宝盖草的繁育系统

宝盖草(Lamium amplexicaule L.)具两种类型的花:短花冠的闭花授粉的花和长花冠的开花授粉的花。本文研究了在3个不同生态环境的自然群体中这两种类型花的数量和开花时间的变化,并通过套袋试验,种子萌发试验和刈割试验,获得了有关开花节律和生活周期以及与环境条件和人类干扰的关系的结果。宝盖草的种子绝大部分是自花授粉的产物,但也有一定程度的异花授粉。自花授粉有利于开拓新的定居环境,而异花授粉则使后代能适应多种环境条件,而这两种授粉方式的结合是使宝盖草成为一种成功的杂草的主要因素。     

INBREEDING EFFECTS IN CLARKIA TEMBLORIENSIS (ONAGRACEAE) POPULATIONS WITH DIFFERENT NATURAL OUTCROSSING RATES

Inbreeding depression is commonly observed in natural populations. The deleterious effects of forced inbreeding are often thought to be less pronounced in populations with self-pollinating mating systems than in primarily outcrossing populations. We tested this hypothesis by comparing the performance of plants produced by artificial self- and cross-pollination from three populations whose outcrossing rate estimates were 0.03, 0.26, and 0.58. Outcrossing rates and inbreeding coefficients were estimated using isozyme polymorphisms as genetic markers. Analysis of F statistics suggests that biparental inbreeding as well as self-fertilization contribute to the level of homozygosity in the seed crop. Biparental inbreeding will reduce the heterozygosity of progeny produced by outcrossing, relative to random outcrossing expectations, and hence will reduce the effects of outcrossing versus self-fertilization. Heterotic selection may increase the average heterozygosity during the life history. Selfed and outcrossed seeds from all three populations were equally likely to germinate and survive to reproduce. However, inbreeding depression was observed in fecundity traits of plants surviving to reproduction in all three populations. Even in the population whose natural self-fertilization rate was 97%, plants grown from seed produced by self-pollination produced fewer fruits and less total seed weight than plants grown from outcrossed seed. There was no detectable inbreeding depression in estimated lifetime fitness. Inbreeding effects for all reproductive yield characters were most severe in the accession from the most outcrossing population and least severe in the accession from the most self-fertilizing population.     

Plantago ser. Hispidulae, a taxonomic revision Knud Rahn

Plantago ser. Hispidulae Rahn belongs to subgen. Psyllium (Juss.) Harms sect. Gnaphaloides Barn. 5 species are recognized, they are all narrow–leaved annuals confined to the area west of the Andes. P. limensis Pers. occurs in Peru; P. litorea Phil., P. hispidula Ruiz & Pav., and P. rancagua Steud. in those parts of Chile with drought during the summer, and P. lundborgii Sparre on the island of San Ambrosio. Flowers are cleistogamic with small corolla lobes and small anthers, very rarely chasmogamic with larger corolla lobes and anthers. Experiments demonstrated that pollination is necessary for development of seeds in cleistogamic flowers, and that transfer of pollen from one such flower to another is almost impossible.     

REPRODUCTIVE BIOLOGY OF SEVERAL BROMEGRASSES (BROMUS): BREEDING SYSTEM,PATTERN OF FRUIT MATURATION,AND SEED SET

Five species of Bromus were tested for self-fertility when bagged. Bromus inermis was self-incompatible. Bromus tectorum, B. kalmii, B. ciliatus, and B. latiglumis were capable of self-fertilization. Further evidence from flowering behavior suggested that B. tectorum almost always self-pollinated and that the remaining self-fertile species sometimes outcrossed. Florets in different positions on a spikelet varied in the proportion that set seed. Open-pollinated B. inermis plants had lower seed set than the self-fertile species, in agreement with the pattern in a variety of hermaphroditic plants.     

Introduced populations of <Emphasis Type="Italic">Genista monspessulana</Emphasis> (French broom) are more dense and produce a greater seed rain in California,USA, than native populations in the Mediterranean Basin of Europe

Some invasive plants perform better in their area of introduction than in their native region, and this is often attributed either to phenotypic responses and/or to adaptive evolution following exposure to new environmental conditions. Genista monspessulana (French broom) is native to Europe, but highly invasive and abundant along the Pacific Coast of the USA. In this study, the population density and age structure, plant growth and reproductive traits, and seed bank characteristics of 13 native (Mediterranean Basin) and 15 introduced (California, USA) field populations of G. monspessulana were compared. Mean population density, plant height and stem diameter were greater in introduced populations, with the latter two traits explained by a greater mean plant age. Age structure also showed a greater percentage of seedling plants in introduced populations. Fecundity was higher in introduced populations when measured in terms of mature seeds per pod, but lower when comparing seed production per plant (number of pods and mature seeds). Thus, seed rain and seed bank size was considerably higher in introduced populations. Results from this study indicate that G. monspessulana performs better in its introduced region. We hypothesize that release from natural enemies and competitors together with more favorable environmental conditions in the introduced region may explain the invasion success of G. monspessulana. As a result, an integrated management approach using introduced seed predators to suppress seed production and selected management practices to reduce seed banks may be needed for effective long-term control in California.     

Seed predation,pathogen infection and life-history traits in Brassica rapa

Herbivory and disease can shape the evolution of plant populations, but their joint effects are rarely investigated. Families of plants of Brassica rapa (Brassicaceae) were grown from seeds collected in two naturalized populations in an experimental garden. We examined leaf infection by the fungus Alternaria, seed predation by a gall midge (Cecidomyiidae) and plant life-history traits. Plants from one population had heavier seeds, were more likely to flower, had less fungal infection, had more seed predation and were more fecund. Fungal infection score and seed predation rate increased with plant size, but large plants still had the greatest number of undamaged fruits. Spatial heterogeneity in the experimental garden was significant; seed predation rate and fecundity varied among blocks. An apparent tradeoff existed between susceptibility to disease and seed predation: plants with the highest fungal infection score had the lowest seed predation rate. Alternaria infection varied between populations, but the disease had no effect on fecundity. Seed predation did reduce fecundity. Damaged fruits had 31.4% fewer intact seeds. However, evidence for additive genetic variation in resistance to seed predation was weak. Therefore, neither disease nor seed predation was likely to be a strong agent of genetically based fecundity selection.     

THE BALANCE BETWEEN CHASMOGAMY AND CLEISTOGAMY IN COLLOMIA GRANDIFLORA (POLEMONIACEAE)

The extent of cleistogamy (CL) and chasmogamy (CH) was investigated in natural populations of Collomia grandiflora Dougl. ex Lindl. The ubiquitous occurrence of CL flowers was documented in 18 populations from throughout the geographic range of the species. The interrelationship of environmental perturbation, vegetative weight, and the balance between CH and CL flowers was investigated by means of a factorial experiment which varied plant density, incident light and soil mix. An ANOVA revealed that total plant weight increased significantly under conditions of low density, low light and low sand content, with density accounting for most of the variance in plant weight. Interactions involving density were not significant. The number of CH and CL flowers and estimated percent outcrossing increased significantly with increased plant weight, low density, low light, and low sand content, although an analysis of covariance showed that plant weight and density accounted for most of the variance in these traits. Among large, multibranched plants, the proportion of CL flowers increased in a basipetal pattern, whereas most of the CH flowers were restricted to the terminal and upper inflorescences. The smallest plants produced only a single terminal inflorescence comprising entirely CL flowers. The proportion of reproductive effort allocated to outcrossing was dependent upon plant size as estimated from total weight, whereas the proportion allocated to ***selfing was relatively independent. Studies of outcrossed vs. inbred progenies of five populations revealed little evidence for immediate inbreeding depression with respect to plant weight and the number of CH or CL flowers, but indicated that significant quantitative genetic differences exist for these traits among populations of diverse ecological or geographical origin.     

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

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

Adaptive significance of self-fertilization in a hermaphroditic perennial, Trillium camschatcense (Melanthiaceae)

The evolution of self-fertilization from primarily outcrossing ancestors is one of the most common evolutionary transitions in plants; however, the ecological mechanisms that maintain self-fertilization have remained controversial. Theoretical studies suggest that selfing is advantageous over outcrossing in terms of genetic transmission and assurance of seed production under pollen-limited circumstances. Trillium camschatcense is a herbaceous perennial distributed in Hokkaido and northern Honshu, Japan. Geographical variation in the breeding system (self-compatible, SC; or self-incompatible, SI) has been reported in populations in Hokkaido. Here, we used several SC and SI populations of T. camschatcense to investigate the adaptive significance and the evolutionary basis of self-fertilization. Pollination experiments and genetic analyses demonstrated that the potential availability of outcross pollen in SC populations was sufficient and that the number of pollen donors was equal to that of SI populations. However, despite the high availability of outcross pollen, the SC populations produced seeds predominantly by selfing and so underwent severe inbreeding depression. Although none of the suggested advantages for self-fertilization were supported by our analyses, we propose two possible scenarios for the evolution of self-fertilization in T. camschatcense.     

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

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

Comparison of genetic variation in populations of wild rice, Oryza rufipogon, plants and their soil seed banks

Wild rice, Oryza rufipogon, has endangered species conservation status and it is subject to in situ conservation in China. To understand the potential of the seed bank in species conservation and population restoration, this study compared the genetic diversity of O. rufipogon plants with that of its soil seed banks in two marshes. A total of 11 pairs of rice SSR primers were used and 9 were polymorphic. Allele frequencies of the seeds differed significantly from those of surface plants and varied between soil layers. Relatively more alleles and higher genetic diversity (H _e) were found in plant populations, relative to seed banks. The numbers of germinable seeds and the level of genetic variation in seed banks decreased with the increasing of soil depth, indicating a rapid seed loss. Genetic differentiation was detected between sites and between plant and seed populations, as well as among seeds of different soil strata. Rapid seed loss, partly dormancy loss, and nonrandom seed mortality are discussed as the possible contributors to the pattern of reduced genetic variation within seed banks, compared to plants. These could also be responsible for the considerable genetic differentiation between populations. The seed population held about 72% of the total genetic variation of O. rufipogon in each marsh, indicating the potential of seed banks for restoring population variabilities if the plant populations were lost.     

Plant reproductive strategies vary under low and high pollinator densities

Long‐term variation in the population density of honey bees Apis mellifera across landscapes has been shown to correlate with variation in the floral traits of plant populations in these landscapes, suggesting that variations in pollinator population density and foraging rates can drive floral trait evolution of their host plants. However, it remained to be determined whether this variation in plant traits is associated with adaptive variation in plant reproductive strategies under conditions of high and low pollinator densities. Here we conducted a reciprocal transplant experiment to examine how this variation in floral traits, under conditions of either high and low pollinator density, impacted seed production in the Tibetan lotus Saussurea nigrescens. In 2014 and 2015, we recorded the floral traits, pollinator visitation rates, and seed production of S. nigrescens populations grown in both home sites and foreign sites, where sites varied in honey bee population density. Our results demonstrated that the floral traits reflected those of their original population, regardless of their current location. However, seed production varied with both population origin and transplant site. Seed number was positively correlated with flower abundance in the pollinator‐rich sites, but with nectar production in the pollinator‐poor sites. Pollinator visitation rate was also positively correlated with flower number at pollinator‐rich sites, and with nectar volume at pollinator‐poor sites. Overall, the local genotype had higher seed production than nonlocal genotypes in home sites. However, when pollen is hand‐supplemented, plants from pollinator‐rich populations had higher seed production than plants from pollinator‐poor populations, regardless of whether they were transplanted to pollinator‐rich or ‐poor sites. These results suggest that the plant genotypic differences primarily drive variation in pollinator attraction, and this ultimately drives variation in seed: ovule ratio. Thus, our results suggest that flowering plant species use different reproductive strategies to respond to high or low pollinator densities.     

Reproductive assurance through self-fertilization does not vary with population size in the alien invasive plant Datura stramonium

Autonomous self-fertilization is suggested to be associated with invasiveness in plants because it offers reproductive assurance when there is a shortage of suitable mates or pollinators. Given that shortages of mates and pollinators are a common cause of Allee effects in small plant populations, we predict that the benefits of self-fertilization in terms of reproductive assurance should be greatest in small populations. We tested this idea for the invasive herb Datura stramonium , a self-fertilizing species which is also cross-pollinated to some extent by insects (mainly hawkmoths and honeybees). During two consecutive years, we studied 20 and 55 populations, respectively, of different sizes. Untreated flowers of D. stramonium showed high levels of fruit and seed set in all populations studied. Although, fruit and seed set were generally reduced by about 90% in flowers in which self-fertilization was prevented through emasculation, this effect did not vary according to population size. By using a natural color (anthocyanin) dimorphism in 12 populations, we showed that the average outcrossing rate was low (1.3%) and that there was no relationship between outcrossing rate and population size. Pollen removal from flowers also did not vary according to population size, suggesting that the pollinator visitation rate is not lower in small populations. However, decreasing deviations of observed from expected fruit set with population size imply that small populations may have an increased chance of extinction due to demographic stochasticity. Overall, our results suggest that reproductive assurance through self-fertilization in invasive plants may be important for all stages of population establishment, and not just in the founder population.     

Isolation affects reproductive success in low-density but not high-density populations of two wind-pollinated <Emphasis Type="Italic">Thalictrum</Emphasis> species

Since pollen usually travels limited distances in wind-pollinated plant species, plants growing at low density may become pollen limited. We examined how local pollen availability and population density affect reproductive success in two wind-pollinated, dioecious species, Thalictrum fendleri and Thalictrum dioicum. Distance to the nearest flowering male, the number of flowering males within 2 m, and flower number on those males served as measures of local pollen availability. Increased distance from pollen donors reduced seed set in the lowest-density population of each species, but seed set in high-density populations was not correlated with local pollen availability. For plants in high- and low-density populations at similar distances from pollen donors, this distance only affected seed set in low-density populations. To ensure that differences in resource availability were not causing spurious correlations between seed set and plant density, we constructed low-density artificial arrays in populations of T. dioicum. In these, seed set decreased rapidly with increases in distance from pollen donors. Despite these effects, the density of males in a population was not correlated with average seed set in T. dioicum, and hand pollination in the T. dioicum populations also failed to increase seed set over natural levels. These results suggest that pollen receipt only limits seed set on isolated plants within low- density populations of T. dioicum and T. fendleri.     

Floral density,pollen limitation,and reproductive success in<Emphasis Type="Italic"> Trillium grandiflorum</Emphasis>

Decreases in floral density can disrupt mutualistic interactions between plants and their pollinators, and decrease reproductive success. I addressed the relationship between floral density and plant reproductive success using two experimental approaches: a pollen supplementation experiment in 12 populations of Trillium grandiflorum that naturally varied in floral density, and a transplant experiment in which floral density was manipulated in plots at four experimental sites. In the pollen supplementation experiments, the degree of pollen limitation, in terms of fruit set and seed set, decreased with floral density. Further, in the experimental sites, plant reproductive success increased asymptotically with floral density. These results demonstrate the value of simultaneously conducting experiments in both experimental sites and natural populations to understand how population density influences plant reproductive success. Factors that reduce the density of this perennial herb, such as habitat fragmentation and herbivory by white-tailed deer (Odocoileus virginianus), should be expected to limit its reproduction.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

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.     

GAMETOPHYTIC SELF-FERTILIZATION IN HOMOSPOROUS PLANTS: DEVELOPMENT,EVALUATION, AND APPLICATION OF A STATISTICAL METHOD FOR EVALUATING ITS IMPORTANCE

The methods described here make it possible to use data on sporophytic genotype frequencies to estimate the frequency of gametophytic self-fertilization in populations of homosporous plants. Bootstrap bias reduction is effective in reducing or eliminating the bias of the maximum likelihood estimate of the gametophytic selfing rate. The bias-corrected percentile method provides the most reliable confidence intervals for allele frequencies. The percentile method gives the most reliable confidence intervals for the gametophytic selfing rate when selfing is common. The maximum likelihood intervals, the percentile intervals, the bias-corrected percentile intervals, and the bootstrap t intervals are all overly conservative in their construction of confidence intervals for the gametophytic selfing rate when self-fertilization is rare. Application of the recommended methods indicates that gametophytic self-fertilization is quite rare in two sexually reproducing populations of Pellaea andromedifolia studied by Gastony and Gottlieb (1985).     

Density-dependent flowering phenology,outcrossing, and reproduction in Impatiens capensis

Summary We investigated the effects of plant density on cleistogamous (CL) and chasmogamous (CH) flowering phenology and seed production in a natural Impatiens capensis population, by censusing individually marked plants at experimentally reduced and natural densities. CL flowering was earlier at natural density. This plastic density response may have resulted from a stress-related threshold for CL flowering; slower growing plants at natural density flowered earlier. Although apparently triggered by slow early growth, early CL flowering also involved an additional cost for later growth rate. In contrast, CH flowering was unrelated to relative growth rate, but apparently required a size threshold. Experimental density reduction resulted in earlier CH flowering and a dramatic increase in the percentage of plants producing CH flowers. Individual CL and CH flowering duration and flower production were greater at reduced density. These density-dependent effects caused differences between treatments in the shape and location of population flowering phenology curves. Moreover, the percentage of CH seeds produced per individual was much higher at reduced density. At natural density total seed production per plant was lower and more hierarchical than at lower density, suggesting that dominance and suppression shape jewelweed fitness distributions.     

CORRELATED EVOLUTION OF SELF-FERTILIZATION AND INBREEDING DEPRESSION: AN EXPERIMENTAL STUDY OF NINE POPULATIONS OF AMSINCKIA (BORAGINACEAE)

The relation between inbreeding depression and rate of self-fertilization was studied in nine natural populations of the annual genus Amsinckia. The study included two clades (phylogenetic lineages) in which small-flowered, homostylous populations or species are believed to have evolved from large-flowered, heterostylous, self-compatible ones. In one lineage the small-flowered species is tetraploid with disomic inheritance. Rates of self-fertilization were 25% to 55% in the four large-flowered, heterostylous populations; 72% in a large-flowered but homostylous population; and greater than 99.5% in the four small-flowered, homostylous populations, which produce seed autonomously. When present, inbreeding depression occurred in the fertility but not the survival components of fitness. Using a cumulative fitness measure incorporating both survival and fertility (flower number), we found inbreeding depression to be lower in the four very highly self-fertilizing populations than in the five intermediate ones. The Spearman rank correlation between inbreeding depression and selfing rate for the nine populations was –0.50, but was not statistically significant (P = 0.12). Inbreeding depression was greater in the two tetraploid populations than in the very highly self-fertilizing, diploid ones. Phenotypic stability of progeny from self-fertilization tended to be higher in populations with lower inbreeding depression. We conclude that levels of self-fertilization and inbreeding depression in Amsinckia are determined more by other factors than by each other. Estimates of mutation rates and dominance coefficients of deleterious alleles, obtained from a companion study of the four highly self-fertilizing populations, suggest that a strong relationship may not be expected. We discuss the relationship of the present results to current theory of the coevolution of self-fertilization and inbreeding depression.     

EFFECTS OF POSTDISPERSAL SEED PREDATION ON SPATIAL INEQUALITY AND SIZE VARIABILITY IN AN ANNUAL PLANT,ERODIUM CICUTARIUM (GERANIACEAE)

By decreasing seed density, ants introduced into flats of uniformly sown seeds of Erodium cicutarium (Geraniaceae) created differences in the neighbor-free area available to individual plants. The changes in spatial patterns brought about by the ants were greater when a higher proportion of seeds was removed but were independent of initial seed density. These spatial changes and differences in seed density were examined for their effects on plant size and reproduction. Gini values were calculated to determine inequalities. As the inequality in space among individual plants increased, the variation in final biomass increased. The number of individuals reproducing was constant among treatments, and yet seed production per plant was significantly greater for populations in which the spatial pattern was influenced by seed predation. The decrease in density and changed spatial pattern, due to previous seed predation, resulted in a few individuals having much more space than others and consequently producing many more seeds. The increase in reproductive effort per flat was much greater than could be explained by the changing density alone. Our experiment demonstrates that spatial inequality, such as that generated by seed predators, can be more important than density in generating size inequalities in plant populations. This result can profoundly alter the competitive interactions between plants and determine which plants produce seed for the next generation.