Cleistogamy inCentaurea melitensis (Asteraceae): Capitulum variability and spatio-temporal development patterns

Centaurea melitensis is a cleistogamous species of Mediterranean distribution, which normally produces a wide range of capitula varying in terms of morphology, size, position in the shoot system and other aspects. For that reason, and particularly in view of the pseudanthic nature of the flower head in Asteraceae, a preliminary study to characterise cleistogamy in this species was considered of interest. Capitular heteromorphism observed inC. melitensis was confirmed by principal component analysis (PCA). The five capitulum types differentiated [initial cleistogamous (iCL), intermediate cleistogamous (xCL), final cleistogamous (fCL), pseudocleistogamous (PCL) and chasmogamous (CH)] fit development patterns specific for both placement on the plant and timing. The balance between cleistogamy and chasmogamy is correlated with the final height of the plant. All these features are consistent with those reported for other cleistogamous species.     

Differential outcrossing rates in dispersing and non-dispersing achenes in the heterocarpic plant Crepis sancta (Asteraceae)

The association between mating systems and dispersal in plants has been studied mostly in cleistogamous species where, generally, seeds produced by cleistogamous (selfed) flowers are less dispersed than seeds produced by chasmogamous (potentially outcrossed) flowers. In heterocarpic Asteraceae, non-dispersing fruits (achenes) are produced at the periphery of the capitulum (outer florets) whereas dispersing achenes are produced by inner florets in the same capitulum. Since all the florets are protandrous, the outer floret developing first are in female phasis when anthesis of inner florets takes place. Thus, outer florets can be potentially selfed by the inner florets of the same capitulum whereas the latter must be pollinated by flowers of other capitula. Therefore outer florets should be more inbred than inner florets. To test this hypothesis, we measured the natural outcrossing rate in outer and inner florets using allozymes in three populations of the heterocarpic Crepis sancta. The results showed that the outcrossing rate was highest for non-dispersed achenes. Moreover, among the outcrossed achenes within a capitulum it was observed that the number of paternal parents of non-dispersing achenes was higher than for dispersing achenes. The pattern observed was therefore the opposite to the pattern of cleistogamous plants and contradicts the putative pollination mechanism we proposed for Asteraceae. The results agree with the predictions of sib competition theory which considers that outcrossing may minimize competitive interactions among relatives (sibs) falling near the mother plant. Higher outcrossing rate in outer florets could also occur because pollinators are more attracted to these florets.     

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.     

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.     

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.     

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.     

The interactive effects of herbivory and mixed mating for the population dynamics of Impatiens capensis

In this study, we examine the demographic consequences of mixed mating and explore the interactive effects of vegetative herbivory and mating system for population dynamics of Impatiens capensis, a species with an obligate mixed mating system (i.e., individuals produce both obligately selfing cleistogamous and facultatively outcrossing chasmogamous flowers). In two natural populations, we followed seeds derived from cleistogamous and chasmogamous flowers subject to different herbivory levels throughout their life cycle. Using a mating system-explicit projection matrix model, we found that mating system types differed in important vital rates. Cleistogamous individuals had higher rates of germination than did chasmogamous individuals, whereas chasmogamous individuals expressed a fecundity advantage over cleistogamous individuals. In addition, population growth was most sensitive to changes in vital rates of cleistogamous individuals, indicating the demographic importance of selfing for these populations. Herbivory also had demographic consequences; a 33%-49% reduction in herbivory caused the population growth rates to increase by 104%-132%, primarily because of effects on vital rates of selfed individuals. Our results not only uncover a novel consequence of mating system expression, that is, mating system influences population dynamics, but also shed light on the role of herbivores in maintaining mixed mating.     

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.     

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.     

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.     

关于四棱草的二型花的研究

陈介(1964)在四棱草属(Schnabelia)中发现具有二型花,即开花授粉型的花(chasmogamous)和闭花授粉型的花(cleistogamous)。他根据当时所掌握的材料,认为二型花"似乎是异株"。笔者在采自安徽祁门的四棱草(S.oligophylla)上发现,二型花无疑是生在同一个体上,但这两种花的花期和花的着生位置是不同的。开花授粉型花的花期早,持续时间短,为一次性开花;闭花授粉型的花是不断地从叶腋发出的,花期持续时间长。     

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.     

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.     

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.     

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.     

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 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.     

Influence of achene heteromorphism on life-cycle traits in the annual weed gallant soldier (Galinsoga parviflora Cav.)

Galinsoga parviflora (gallant soldier), Asteraceae, produces two morphologically distinct achenes in a single capitulum: peripheral and central. The morphological, phenological and generative reproduction traits of the progeny derived from peripheral and central achenes that were cultivated in greenhouse conditions were analyzed. Differences between the progeny of various morph types were manifested at different stages of life. The plants of both morph types developed at a similar rate and they entered key phenological phases at the same time except for the flowering stage. The average height of the studied plants was similar on the same days of the experiment. The study showed that plants from peripheral achenes realize the generative reproduction in other pattern than plants from central achenes. The plants from central achenes produce more capitula per plant at the beginning of the fruiting stage, whereas plants from peripheral seeds achieve the highest number of capitula per plant at later stages. Finally, on day 130 of the experiment, the number of capitula per plant of both morphs equalized. At the early fruiting stage, the number of achenes per capitulum of plants from the two morphs did not differ significantly. The number of achenes within one capitulum decreases with plant age in both populations, but the rate of that decline is greater in the progeny of peripheral seeds. The number of peripheral achenes per capitulum in plants from both achene types decreases at a comparable rate. In contrast, the number of central achenes is reduced at a faster rate in plants germinated from peripheral achenes. After day 83, the individuals from peripheral diaspores intensified their reproductive effort; they produced more capitula but with fewer number of achenes than in individuals from central diaspores. On day 130, the number of dispores per plant of the two morphs equalized. The changes of fecundity of peripheral and central progeny with the age of the plant (at different stages of the life cycle of a single plant) contribute to an intensive seed rain throughout the fruiting period. By producing a large number of less fertile capitula with a stable number of peripheral achenes, plants derived from peripheral achenes are able to supply a higher share of peripheral achenes than if they would produce fewer but more fertile capitula. This strategy increases the pool of peripheral achenes.     

Relationship between capitulum size and pre-dispersal seed predation by insect larvae in common Asteraceae

The evolution of a showy floral display as an advertisement to pollinators could simultaneously advertise the availability of resources to pre-dispersal seed-predators. The hypotheses tested here are that the incidence of seed predation by bud-infesting insect larvae in capitula of Asteraceae is positively related to (1) capitulum size among species, (2) capitulum size within species, (3) capitulum lifespan, and (4) the degree of flowering asynchrony on individual plants. Three populations of each of 20 common herbaceous species of Asteraceae from disturbed ground and grassland habitats were monitored for the presence of pre-dispersal, seed-eating insect larvae. Mean capitulum size (receptacle width) of each species was measured. In a sub-set of eight species, individual capitula were tagged to determine their flowering phenology and lifespan (from anthesis to seed shedding). From these data an index of flowering synchrony on individual plants was derived. Among species, the incidence of larval infestation increased with capitulum size. Small-flowered species such as Achillea millefolium were largely free of bud-infesting larvae, whilst large-flowered species such as Arctium minus were heavily infested. In three cases investigated in greater detail, bud infestation was found to increase with capitulum size within species, suggesting a potential for natural selection to favour smaller capitula. No relationship was found between infestation levels and either capitulum lifespan or degree of flowering synchrony, and there was no evidence that the relationship between capitulum size and infestation was confounded by correlations with these other features. The results support hypotheses 1 and 2, but not 3 and 4. It is suggested that the characteristic capitulum size of each species may represent a trade-off between the opposing selection pressures of pollinators and pre-dispersal seed predators.     

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