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.     

Chasmogamous and Cleistogamous Pollination in Salpiglossis sinuata

Pollen from chasmogamous flowers of Salpiglossis sinuata L. could not be induced to germinate in vitro unless stigmatal extract was applied to the culture medium. The substance that induces pollen germination in the stigmatal extract is water-soluble and heat-stable. Crosses could not be achieved between chasmogamous and cleistogamous flowers because of structural incompatibility. Pollinated pistils of chasmogamous flowers release a large amount of ethylene. The burst of ethylene release is due to an interaction between pollen tubes and stylar tissue and is directly proportional to the quantity of pollen placed on the stigma. Cleistogamous flower buds also produce a burst of ethylene at the time of pollination within the closed flower. The ethylene release may be a cause of reduced corolla development associated with cleistogamous flowers.     

Evolution of floral dimorphism in a cleistogamous annual, <Emphasis Type="Italic">Impatiens noli-tangere</Emphasis> L. occurring under different environmental conditions

The resource allocation models for the evolution of a mixed production of cleistogamous and chasmogamous flowers were tested in four natural populations of Impatiens noli-tangere L. When we incorporated the estimated parameter values into our basic model, under the assumption that the environmental conditions did not change through the reproductive season, the model predicted the obligate chasmogamy in all four populations. However, including the fluctuation of the fertility of chasmogamous flowers in the model, the seasonal switching of chasmogamous and cleistogamous flower production becomes an evolutionary stable strategy, which explains the reproductive behavior of the individuals in two populations. Only after the relationship between the geitonogamy of chasmogamous flowers and the number of chasmogamous flower production was considered in the model did the simultaneous production of both of the dimorphic flowers become an evolutionary stable strategy. Further, the complete model clarifies the reproductive behavior of the individuals in the other two populations.     

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.     

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.     

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 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.     

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.     

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.     

INTRA-INFLORESCENCE VARIABILITY IN POLLEN/OVULE RATIOS IN THE CLEISTOGAMOUS SPECIES LAMIUM AMPLEXICAULE (LABIATAE)

A comparative study was made of the various floral forms in the main shoot inflorescence of the cleistogamous species Lamium amplexicaule L. (Labiatae). This study revealed significant differences in pollen/ovule (P/O) ratios between successively produced flowers. The variation in P/O ratio is due to a change in pollen count/flower and not in ovule number. The study included two plant populations, one grown in November when only closed flowers are produced, and the other grown in April when open flower production occurs at the upper nodes. The intra-inflorescence variation in P/O ratio among the spring population showed a correlation between P/O ratio and breeding system. The cleistogamous flowers had low P/O ratios of about 200 while the chasmogamous flowers had higher P/O ratios of about 600.     

Divergent pollination systems in the cleistogamous species,Collomia grandiflora (Polemoniaceae)

Summary A structural study of pollination in the dimorphic flowers ofCollomia grandiflora, a cleistogamous species, reveals significant differences in stigma behavior during pollination, stylar structure, the timing of generative cell division, and pollen tube growth rate patterns. The cleistogamous flower shows a loss of protandry and the stigma is receptive only after reflexing and closing of its lobes. In contrast, the chasmogamous stigma is receptive when reflexed and closes when pollen has been deposited on the lobes. Pollen tube penetration of the dry stigma papillae and entry into the style is similar in the two morphs. The chasmogamous style is solid and the cleistogamous style partly hollow. The matrix of secretion produced by the transmitting tract cells is mainly carbohydrate with a trace of lipids. It is fibrillar in nature and appears to be partly comprised of wall material from the transmitting tract cells. In the chasmogamous pollen, the generative cell enters the tube before division, which occurs between 30 and 60 min after pollination. This division correlates with an increased growth rate for the pollen tube. In the cleistogamous pollen, contact with the stigma triggers generative cell division inside the hydrated pollen grain before germination. The two resulting sperm cells exit the grain 15–30 min after pollination when the pollen tube is in the stigma lobes. The cleistogamous pollen tube shows only one phase of growth which occurs at a rate similar to that of the slow, first phase of the chasmogamous pollen.Abbreviations CH chasmogamous - CL cleistogamous - DAPI 4, 6-diamidino-2-phenylindole     

CLEISTOGAMY IN SALPIGLOSSIS SINUATA

Cleistogamy in Salpiglossis sinuata L. (Solanaceae) involves a series of developmental events that includes germination of pollen tetrads en masse inside the anther, penetration of pollen tubes through the anther wall toward the stigma which is in contact with the anther, growth of pollen tubes down the style, and, finally, fertilization. This occurs within 6 days from the first appearance of the flower as compared to 10–12 days for normal anthesis of chasmogamous flowers. Coincident with these events in cleistogamous flowers are lack of corolla development and reduced size of androecium and gynoecium.     

Herbivory and competition interact to affect reproductive traits and mating system expression in Impatiens capensis

As a step toward understanding how community context shapes mating system evolution, we investigated the combined role of two plant antagonisms, vegetative herbivory and intraspecific competition, for reproduction and mating system expression (relative production of selfing, cleistogamous and facultatively outcrossing, chasmogamous flowers and fruits) of Impatiens capensis. In a survey of I. capensis populations, we found that vegetative herbivory and intraspecific competition were positively correlated. In a greenhouse experiment where leaf damage and plant density were manipulated, multispecies interactions had dramatic effects on reproductive and mating system traits. Despite having additive effects on growth, herbivory and competition had nonadditive effects for mating system expression, chasmogamous fruit production, flower number and size, and cleistogamous flower production. Our results demonstrate that competitive interactions influence the effect of herbivory (and vice versa) on fitness components and mating system, and thus antagonisms may have unforeseen consequences for mating system evolution, population genetic diversity, and persistence.     

Reproductive ecology of Viola mirabilis var. subglabra representing intermediate flowering characteristics between stemless and stemmed Viola

Viola (Violaceae) is one of the largest genera in angiosperms. This genus is essentially classified into stemless and stemmed groups based on growth morphology. However, Viola mirabilis var. subglabra is an exception in having intermediate flowering characteristics; cleistogamous (CL) flowers are formed in the axils of stem leaves, whereas chasmogamous (CH) flowers arise from basal rosettes (radical CH (CH(r)) flowers) and also in the axils of the stem (axially CH (CH(a)) flowers). To understand why the pattern of flower production varies in this Viola species, flower production was investigated in 10 Japanese populations from Hokkaido to the western part of Honshu in 2014 and 2015. Furthermore, flower characteristics were also compared between CH(r) and CH(a) flowers in Hokkaido. In this species, the production of CH flowers varied among individuals, and they were categorized into three groups, individuals that produced (i) only CH(r) flowers, (ii) only CH(a) flowers and (iii) both CH flowers. The frequency of these groups differed among populations, but some individuals changed the category between 2014 and 2015. Thus, the production of CH(r) and CH(a) flowers plastically changes depending on individual conditions and/or environmental factors. On the other hand, CH(r) and CH(a) flowers differed in flower size and flowering phenology. These results suggest that two types of CH flowers may play different roles in reproduction in each population, but fruit sets and seed sets did not differ between two types of CH flowers.     

Reproductive effort and sex allocation strategy in Commelina benghalensis L., a common monsoon weed

Commelina benghalensis L. exhibits variability in both foliar and floral features; every plant bears three types of branches and four types of flowers. The branches are negatively geotropic, positively geotropic and diageotropic. The flowers are uni- or bisexual, chasmogamous and cleistogamous. This variability influences the breeding system as well as resource allocation to male and female functions. The plants allocate c. 15% of their total resources to reproduction, the major part of which (68.9%) is devoted to production of aerial branches. The proportion of reproductive effort (RE) allocated to various branch systems is correlated with the availability of resources at the time of their differentiation. The pollen/ovule (P/O) ratio, female : male biomass ratio and reproductive output vary between different flower and branch types; variation is more pronounced in the latter. These variations notwithstanding, the results are in line with Charnov's sex allocation theory. The cleistogamous flowers of aerial branches are, however, an exception, being male- rather than female-biased. The reason behind the deviation is, in all probability, their recent evolution from chasmogamous flowers.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 403−413.     

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.     

Flower variation and breeding systems in theCistaceae

Fourteen common southern Spanish species in theCistaceae were examined for quantitative variations in floral traits. These included the diameter of the corolla, number and size of anthers, number of ovules, and the size and shape of the gynoecium. The variables were in most instances positively correlated, species with larger flowers having more and larger anthers, more ovules, etc. Shrubs were observed to possess the largest, annuals the smallest, and subshrubs medium-sized flowers. Detailed observations of flower structure and function in a shrub (Cistus salvifolius), a spring-annual (Tuberaria guttata) and a winter-annual (T. inconspicua), revealed substantial variations in the breeding system. WhileC. salvifolius (and probably most woody species in this family) presents self-incompatibility, annual species ofTuberaria are self-compatible. In the studied population,T. inconspicua plants bore only reduced, cleistogamous flowers with no sign of a corolla.T. guttata has chasmogamous flowers that can facultatively self-pollinate.     

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.     

Is plasticity across seasons adaptive in the annual cleistogamous plant Lamium amplexicaule?

Background and aims Many angiosperms exhibit cleistogamy, the production of both cleistogamous flowers (CL), which remain closed and obligately self-pollinated, and chasmogamous flowers (CH), which are potentially open-pollinated. The CH proportion can be plastic. Plasticity is adaptive if environmental changes can be reliably assessed and responded to with an appropriate phenotype and if plastic genotypes have higher fitness in variable environments than non-plastic ones.Methods We studied the plastic response of four natural populations from northern and southern France of an annual cleistogamous plant, Lamium amplexicaule, to predictable seasonal variation. Plants were grown in a semi-controlled environment in spring and in autumn. We assessed the variation in flower number, phenology and cleistogamy-related traits, which were all plastic with respect to season. The CH proportion was higher in spring than in autumn in all four populations.Key Results We showed significant stabilizing selection for cleistogamy traits, with higher optimal CH proportions and more pronounced stabilizing selection in spring than in autumn. Observed CH proportions were close to the predicted optimal CH proportions in each season except in autumn for southern populations, which do not experience the autumnal growing season in nature.Conclusions These results are consistent with adaptive plasticity across seasons of cleistogamy in L. amplexicaule. We propose that adaptive plasticity of cleistogamy could be driven by pollination environment variation, with CL flowers providing reproductive assurance when pollinators are scarce and CH flowers reducing the inbreeding depression in offspring when pollinators are abundant.