Demographic consequences of greater clonal than sexual reproduction in Dicentra canadensis

Clonality is a widespread life history trait in flowering plants that may be essential for population persistence, especially in environments where sexual reproduction is unpredictable. Frequent clonal reproduction, however, could hinder sexual reproduction by spatially aggregating ramets that compete with seedlings and reduce inter‐genet pollination. Nevertheless, the role of clonality in relation to variable sexual reproduction in population dynamics is often overlooked. We combined population matrix models and pollination experiments to compare the demographic contributions of clonal and sexual reproduction in three Dicentra canadensis populations, one in a well‐forested landscape and two in isolated forest remnants. We constructed stage‐based transition matrices from 3 years of census data to evaluate annual population growth rates, λ. We used loop analysis to evaluate the relative contribution of different reproductive pathways to λ. Despite strong temporal and spatial variation in seed set, populations generally showed stable growth rates. Although we detected some pollen limitation of seed set, manipulative pollination treatments did not affect population growth rates. Clonal reproduction contributed significantly more than sexual reproduction to population growth in the forest remnants. Only at the well‐forested site did sexual reproduction contribute as much as clonal reproduction to population growth. Flowering plants were more likely to transition to a smaller size class with reduced reproductive potential in the following year than similarly sized nonflowering plants, suggesting energy trade‐offs between sexual and clonal reproduction at the individual level. Seed production had negligible effects on growth and tuber production of individual plants. Our results demonstrate that clonal reproduction is vital for population persistence in a system where sexual reproduction is unpredictable. The bias toward clonality may be driven by low fitness returns for resource investment in sexual reproduction at the individual level. However, chronic failure in sexual reproduction may exacerbate the imbalance between sexual and clonal reproduction and eventually lead to irreversible loss of sex in the population.     

食虫植物光萼茅膏菜的生物学研究

通过２年野外观察和栽培实验,对光萼茅膏菜的生活史特征、种群动态以及食虫习性进行了初步研究。结果表明：光萼茅膏菜杰早春植物;该种以球茎行营养繁殖,虽然种子产量较大,但实生苗更新能力较低;花期植物捕虫率最高;食虫对植物生长和繁殖有一定益处。     

Analyses of spatial patterns and population processes of clonal plants

The nonrandom spatial structure of terrestrial plants is formed by ecological interactions and reproduction with a limited dispersal range, and in turn this may strongly affect population dynamics and population genetics. The traditional method of modelling in population ecology is either to neglect spatial pattern (e.g. in transition matrix models) or to do straightforward computer simulation. We review here three analytical mothods to deal with plant populations in a lattice-structured habitat, which propagate both by seeds that scatter over the whole habitat and by vegetative reproduction (producing runners, rhizomes, etc.) to neighboring vacant sites. [1]Dynamics of global and local densities: Dynamical equations of population density considering nearest-neighbor correlation (spatial clumping) are developed as the joint dynamics of global average density and local density (comparable to mean crowding) based onpair approximation. If there is a linear trade-off between seed production and vegetative reproduction, the equilibrium abundance of the population may be maximized by engaging both means of reproduction. This result is accurately predicted by the pair approximation method, but not by mean-field approximation (neglect of spatial structure). [2]Cluster size distributions: Using global and local densities obtained by pair approximation, we predicted cluster size distribution, i.e. the number of clusters of occupied sites of various sizes. [3]Clonal identity probability decreasing with distance: Multi-locus measurement of allozymes or other neutral molecular markers tells us whether or not a given pair of individuals belong to the same clone. From the pattern of clonal identity probability decreasing with the distance between ramets, we can estimate the relative importance of two modes of reproduction: vegetative propagation and sexual seed production.     

Evaluation of the population dynamics of the forage legume (Lotus corniculatus), using matrix population models

The population dynamics of perennial crop plants are influenced by numerous factors, including management practices. Conditions in the field vary from year to year, and matrix population models are useful for evaluating population behaviour in relation to environmental variability. In Missouri, the stand persistence of birdsfoot trefoil ( Lotus corniculatus ), a perennial legume, is often limited by disease and poor seed production. A stage-based, matrix population model was developed to evaluate the population dynamics of birdsfoot trefoil in relation to clipping treatment. The plant growth stages represented in the model were seeds, seedlings, mature vegetative and reproductive plants. Two phases of population growth were evaluated in clipped and unclipped stands. Establishment-phase populations were characterized by relatively high mortality and low reproduction. Elasticity analysis indicated that growth of these populations was most sensitive to the survival of vegetative plants. Mature vegetative plants and seeds comprised the majority of surviving individuals in clipped and unclipped populations, respectively; however, establishment-phase populations under both management treatments tended toward extinction. Populations in the post-establishment phase of growth were characterized by relatively low mortality and high reproduction. Population growth in this phase of growth was most sensitive to seed production, and most individuals in these populations were at the seed stage.     

Optimum reproduction and dispersal strategies of a clonal plant in a metapopulation: a simulation study with Hieracium pilosella

Clonal spread is favoured in many plants at the expense of seed production in order to expand rapidly into open habitats or to occupy space by forming dense patches. However, for the dynamics of a population in a patchy landscape seed dispersal remains important even for clonal plants. We used a spatially explicit individual-based metapopulation model to examine the consequences of two trade-offs in Hieracium pilosella L: first, between vegetative and sexual reproduction, and second, between short and far-distance dispersal of seeds. Our main question was, what are the environmental conditions that cause a mixed strategy of vegetative and sexual reproduction to be optimal. The model was parameterised with field data on local population dynamics of H. pilosella. Patch dynamics were given firstly by disturbance events that opened patches in a matrix of a clonal grass that were colonisable for H. pilosella, and secondly by the gradual disappearance of H. pilosella patches due to the expanding grass. Simulations revealed opposing selection pressures on traits determined by the two trade-offs. Vegetative reproduction is favoured by local dynamics, i.e. the need for maintenance and expansion of established populations, whereas seed production is favoured by the necessity to colonise empty habitats. Similar pressures act on the proportion of seeds dispersed over short and far distances. Optimum reproductive and dispersal strategies depended on habitat quality (determined by seedling establishment probability), the fraction of dispersed seeds, and the fraction of seeds lost on unsuitable ground. Under habitat conditions supporting moderate to low seedling establishment, between 20% and 40% of reproductive effort in H. pilosella should be devoted to sexual reproduction with at least 10% of the seeds dispersed over distances suitable to attain empty patches. We conclude that in a spatially heterogeneous landscape sexual seed production in a clonal plant is advantageous even at the expense of local vegetative growth.     

Current and historical factors drive variation of reproductive traits in unisexual mosses in Europe: A case study

Unisexual bryophytes provide excellent models to study the mechanisms that regulate the frequency of sexual versusasexual reproduction in plants, and their ecological and evolutionary implications. Here, we determined sex expression, phenotypic sex ratio, and individual shoot traits in 242 populations of the cosmopolitan moss Pseudoscleropodium purum spanning its whole distributional range. We tested whether niche differentiation, sex-specific differences in shoot size, and biogeographical history explained the spatial variation of reproductive traits. We observed high levels of sex expression and predominantly female-biased populations, although both traits showed high intraspecific variation among populations. Sex expression and sex ratio were partly explained by current macroscale environmental variation, with male shoots being less frequent at the higher end of the environmental gradients defined by the current distribution of the species. Female bias in population sex ratio was significantly lower in areas recolonized after the last glacial maximum (recent populations) than in glacial refugia (long-term persistent populations). We demonstrated that reproductive trait variation in perennial unisexual mosses is partially driven by macroscale and historical environmental variation. Based on our results, we hypothesize that sexual dimorphism in environmental tolerance and vegetative growth contribute to sex ratio bias over time, constraining the chances of sexual reproduction, especially in long-term persistent populations. Further studies combining genetic analyses and population monitoring should improve our understanding of the implications of the intraspecific variation in the frequency of sexual versusasexual reproduction in bryophyte population fitness and eco-evolutionary dynamics.     

Lattice population dynamics for plants with dispersing seeds and Vegetative propagation

The population dynamics of plants in a lattice structured habitat are studied theoretically. Plants are assumed to propagate both by producing seeds that scatter over the population and by vegetative reproduction by extending runners, rhizomes, or roots, to neighboring vacant sites. In addtion, the seed production rate may be dependent on the local density in the neighborhood, indicating beneficial or harmful crowding effects. Two sets of population dynamical equation(s) are derived: one based onmean-field approximation and the other based onpair approximation (tracing both global and local densities simultaneously). We examine the accuracy of these approximate dynamics by comparing them with direct computer simulation of the stochastic lattice model. Pair approximation is much more accurate than mean-field approximation. Mean-field approximation overestimates the parameter range for persistence if crowding effects on seed production are harmful or weakly beneficial, but underestimates it if crowding effects are highly beneficial. Dynamics may show bistability (both population persistence and extinction) if the effect of crowding is strongly beneficial. If there is a linear trade-off between seed production and vegetative reproduction, the equilibrium abundance of the population may be maximised by a mixture of seed production and vegetative reproduction, rather than by pure seed production or by pure vegetative reproduction. This result is correctly predicted by pair approximation but not by mean-field approximation.     

Variation and covariation among life-history traits in Rumex acetosella from a successional old-field gradient

In this study morphological variation and the potential for competition to affect biomass and seedling selection of the families of five populations of Rumex acetosella L. sampled along a successional old-field gradient have been investigated. Seeds from 25 families were submitted to four competitive regimes: no competition (one plant per pot), medium competition (two plants/ pot taking plants from the same population), high within-population competition (four individuals from the same population in a pot) and high between-population competition (four individuals from two different populations in a pot). Eight traits were analysed after 3 months of growth for variation among families within populations. A significant difference among families within the two older populations was recorded for sexual biomass and related components. High sensitivity of these traits to density was observed in all populations except the youngest, suggesting specialization to particular environmental conditions in late successional populations, and a good adaptive capacity to buffer environmental variation in the pioneer population. Little significant interaction between competitive regimes and families within populations was found, i.e. genotypes within each population showed little variation in their response to environmental variation. Genotypic variance decreased with increasing competitive conditions for the majority of the traits. However, the percentage of variance in sexual reproduction explained by family was stable among treatments. Tradeoffs between vegetative reproduction and sexual reproduction were recorded at the population level along the successional gradient, with increasing competitive conditions. As succession proceeds, we observed a decrease in sexual reproduction and an increase in vegetative reproduction. At the family level, correlation among traits were similar when plants were grown in the absence of competition and at high density, with a significant negative correlation between sexual reproduction and vegetative reproduction. For both sprout number and sexual biomass, the performance of families grown under all the treatments was positively correlated. Together these results indicate allocational constraints on the reproductive biology of R. acetosella that may be favoured by natural selection and have influenced population differentiation along the successional gradient. However, they also revealed that the potential exists for evolutionary specialization through plasticity, in response to variation in environmental conditions.     

Biology of Elodea canadensis mich. and its management in Australian irrigation systems

The seasonal growth and decline of a population of Elodea canadensis Mich. growing in an irrigation drain are described, together with some characteristics of the aquatic environment (turbidity, photosynthetically-available radiation, temperature and pH). Overwintering buds (up to 5000 m^?2) in the form of swollen dormant stem apices are produced in autumn with the onset of cold weather, remain in the mud, and grow out in the spring. In late summer vegetative reproduction also occurs when large numbers of the leafless stem portions which are capable of growing into independent plants are swept downstream from established populations. Results suggest that control measures should be applied in early summer when populations are approaching nuisance size, and again in late summer before fragmentation occurs and overwintering propagules are initiated. In irrigation channels in Australia, where draining and drying are not feasible, biomass in successfully reduced by widespread use of herbicides.     

Population dynamics in Oxalis acetosella: the significance of sexual reproduction in a clonal, cleistogamous forest herb

I studied demography in three populations of the clonal, cleistogamous herb Oxalis acetosella during three growing seasons, to assess the impact of seedling recruitment relative to ramet recruitment on its population dynamics. I followed each plant in permanent plots, and stage-classified it according to origin (sexually or vegetatively derived) and reproductive status. I calculated between-year transition probabilities, and sexual and vegetative fecundities, and used them in projection matrix analyses to simulate the development of populations and obtain the elasticity of each life-history event.
In general, non-flowering adults was the most numerous stage class and flowering new ramets the rarest, whereas the proportions of the other stage classes varied among populations and years. The negative correlation between plant density and survival was not very strong, and mortality seemed to be largely density-independent. Simulated population growth rates (λ) indicated that one of the populations is declining and the other two are growing, although slowly. The highest elasticities were usually those of stasis in adult plants, whereas elasticities of other matrix entries were low and highly variable. λ was positively correlated with the elasticities of seedling production and survival but negatively correlated with the elasticity of adult survival, indicating that seedling recruitment is more favourable for population growth than is adult stasis. There were no correlations between λ on the one hand and the elasticities of ramet production and survival on the other.
The results suggest that recruitment from seedlings is important for the growth and long-term maintenance of O. acetosella populations, as a complement to the rather low vegetative propagation. In this context cleistogamy is an adaptive strategy, maximizing total seed output.     

浙江天目山苦槠种群结构和动态研究

选取了6个代表性样地对浙江天目山的苦槠种群结构和动态进行了研究,研究结果表明：苦槠种群的结构为衰败型：种群结构存在着幼龄种苗数量较少,部分样地有缺失现象。种群结构总体分布格局为随机型,不同阶段幼苗—幼树—中树—大树动态格局趋势表现为聚集—聚集—随机过渡—随机型。造成这种现象的原因一方面是由于大量的结实种子易被啮齿类捕食所致;另一方面也与苦槠的生物学特性和生境密切相关。苦槠种群维持和更新繁殖方式的适应性特点表现为无性萌生繁殖贡献率远远大于有性繁殖,种群主要通过萌生的方式来维持种群的更新和存活,并在一定程度上影响了演替的方向。衰败型苦槠种群的生态恢复可通过控制鼠害和移栽幼苗的策略。     

Differences in vital demographic rates in three populations of the epiphytic bromeliad,Werauhia sanguinolenta

Since the response to differences in resource availability is most pronounced in smaller individuals of vascular epiphytes such as Werauhia sanguinolenta Cogn. et Marchal (Bromeliaceae), I expected variation in growth and survival of small individuals to play an important role in the dynamics of entire populations. Four annual censuses (2002–2005) of three study populations, which were located across the isthmus of Panama, allowed me to construct stage transition matrices, and to conduct growth analysis and elasticity analysis. Differences between populations were highly consistent through time, but, contrary to expectations, hardly related to the comportment of smaller plants. For example, although average mortality rates were highest at the driest site, close to the Pacific, small plants were not predominantly affected. Similarly, although the highest relative growth rates (RGR) of individuals and the highest population growth rates (λ) were found in the population with the highest moisture input, which was located close to the Atlantic coast, this was not due to a particularly strong stimulation of RGR in small plants. Elasticity analysis indicated rather small differences in the importance of the three demographic processes growth, survival, and reproduction for population growth in the three populations, but invariably identified the survival of large tanks as the single most important process determining λ.     

加拿大一枝黄花的入侵机理研究进展

我国是世界上遭受外来生物入侵最为严重的国家之一。加拿大一枝黄花(Solidago canadensis L.)是一种世界性的入侵植物,侵入我国已有70多年的历史,目前国内外对其入侵机理已有许多研究。结合近年来的研究热点和重要进展,从加拿大一枝黄花的生物学特点、入侵地环境的可入侵性、化感效应以及植物与土壤的反馈作用等4个方面综述了其成功入侵的机理,分析和评价了各种机理之间的内在联系。认为生物入侵是一个涉及多个营养级的复杂生物学过程,是多种机制综合作用的结果,人类活动的干扰是入侵事件频发的深层次原因。加拿大一枝黄花强大的入侵能力、高度的抗逆性、对本地植物和土传病原菌的化感抑制、入侵地环境的脆弱性以及菌根真菌等有益微生物的协助在入侵过程中起着关键作用,但天敌释放并未使其获得更大竞争优势。     

The relationship between climatic conditions and generative reproduction of a lowland population of Pulsatilla vernalis: the last breath of a relict plant or a fluctuating cycle of regeneration?

The alpine-boreal plants which have survived in Central European lowlands during glacial periods depend both on the preservation of their refugial habitat, and their capability for vegetative and generative reproduction and dispersion. Pulsatilla vernalis (L.) Mill. is a model species which occurs throughout most of the European alpine system, as well as in isolated populations in the lowlands. At present, the relict lowland localities of this species often have a historic character. In the Polish lowlands, only the population located in Rogowiec is characterized by effective recruitment. It presents a large number of rosettes and a diversified demographic structure, with plants in all stages of development. The study examines the population in Rogowiec from 2002 to 2008 with regard to the number of flowering and fruiting shoots, new seedlings, and loss of juvenile rosettes, and the obtained data are correlated with climatic conditions. Three years were found to demonstrate effective recruitment, with numerous seedlings and little loss of juvenile rosettes. No significant relationship was found between seedling dynamics and the flowering–fruiting process. However, correlations were found between effective renewal and some climatic factors. Temperature, water balance, and solar radiation were found to have a limiting effect on the reproduction and regeneration of the analyzed relict population. Due to the longevity of the rosettes, favorable climatic conditions occurring every few years are sufficient for survival of the species in this location. However, serious threats are posed by the climate change trends expected in Europe over the coming decades, and the fact that due to its small-scale dispersal ability, the analyzed species occupies only a small area in Rogowiec.     

An allelopathic investigation of the domination of the introduced invasive Conyza canadensis L.

The introduced, invasive species Conyza canadensis L. covers large areas of the sandy levees next to the River Tamiš (Serbia), forming dense microcomplexes and dominating the other herbaceous species in the ruderal phytocoenosis with its aboveground mass and abundance. In addition to this species, a further 28 plant species were found, but the abundance and cover of these was significantly lower. The allelopathic influence of the species C. canadensis was investigated through analyzing the total phenolics and phenolic acids, as the main allelochemicals, in dead and vegetative parts and the soil beneath them. Seed germination and seedling growth of the target plants (Dactylis glomerata L. and Trifolium repens L.), which grow in this community, served as a measure of the inhibitory capacity of this species. It was established that the content of total phenolics and phenolic acids (p-coumaric, ferulic, p-hydroxybenzoic, vanillic and syringic) varies, following the order: vegetative plant parts > dead plant parts > sandy soil under C. canadensis. Water leachate and soils inhibited seed germination and seedling growth of the test plants to varying degrees, following the order: vegetative parts > dead parts > sandy soil, which is directly related to the content of total phenolics and phenolic acids in them. It was concluded that the pioneer species C. canadensis plays a decisive role in the first phases of vegetation succession and the process of soil formation on the barren sandy levees, owing to the synthesis of secondary phenolic metabolites.     

Pollen flow and the intrapopulation genetic structure of Heloniopsis orientalis on the forest floor as determined using microsatellite markers

Pollenflow, seedling dynamics, and the genetic structure of Heloniopsis orientalis (Thunb.) C. Tanaka (Liliaceae) were analyzed as part of an effort to understand the reproduction system of perennial herbs foundon the forest floor by means of microsatellite markers. It was possible to assign paternity to all offspring within this population by paternity analysis. Although no ordinary pollinator visits were observed during the flowering period, effective pollen flow occurred throughout the population. It appears that low frequencies of pollinator visits nonetheless permitted adequate pollen flow. Studies of seedling dynamics suggested restricted recruitment by means of seed. Gene flow in the present population seemed to depend mainly on pollen. Local genetic structuring caused by seeds falling near the mother plant and by vegetative reproduction was detected by means of spatial autocorrelation. Three main modes of development of genetic structure were found for this species: (1) asexual reproduction by adventitious buds, which was dominant (up to 20 cm from the parent plants); (2) sexual reproduction by seeds that fell from flower stalks (up to 60 cm from the parent plants); and (3) pollen that dispersed over a wide area (throughout the population). Received: 13 January 2000 / Accepted: 21 March 2000     

Genetic Allee effects on performance, plasticity and developmental stability in a clonal plant

Negative effects of small population size on fitness, so-called Allee effects, may threaten population persistence even in intact habitat remnants. We studied genotypes of 14 isolated populations of the clonal plant Ranunculus reptans, for which molecular genetic (RAPD-) variability is higher for large than for small populations. In a competition-free greenhouse environment vegetative offspring of genotypes from large populations produced more rosettes and flowers, indicating higher fitness. Within-genotype coefficients of variation in performance traits, indicating developmental instability, were lower for genotypes from populations with higher RAPD-variability. In competition with a taller grass, we found relative reduction in leaf length less pronounced for plants from large populations, suggesting higher adaptive plasticity. Our experimental study of a plant with predominantly vegetative reproduction suggests, that negative genetic effects of recent habitat fragmentation, which so far rather were expected in plants with frequent sexual reproduction, are more severe and more common than previously acknowledged.     

Adaptive genetic differentiation in life-history traits between populations of <Emphasis Type="Italic">Mimulus guttatus</Emphasis> with annual and perennial life-cycles

The optimal allocation to sexual and vegetative reproduction as well as the optimal values of other life-history characteristics such as phenology, growth and mating system are likely to depend on the life-cycle of the organism. I tested whether plants of Mimulus guttatus originating from temporarily wet populations where the species has an enforced annual life-cycle have higher allocation to sexual reproduction, lower allocation to vegetative reproduction, more rapid phenology, faster growth, and floral traits associated with a self-fertilizing mating system than plants from permanently wet populations where the species has a perennial life-cycle. I grew a total of 1377 plants originating from three populations with an annual life-cycle and 11 populations with a perennial life-cycle in a greenhouse under permanently and temporarily wet conditions. Plants of M. guttatus in permanently wet conditions had significantly more vegetative reproduction and tended to have a faster growth than plants in the temporarily wet conditions, indicating plasticity in these life-history traits. Plants from populations with an annual life-cycle invested significantly more in sexual reproduction and significantly less in vegetative reproduction than the ones from populations with a perennial life-cycle. Moreover, this study showed that plants originating from populations with an annual life-cycle have a significantly faster development and floral traits associated with autonomous self-fertilization. In conclusion, this study suggests that there has been adaptive evolution of life history traits of M. guttatus in response to natural watering conditions that determine the life span of the species.     

为什么自然条件下沙地柏种群以无性更新为主

 基于调查、实验和文献资料,作者探讨了沙地柏(Sabina vulgaris)种群的繁殖特征和更新特征。1)根原基是沙地柏进行营养繁殖的结构基础,由根原基发育而成的不定根能够吸收充足的土壤资源供给萌生苗需要,因此定居的萌生苗能够独立维持。2)沙地柏同时具有营养繁殖和有性繁殖,因而沙地柏种群可通过无性更新和有性更新实现维持。3)种子质量极差、萌发率低和实生苗存活率低是制约沙地柏种群有性更新的3个“瓶颈”。4)沙地柏的营养繁殖力强、萌生苗存活率高、繁殖体的产生途径多样,这使自然条件下沙地柏种群以无性更新为主。     

Population genetics of the invasive water weed Elodea canadensis in Finnish waterways

Invasions of exotic species often involve a rapid evolutionary change in the introduced populations. Elodea canadensis is an invasive aquatic weed native to North America. Our aims were to reveal the evolutionary consequences of invasion to the population genetic structure of the presumably clonal E. canadensis in Finland and to test the hypothesis that the whole Finnish population originates from the first introduction of the species. We used ten polymorphic microsatellite markers to analyze the genetic characteristics of seven introduced E. canadensis populations in Finland. Despite the species' totally asexual mode of reproduction in Finland, two to five alleles per locus were detected in Finnish populations, and the expected heterozygosities varied between 0.19 and 0.37. The majority of variation was found within populations. Except for one, all pairwise values of population differentiation (F _ST) were significant, indicating restricted gene flow among the Finnish populations. In addition, a Bayesian analysis of population structure revealed the presence of regional population structuring. Genetic analyses indicate that E. canadensis could have been introduced to Finland multiple times. However, the amount of genetic variation and regional clustering detected could also be explained by post-establishment evolution, and based on this study it is not possible to exclude one introduction event followed by rapid evolution. We also tested the usability of the microsatellite markers for native North American samples in order to compare the within-population genetic characteristics of introduced and native populations. However, in native populations only four microsatellite markers amplified reliably, indicating sequence variation within primer-binding regions and, thus, genetic differentiation among populations of E. canadensis.