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.     

荒漠植物角果藜的地上地下结果性

对分布于准噶尔荒漠中角果藜(Ceratocarpus arenarius)果实的形态特征、空间位置、发育特性及散布特性进行了比较研究,结果表明:角果藜具有地上地下结果性,地上果实和地下果实在颜色、形状、大小、质量、数量、空间位置、成熟期和散布特性上均有显著差异。角果藜地上地下结果性是植物适应荒漠多变环境所具有的一种特殊的繁殖方式。这种独特的繁殖方式增加了物种在荒漠极端环境中成功生存的机率,保证了物种的顺利繁衍。这是首次在藜科植物中发现的地上地下结果性现象。     

On the ecology and evolution of annual plants in disturbed environments

This paper is concerned with the effect of disturbance on some crucial characteristics of annual plants. The theoretically optimal life-history traits that maximize individual fitness in disturbed environments are described and critically evaluated. It seems that none of them holds for all annual species.Self-pollination and especially seed polymorphism are considered important adaptations to life in unpredictable environments. The thesis is put forward that amphicarpic annuals, which exhibit both self-pollination and extreme seed polymorphism, are best adapted to life in hazardous habitats. The hypothetical course of the evolution of amphicarpy is demonstrated on the grounds of the comparison of contemporary annual species producing chasmogamous and cleistogamous flowers on a single individual.     

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.     

Disentangling sources of maternal effects in the heterocarpic species Rumex bucephalophorus

Maternal effects are one of the most important influences on the offspring phenotype and their adaptive significance has been increasingly recognized. In this study we document the role of two sources of maternal effects, position and heterocarpy, on juvenile and adult traits of Rumex bucephalophorus, an annual plant that occurs in heterogeneous and unstable environments. Up to three different diaspore types can be found in populations of this species: buried diaspores (BD) at the plant base, and fixed diaspores (FD) and long dispersible diaspores (LD) both borne along aerial stems. The two sources of maternal effects affected most stages of the life cycle of R. bucephalophorus: seed size, dispersability, seedling size and survival, and adult plant fecundity. Maternal effects due to position affected mainly juvenile traits, while those due to heterocarpy were stronger and more persistent. In general, seedlings from FD showed higher survival and originated plants with higher fecundity but with lower dispersability than those derived from BD and LD. Dispersability was markedly affected by the two maternal effects at two different stages: first, there was a trend to place the slighter and more easily detachable diaspores (LD) at the optimal positions for dispersal, and second maternal effect due to heterocarpy surprisingly affected dispersability of the second generation progeny. Our study shows that maternal effects can play an important role in ecology, dynamic and evolution of populations and it adds to growing evidence about the important role of maternal effects.     

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.     

Reproductive versatility in legumes: the case of amphicarpy in Trifolium polymorphum

Amphicarpy is a fascinating reproductive strategy, defined as fruit produced both below the soil surface and as aerial fruit on the same plant. Trifolium polymorphum is a grassland species subject to herbivory that combines amphicarpy with vegetative reproduction through stolons. Underground flowers have been described as obligate autogamous and aerial ones as self‐compatible allogamous, with aerial floral traits favouring cross‐pollination. In the present work we performed different pollination treatments on aerial flowers to analyse rates of pollen tube development and offspring fitness, measured as fruit set, seed production and germination percentage. This last variable was compared to that of seeds produced underground. No significant differences were found between fruit set in self‐ and cross‐pollinations. Seed production was higher in self‐pollinations, which is consistent with the higher rate of pollen tube development observed in self‐crosses. Spontaneous self‐pollination is limited in aerial flowers; thus pollen transfer by means of a vector is required even within the same flower. Germination tests showed that aerial seeds produced after self‐ and cross‐pollination did not differ in fitness, but underground seeds had higher germination percentage than aerial ones. Thus, we conclude that T. polymorphum has a mixed mating system. In grasslands with heavy grazing pressure, clonal propagation and underground seed production ensure persistence in the field. An intermediate level of selfing in aerial flowers ensures offspring, but morphological (herkogamy) and functional (dicogamy) floral traits maintain a window to incorporate genetic variability, allowing the species to tolerate temporal and spatial pressures.