Evolutionarily stable allocation to vegetative and sexual reproduction in plants

I apply the resource allocation approach to construct an ESS model aimed at predicting the rate of vegetative and sexual reproduction in plants. The model provides an integrated explanation of why the mode of reproduction should vary between populations and species, and what factors should affect the trade-off between ramet and seed production. It follows that seed production, usually considered to be less effective and more costly than ramet production, may be maintained in the population. The production of ramets should be favored when their recruitment rate is high or the costs are low. The intensity of vegetative reproduction is also related to conditions which affect seed production directly. The rate of ramet production should be inversely correlated with factors favoring sexual reproduction, such as lower cost of seeds or high opportunity for colonization of new patches, etc. Under such conditions it is beneficial to allocate more resources available for reproduction into seeds, at the expense of ramet production. The model emphasizes the need to consider both modes of plant reproduction whenever one of them is theoretically or empirically studied.     

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.     

Optimal resource allocation to seeds and vegetative propagules under density-dependent regulation in Syneilesis palmata (Compositae)

Syneilesis palmata reproduces by both seeds and vegetative propagules (short rhizomes). The latter result in the production of new plants that are larger in size and hence have a higher survival probability and a higher growth rate than seeds. A previous study predicted that the optimal reproductive strategy, in terms of maximizing population growth rate (a fitness measure under no density regulations), was pure vegetative reproduction. However, high resource investment to vegetative propagules can cause local crowding resulting in reduced demographic performances of the plants, because the vegetative propagules of Syneilesis are produced close to one another. We examined, in this situation, the impact of allocating a certain proportion of reproductive resource to seeds with relatively greater capacity for dispersal. We simulated dynamics of hypothetical Syneilesis populations with various reproductive resource allocation balances (from pure seed to pure vegetative reproduction), using a density-dependent matrix model. In the model, it was assumed that plants from vegetative propagules experienced density-dependent reduction in their survival probabilities, but this was not the case for plants originating from seeds. Each allocation strategy was evaluated based on an equilibrium population density, a fitness measure under density-dependent regulations. The optimal reproductive strategy predicted was pure vegetative reproduction. Unrealistic conditions were required for seed reproduction to be favoured, such as the production of seeds one hundred times the normal number per unit resource investment. However, the conditions were fairly relaxed compared with those required in the model where no density effects were incorporated. This indicates that escape from local crowding is likely to be one of the roles of seed production in Syneilesis.     

Sexual and vegetative reproduction in the aboveground part of a dioecious clonal plant, <Emphasis Type="Italic">Dioscorea japonica</Emphasis> (Dioscoreaceae)

In dioecious clonal plants, the reproductive effort required to set seeds will be responsible for the larger investment in sexual reproduction by females. If there will be a trade-off in resource allocation between sexual and clonal reproduction, this differential sexual reproduction will lead to sexual differentiation in the relative amount of clonal reproduction. To test this prediction, we studied differences between the sexes in their phenologies and investments in sexual and vegetative reproduction (clonal reproduction by means of bulbils) with respect to ramet size in a dioecious clonal plant, Dioscorea japonica Thunb. The period of bulbil production overlapped the period during which infructescences developed. Females flowered later, produced heavier inflorescences, and fewer flowers per inflorescence than did males. Regression analysis using the size of the individual plants demonstrated that large females made smaller investments in inflorescences and larger investments in sexual reproduction than did large males. In contrast, females invested fewer resources in vegetative reproduction than did males. However, the total investments in sexual and vegetative reproduction did not differ between the sexes. These results supported our hypothesis on the sexual differentiation in sexual and clonal reproduction.     

Changes in reproductive investment with altitude in an alpine plant

Aims In perennial species, the allocation of resources to reproduction results in a reduction of allocation to vegetative growth and, therefore, impacts future reproductive success. As a consequence, variation in this trade-off is among the most important driving forces in the life-history evolution of perennial plants and can lead to locally adapted genotypes. In addition to genetic variation, phenotypic plasticity might also contribute to local adaptation of plants to local conditions by mediating changes in reproductive allocation. Knowledge on the importance of genetic and environmental effects on the trade-off between reproduction and vegetative growth is therefore essential to understand how plants may respond to environmental changes.Methods We conducted a transplant experiment along an altitudinal gradient from 425 to 1?921 m in the front range of the Western Alps of Switzerland to assess the influence of both altitudinal origin of populations and altitude of growing site on growth, reproductive investment and local adaptation in Poa alpina .Important findings In our study, the investment in reproduction increased with plant size. Plant growth and the relative importance of reproductive investment decreased in populations originating from higher altitudes compared to populations originating from lower altitudes. The changes in reproductive investment were mainly explained by differences in plant size. In contrast to genetic effects, phenotypic plasticity of all traits measured was low and not related to altitude. As a result, the population from the lowest altitude of origin performed best at all sites. Our results indicate that in P. alpina genetic differences in growth and reproductive investment are related to local conditions affecting growth, i.e. interspecific competition and soil moisture content.     

No long-term costs of meristem allocation to flowering in stoloniferous Trifolium species

Clonal plants propagate by means of clonal growth and sexual reproduction. The commitment of meristems to branching and flowering govern the expression of these two mutually exclusive life-history functions. We used a modelling and an experimental approach to examine the consequences of a structural trade-off between flowering and clonal growth on future growth and fitness in stoloniferous species with a determinate module architecture. The model revealed negative effects of flowering on vegetative growth due to a structural trade-off at the meristem level. Total fecundity was maximized at intermediate flowering frequencies. In addition, optimal meristem commitment to flowering depended strongly on the time available for growth and reproduction. This indicates an interaction between optimal flowering frequency, the length of the growing period and the rate of ontogenetic development. The greenhouse study made use of 15 genotypes of two closely related, stoloniferous Trifolium species. Despite the existence of a structural trade-off at the meristem level, we found no evidence for costs of flowering on the whole-plant level. High allocation to flowering did not result in reduced plant performance (biomass and module production) and total fecundity, indicating that there were no demographic costs of meristem investment to different life-history functions. Flowering frequencies never exceeded the model prediction for optimal commitment of meristems to sexual reproduction, suggesting strong past selection to eliminate high levels of meristem allocation to flowering. Hence, clonal growth seems to have evolutionary priority over sexual reproduction in our species. This revised version was published online in November 2006 with corrections to the Cover Date.     

海拔对青藏高原东缘弯齿风毛菊繁殖特征的影响

研究了海拔对青藏高原东缘不同海拔12个居群弯齿风毛菊繁殖特征的影响.结果表明: 弯齿风毛菊的个体大小、繁殖器官和营养器官生物量、每株头状花序数量及种子总数量均随海拔的升高而减小,而每个头状花序质量和种子百粒重随海拔的升高而增加,表明弯齿风毛菊可以通过减小个体大小来降低资源消耗,把获取的有限资源分配到繁殖部分以达到繁殖成功.为了保证顺利完成有性繁殖,繁殖分配随海拔升高而增大.弯齿风毛菊头状花序的数量与大小、种子数量与百粒重之间均存在资源分配上的权衡,以此来适应环境胁迫,提高自身的适合度.     

新疆郁金香营养生长、个体大小和开花次序对繁殖分配的影响

植物有性繁殖与资源分配的关系研究对于揭示植物生活史特征及繁育系统进化具有重要意义。新疆郁金香(Tulipa sinkiangensis)是新疆天山北坡荒漠带特有的一种多年生早春短命植物。在自然生境中,该物种仅以有性繁殖产生后代,每株能产生1-8朵花,且不同植株上的花数及果实数以及花序不同位置上的花与果实大小明显不同。本文通过对新疆郁金香有性繁殖与营养生长及植株大小的关系以及花序中不同位置花及果实间的资源分配研究,旨在揭示营养生长、个体大小及开花次序对其繁殖分配的影响。结果表明:在开花和果实成熟阶段,新疆郁金香植株分配给营养器官(鳞茎和地上营养器官)与繁殖器官的资源间均存在极显著的负相关关系(P<0.01),说明其植株的营养生长与生殖生长间存在权衡关系。多花是新疆郁金香的一个稳定性状,其植株上花数目、花生物量、果实生物量和种子数量与植株生物量间均呈极显著的正相关关系(P<0.01),说明新疆郁金香植株的繁殖分配存在大小依赖性。在具2-5朵花的新疆郁金香植株中,花序内各花的生物量、花粉数和胚珠数、结实率、果实生物量、结籽数、结籽率及种子百粒重按其开花顺序依次递减,说明花序内各花和果实的资源分配符合资源竞争假说。植株通过减少晚发育的花或果实获得的资源来保障早发育的花或果实获得较多的资源,从而达到繁殖成功。     

The role of sexual and vegetative reproduction in the population maintenance of a monocarpic perennial herb,Cardiocrinum cordatum var. glehnii

Because monocarpic perennial plants have only one reproductive opportunity in their entire life, they need to ensure offspring production. Some plants reproduce both sexually and vegetatively, and vegetative reproduction could possibly compensate for seed production. Therefore, the role and significance of these reproductive modes is likely to differ between monocarps and polycarps, which can reproduce many times. Cardiocrinum cordatum var. glehnii is a monocarpic perennial that reproduces both sexually and vegetatively (bulblet formation). Here, we investigated the characteristics and contribution to population maintenance of sexual and vegetative reproduction to reveal the significance of these two reproductive modes in this species. First, we found that bulblet formation occurred in plants after the three‐leaved rosette stage. Second, resource allocation experiments revealed that although resources were mainly invested in fruit maturation after the flowering season, resource allocation was switched from sexual reproduction to vegetative reproduction if seed production was insufficient. Third, the outcrossing rate in this species varied greatly according to the environment surrounding the population. However, reproductive assurance by selfing kept seed production stable even if flowers did not receive sufficient pollen for full seed set via outcross pollination, and moreover, there was no intensive inbreeding depression. Finally, genotypic identification of ramets suggested that daughter ramets derived from vegetative reproduction received the space that the mother flowering ramet had occupied until the previous year.     

The role of vegetative spread and seed dispersal for optimal life histories of clonal plants: a simulation study

Sexual and vegetative reproduction of clonal plants phenotypically differ in dispersal distance, in the phenology of offspring production and establishment, and in the success of establishment. We applied a combination of analytical and of spatially explicit individual-based simulation modelling to calculate long-term fitness. We predicted optimal clonal plant life histories in a parameter space spanned by stolon number, stolon internode length, and relative allocation to sexual and vegetative reproduction. For a given allocation to sexual reproduction and number of stolons, fitness was optimised for rather short internode lengths under small disturbances, and for the longest possible internodes under larger disturbances. A trade-off between length and number of vegetative spacers drew parameters away from their unrestricted optima. Now, intermediate length and number of spacers led to maximum fitness under large disturbances. Simultaneous trade-offs between sexual and vegetative reproduction and between the length and number of spacers could also lead to fitness optima at intermediate parameter values, depending on the success of seedling establishment. We demonstrated that spatial habitat structure (1) selects for an efficient use of available space either by optimum internode length or by investment into seeds, which disperse farther than vegetative spacers, and (2) leads to an interaction between trade-offs. We conclude, that dispersal distance, i.e. a spatial life-history component, and trade-offs must be included in considerations on adaptive evolution of clonal life histories.     

对虾杆状病毒病暴发式大流行的生态机理初步研究

初步探讨中国和沿海和对虾杆状病毒病暴发式大流行的生理机理。分析了病毒病特性与对虾免疫功能的关系;病毒在虾池间的传染机制。着重说明了虾病的群体感染过程,给出模式化的群体传染模型,简要阐述了与虾病相关的社会生态经济问题。     

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。     

Trade-offs between sexual and asexual reproduction in a monoecious species Sagittaria pygmaea (Alismataceae): the effect of different nutrient levels

Available resources could influence the trade-offs among different reproductive components in plants. Here, we created three nutrient levels to test the nutrient effects on trade-offs among sexual reproduction, clonal propagation and vegetative growth in a monoecious clonal herb Sagittaria pygmaea. The results of this study showed that the plant exhibited different trade-off patterns among different nutrient levels. When the nutrient level was low, there were weak trade-offs between sexual reproduction and vegetative growth and between clonal propagation and vegetative growth; when the nutrient level was moderate, we found a strong trade-off between sexual reproduction and clonal propagation; but when the nutrient level was high, we found no trade-offs among these three different reproductive components. These results indicated that the plant could adjust its trade-off patterns to fit the nutrient variation and suggested that trade-offs are unlikely to constrain the evolution of reproductive strategy in this species.     

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.     

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.     

What drives selection on flowering time? An experimental manipulation of the inherent correlation between genotype and environment

The optimal timing of the seasonal switch from somatic growth to reproduction can depend on an individual's condition at reproduction, the quality of the environment in which it will reproduce, or both. In annual plants, vegetative size (a function of age at flowering) affects resources available for seed production, whereas exposure to mutualists, antagonists, and abiotic stresses in the environment (functions of Julian date of flowering) influences success in converting resources into offspring. The inherent tight correlation between age, size, and environment obscures their independent fitness contributions. We isolated the fitness effects of these factors by experimentally manipulating the correlation between age at flowering and date of flowering in Brassica rapa. We staggered the planting dates of families with differing ages at flowering to produce experimental populations in which age at flowering and date of flowering were positively, negatively, or uncorrelated. In all populations, plants with an early date of flowering produced more seed than those flowering late, regardless of age or size at flowering onset. The temporal environment was thus the principal driver of selection on flowering time, but its importance relative to that of age and size varied with the presence/absence of herbivores and seed predators.     

Optimal allocation to vegetative and sexual reproduction in plants: the effect of ramet density

Vegetative reproduction is a very common alternative by which plants can contribute to the next generations. There are many considerations predicting which mode of reproduction, vegetative or sexual, should be favored and numerous experimental studies to verify them. However, the results are inconsistent especially when the effect of plant density is considered. I apply here a dynamic optimization model to predict the rate of vegetative and sexual reproduction in plants as a response to changes in the local plant density. The population is assumed to occupy a heterogeneous environment consisting of patches in which growth and reproduction of plants are possible and unfavorable space between them. As the environment is globally stable, the seeds, which can disperse without restriction, exhibit a constant recruitment rate. The ramets are assumed to settle only within the patch of the mother plant. The rate of ramet production effects local density, which in turn determines ramet recruitment. The optimal strategy maximizes the expected lifetime genetic contribution, realized via both vegetative and sexual reproduction. The solutions obtained under these assumptions are dualistic. The model predicts that different approaches applied in studying the effect of ramet density should give opposite outcomes. When the comparison is between patches in natural populations, a positive relationship between relative ramet allocation and density is expected. When the density is experimentally manipulated or its effect is analyzed across different successional stages, a negative relationship should be found. The results seem to be confirmed by empirical studies.     

不同生境下珠芽蓼（蓼科）的繁殖策略比较

在青藏高原横断山区以海拔相同的高山流石滩和高寒草甸中的珠芽蓼（Polygonum viviparum）居群为研究对象,探讨该物种在不同生境条件下的繁殖策略以及无性繁殖和有性繁殖间的关系。结果发现,流石滩居群的珠芽蓼植株高度和珠芽数量显著低于高寒草甸居群,而珠芽和花总数量、花数量和每个花序上的花比例却显著高于高寒草甸居群。研究结果表明珠芽蓼在环境更为恶劣的高山流石滩生境中增加了对繁殖器官和花资源的投资,说明在高山植物中繁殖优先于营养生长,且有性繁殖比无性繁殖具有更为重要的作用。而对每个植株上花数量和珠芽数量的统计结果表明,两者呈显著的负相关关系,进一步证明了同一植株内无性繁殖和有性繁殖的权衡关系。     

Trade-offs between sexual and clonal reproduction in an aquatic plant: experimental manipulations vs. phenotypic correlations

That trade-offs result from the allocation of limited resources is a central concept of life history evolution. We quantified trade-offs between sexual and clonal reproduction in the aquatic plant, Butomus umbellatus, by experimentally manipulating sexual investment in two distinct nutrient environments. Increasing seed production caused a significant but nonlinear trade-off. Pollinating half of all flowers strongly reduced clonal bulbil production, but pollinating the remaining flowers did not cause any further trade-off. Trade-offs were not stronger under low nutrient conditions that clearly limited plant growth. Experimentally induced trade-offs were not reflected in negative phenotypic correlations between sexual and clonal allocation among plants within eight populations grown in a uniform greenhouse environment. Diminishing effects of increased sexual allocation plus a lack of accord between experimental manipulations and phenotypic correlations suggest that trade-offs between sexual and clonal reproduction are unlikely to constrain the evolution of reproductive strategy in this species.     

Costs of reproduction as related to the timing of phenological phases in the dioecious Shrub Pistacia lentiscus L

Females of woody dioecious species usually devote more resources to reproduction than males. This may lead to a decrease in female survival and growth. The costs of reproduction, however, can be lightened through a number of mechanisms, as for example avoiding the temporal coincidence of reproduction and vegetative growth. The aim of this study was to evaluate whether males and females of P. lentiscus differ in the timing of their vegetative growth, and to assess whether the sequencing of vegetative growth and reproduction reduces reproductive costs. We monitored phenology in males and females. We also compared male and female allocation of nutrients and biomass in the branch, and the developmental stability of the growing shoots. We did this both prior to and at the end of the fruiting period. Males and females showed similar vegetative and flowering phenologies. Males invested more biomass in flowering, but the sexes showed equal vegetative biomass and nutrient content prior to the fruiting period. In female branches, no trade-off was found between fruit load and current-year vegetative growth. In P. lentiscus, avoiding the overlap of flowering, vegetative growth and fruiting probably contributes to reduce the immediate costs of reproductive efforts, both in males and females.