has a distinct phenotype, but the divergence of these groups differs from the varietal divisions previously identified based on morphological characters. Like previous authors, we argue that the taxonomy of this species should be revised. Despite extensive sympatry, estimates of among‐group migration rates are low, and hybrid individuals were at low frequency (<2%) in our dataset. Restricted gene flow likely results from high selfing rates and partial reproductive incompatibility as evidenced by the U‐shaped distribution of pairwise F_ST values reflecting “islands” of genomic divergence. These islands may be associated with hybrid incompatibility loci that arose in allopatry. The coexistence of lineages within sites may reflect density‐dependent attack by species‐specific strains of pathogenic fungi and/or root‐nodulating bacteria specializing on distinct genotypes.     

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.     

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.     

Sex expression and breeding strategy in <Emphasis Type="Italic">Commelina benghalensis</Emphasis> L.

This paper describes the results of a series of experiments conducted to unravel the patterns of sex expression and reproductive output in a fascinating species with high variation in sexuality. Commelina benghalensis L., an andromonoecious rainy season weed, bears male and bisexual flowers in axillary spathes of all the plants investigated. Bisexual flowers are of two types; chasmogamous (CH) and cleistogamous (CL). The former are borne on subaerial and the latter on subterranean shoots, in addition to those on aerial spathes. Three populations of the species, designated JU1, JU2 and JU3, were scanned for three consecutive years from 1996 to 1998, and the number and distribution of male, CH and CL flowers per plant were found to vary. The mere number of CH/CL flowers per plant is by itself not an accurate measure of mixed mating. It is necessary to confirm that CH flowers actually outcross and, if they do so, to what extent. Comparison of the pollen/ovule (P/O) ratio and percentage pollen germination on the stigmas of the CH and CL flowers have been used as indices of the pollination system. Confirmation of this was sought from the fruit and seed sets obtained after manual pollination of emasculated flowers with self- and cross-pollen. Results so obtained were compared with those of natural pollination. In the majority of CH flowers, the male and female reproductive phases (i.e. anther dehiscence and stigma receptivity) overlap, providing for self-pollination. However, two exceptions to this general behaviour were found in some plants of all the three populations. In some CH flowers, the female phase matures prior to anther dehiscence while in others, the anthers are sterile. Such plants, designated as variants 1 and 2, respectively, facilitate cross-pollination. While the CL flowers contribute to the production of selfed progeny, the variants of CH ones permit formation of outcrossed progeny, indicating a mixed mating strategy in C. benghalensis.     

FACTORS INFLUENCING SEED WEIGHT IN IMPATIENS CAPENSIS (BALSAMINACEAE)

I analyzed within- and between-plant variation in seed weight from one greenhouse and four natural populations of Impatiens capensis Meerb. Most variation in seed size in natural populations occurred within populations and within plants. Weights of seeds from cleistogamous flowers on greenhouse plants increased with height on the plant and proximity to the main stem, a result that may be related to the local supply of photosynthate. Mean seed weight in the natural populations declined during seed set, then increased again. Seeds from chasmogamous flowers were significantly larger in one of the four populations. Maternal parents differed significantly in both mean seed weight and the relative size of the two types of seeds. Combined genetic and general environmental effects accounted for 26% of the total phenotypic variance, which provides an outside estimate of the heritability of this character.     

Fruit dimorphism inPhaseolus sublobatus (Fabaceae) and its evolutionary significance

Populations of the annualPhaseolus sublobatus from different ecogeographical zones are genetically differentiated. In twelve populations from the western ghats (Maharashtra range) chasmogamous flowers are arranged in peduncled capitate racemes borne in leaf axils of higher nodes. One population (Poona Race S₄) has additional inflorescences, also with chasmogamous flowers, on the main axis between the cotyledonary node and the ground. When the pods of these flowers ripen, the inflorescence gradually coils and, eventually, gets bury the fruits in the soil. This phenomenon is not known in any other plant.—The seed-coat patterns as revealed by SEM, and germination behaviour of both aerial and subterranean seeds are similar, and both types of seeds lack dormancy.—Heavy seed predation selection pressure is probably the principal cause for the evolution of fruit dimorphism inP. sublobatus. The origin of geocarpy in relation to fruit dimorphism and seed predation is discussed.     

青藏高原东缘块茎堇菜鳞茎分配的个体大小依赖性

块茎堇菜(Viola tuberifera)为青藏高原特有两型闭锁花植物,属多年生草本,具独特的混和交配系统,既可通过早春开放花异花受精和夏季地上地下闭锁花自花受精有性繁殖,还可通过秋季新鳞茎无性繁殖产生后代。高山环境下,异花受精常因花粉限制而无法正常进行,自花受精和克隆繁殖成为保障植物种群正常繁衍的不二之选,而克隆繁殖更能在植株资源消耗最小的情况下保障子代的存活。该文以青藏高原东缘高寒草甸的混合繁育植物块茎堇菜为研究对象,探索其生长期内鳞茎分配的个体大小依赖性,以及植株如何权衡鳞茎的资源分配以适应个体大小的变化。结果表明：块茎堇菜生活史阶段的鳞茎分配具有个体大小依赖性,鳞茎分配与个体大小呈极显著负幂指数相关关系(P<0.01),个体越大,鳞茎分配越小;反之,个体越小,鳞茎分配越高。即块茎堇菜对鳞茎的资源投入受个体大小的制约,通过鳞茎分配比例的高低响应植株自身资源状况的变化,保障在高寒环境下植物种群的生存和繁衍。该研究结果为高山植物克隆繁殖的生活史进化提供了依据。     

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.     

Plasticity in the reproductive strategy of a clonal cleistogamous species, <Emphasis Type="Italic">Pseudostellaria heterophylla</Emphasis>

The presence of cleistogamous and chasmogamous flowers on the same plant individual is considered to represent a “mixed” reproductive strategy. If a cleistogamous species also exhibits clonal propagation, then competition for limited resources is assumed to exist among the three reproductive modes. To date, however, the relationships and interactions among cleistogamous, chasmogamous, and clonal modes of reproduction have received little attention. In this study, we performed manipulative experiments to investigate the interactions among these different types of reproduction in the perennial herbaceous plant species Pseudostellaria heterophylla. The results showed that 66.4%–87.6% of individuals produce chasmogamous flowers and that the fruiting rates of these flowers in each surveyed population were between 23.5% and 77.4%. Furthermore, we found that 8.3% of the individuals of this species show inbreeding depression. We also detected significant negative correlations between the production of chasmogamous and cleistogamous flowers and between cleistogamous flower production and root tuber mass. However, chasmogamous flower production in an individual plant was found to have little influence on its subsequent clonal propagation. We propose that the plasticity of reproductive strategies observed in P. heterophylla is due to changes in the resource pool and resource allocation.     

VARIATION IN THE DEGREE OF CLEISTOGAMY WITHIN AND AMONG SPECIES OF THE GRASS DANTHONIA

North American species of the grass Danthonia bear distinct chasmogamous and cleistogamous flowers. Four taxa occurring in North Carolina (D. compressa, D. epilis, D. sericea, and D. spicata) were examined for variation in the degree of cleistogamy. The flowering culms bear a single terminal panicle consisting of 4 to 12 spikelets of chasmogamous flowers. In each leaf axil, at the nodes of the flowering culm, is a single spikelet of cleistogamous flowers completely surrounded by the leaf sheath. The percentage of cleistogamous flowers produced on any culm depends on the number of chasmogamous and cleistogamous spikelets and the number of flowers in each type of spikelet. All four characters vary among the taxa examined. Danthonia compressa produces, on the average, 50% cleistogamous flowers; D. spicata, 25% cleistogamous flowers; D. sericea and D. epilis, 5% cleistogamous flowers. The degree of cleistogamy in D. spicata is associated with certain habitat features. Populations from mountain sites, disturbed sites, and non-woodland sites produced higher percentages of cleistogamous flowers than did populations from piedmont, undisturbed, and woodland sites, respectively. Grazing may favor increased cleistogamy because cleistogamous flowers are produced lower on the plant. In North Carolina, the most frequently grazed Danthonia taxa also produce the highest percentage of cleistogamous flowers.     

FIELD GERMINATION AND SEEDLING GROWTH OF CH AND CL PROGENY OF IMPATIENS CAPENSIS (BALSAMINACEAE)

The germination behavior and early growth of chasmogamous (CH) and cleistogamous (CL) progeny of Impatiens capensis were investigated in two eastern Nebraska stands. Field germination of families of buried seeds was scored in mid-April. In one stand a significantly higher proportion of CL seeds germinated than CH seeds while no significant differences were observed in the second stand. Among-family variation in germination rates was significant in both stands. The time course of field germination was similar for CL and CH seeds. A sample of the germinated CL and CH seedlings, matched in size, was planted individually in pots and grown in the greenhouse for five wk. Early growth of CL and CH seedlings was similar. In addition, CL seedlings were as variable as CH seedlings for four morphological traits. Overall, a large fitness advantage for CH progeny was not detected in the germination and early growth of Impatiens capensis seedlings.     

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.