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.     

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.     

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.     

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.     

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.     

Post-burn reproduction of woody plants in a neotropical savanna: the relative importance of sexual and vegetative reproduction

1. Burning typically occurs at intervals of 1–3-years in the Brazilian cerrado, a rate that exceeds the precolonization fire regime. To determine if woody plants of the cerrado successfully reproduce within the short span of time between burns, experimental burns were used to quantify the effects of fire on sexual and vegetative reproduction of six species of resprouting trees and shrubs.
2. Four of the six species reproduce vegetatively by producing root suckers. For three of these species, Rourea induta , Myrsine guianensis and Roupala montana , sucker production was seven to 15 times greater in burned plots than in unburned controls.
3. Fire had a negative impact on sexual reproduction. Fire caused an immediate reduction in sexual reproductive success by destroying developing reproductive structures and seeds. Additionally, five of the six study species exhibited overall reductions in seed production in the years following fire. Fire had this effect by reducing the individual size of all species and, for three species, by reducing size-specific reproductive output. Only the tree Piptocarpha rotundifolia exhibited increased seed production following burning.
4. Fire caused substantial mortality to both seedlings and suckers. Suckers were larger than seedlings and experienced lower mortality rates for two of three species. Fire-induced mortality of seedlings varied greatly among species, ranging from 33% to 100%.
5. The results indicate that vegetative reproduction is much more successful than sexual reproduction under the high fire frequency typical of current fire regimes. It is concluded that current fire regimes must be causing a shift in species composition, favouring species capable of vegetative reproduction.     

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.     

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.     

Reproductive strategies may explain plant tolerance to inundation: A mesocosm experiment using six marsh species

We investigated the impact of inundation duration on sexual and asexual reproduction strategies in six hygrophilic angiosperm species (2 annuals - Ranunculus sardous and Ranunculus ophioglossifolius, 2 rhizomatous - Juncus articulatus and Eleocharis palustris, and 2 stoloniferous species - Glyceria fluitans and Agrostis stolonifera). Plant growth during three inundation durations (natural, +3 weeks, +6 weeks) was assessed in 20 mesocosms. Biomass and reproductive traits were measured at the onset of flowering and at seed set.The six species maintained or increased their total mass in response to increased inundation period, suggesting a good tolerance to inundation stress. No changes were recorded in flowering time for the five species that flowered. The two annual species increased the number of flowers and mass allocated to sexual reproduction with increased duration of inundation, promoting extensive seed production. The response was species-specific in perennial species with an either a positive or a negative Gaussian curve of both traits in response to the increased duration of inundation. The duration of inundation affected ramet production in all perennial species except A. stolonifera, promoting ramet production in two out of the three species. In all species except E. palustris, the length and number of stolons or rhizomes were maintained, suggesting the importance of these connections in the resistance of perennial plants to flooding stress. These species occurring in repeatedly inundated habitats thus appeared to be tolerant to long inundations thanks to both sexual reproduction and vegetative propagation.     

Seed characteristics and germination limitations in the highly invasive <Emphasis Type="Italic">Fallopia japonica</Emphasis> s.l. (Polygonaceae)

The species in the Japanese knotweed complex (Fallopia japonica s.l. and its hybrids) are among the most invasive plants on earth. Their expansion and reproduction in the introduced range have been mostly due to vegetative reproduction, but observations of low seedling numbers and hybridization processes exist. Knowledge of factors affecting germination characteristics is essential if the risk of sexual reproduction is to be assessed, and its impact on the ability of the species to spread and adapt to different environments. This study aims to examine the germination success of Fallopia japonica s.l. seeds of different ages, quality, and storage conditions. Irrespective of age and even after natural overwintering in the soil, seeds germinated quite well (48–79%). Ungerminated seeds collected in autumn of 2008 were characterized by a low weight/length ratio (W/L), low nutrient concentrations, and a greenish tepal coloration. These differences may be due to alternative male taxa participating in the pollination process. Spring collected seeds were subject to strong predation by birds. In contrast to the high germination observed under laboratory conditions, seed germination or early establishment in the field was inhibited and only a few seedlings were observed. Although the factors that inhibit the establishment of mature seeds in the field remain unknown, there is a clear risk that sexual reproduction could gain importance in the future as a result of changing environmental conditions or genetic adaptation. Not only would this facilitate expansive dispersal by wind, but it might also increase the potential for further adaptation of the species complex.     

Effects of pollen and nitrogen availability on reproduction in a woodland herb,Lysimachia quadrifolia

Summary Relationships between pollen loads, resource availability, and fruit and seed production were determined for Lysimachia quadrifolia ramets in two adjacent sites (the scrub site and the open site) in 1982, 1983, and 1984. Pollen loads limited % fruit set and seed production in the open site in 1982 and 1983. Reproduction in the scrub site was resource limited in 1982, as shown by an increase in % fruit set when one-half of the flowers on a ramet were removed prior to fruit initiation. In the scrub site in 1983, pollination of one-half of the flowers on a ramet decreased the % fruit set of the remaining, unpollinated flowers. Fruit production in the same site was limited by pollen in 1984. Addition of nitrogen fertilizer to the scrub site in 1984 had no effect on fruit and seed production. There was more variation in fruit set between sites than between years. There was no trend to greater fruit set or number of seeds/fruit on early flowers compared to late flowers on the same ramet. Correlations between measures of reproduction were positive or insignificant. These results demonstrate year-and site-specific variation in the factors that limit plant reproduction.     

植物种群的繁殖对策

植物种群的繁殖对策钟章成（西南师范大学,重庆６３０７１５）ＲｅｐｒｏｄｕｃｔｉｖｅＳｔｒａｔｅｇｉｅｓｏｆＰｌａｎｔＰｏｐｕｌａｔｉｏｎｓ．￥ＺｈｏｎｇＺｈａｎｇｃｈｅｎｇ（ＳｏｕｔｈｗｅｓｔＣｈｉｎａＴｅａｃｈｅｒｓＵ－ｎｉｖｅｒｓｉｔｙ,Ｃｈｏｎ...     

Studies on Sexual Reproduction and Vegetative Propagation of Eleutherococcus senticosus (Araliaceae)

Eleutherococcus senticosus has two reproduction modes: vegetative propagation and sexual reproduction. In this paper, flower number per inflorescence, seed set and plump seed ratio,some aspects of rhizome morphology and morphogenesis of three morphs with different filament lengths are reported, and the breeding system is studied by hand-pollination. It is clear from the two tables presented in the present paper that plants with long filaments cannot fruit, plants with short filaments can fruit annually, but their seed setting ratio is different in different years and different habitats, plants with medium-long filaments can also fruit but their plump seed ratio may be low. In ordinary years, the seed setting ratio of plants with short filament reaches 40% and plants at the edge of forests have slightly higher seed setting ratio than those on cleared area and those in natural secondary forests. Considering the existence of only a few populations with medium-long filaments in the area studied,we conclude that most of seeds in Eleutherococcus senticosus should be the products of cross pollination;and with the coexistence of vegetative propagation and sexual reproduction, the species of Eleutherococcus senticosus should not be endangered,if not destructed by human beings.     

刺五加的有性生殖与营养繁殖

刺五加 Eleutherococcus senticosus (Rupr.et Maxim.)Maxim,既可进行有性繁殖,又可进行营养繁殖。本文报道其三类花丝长度不同的植株每个花序的花朵数量、座果率、饱满种子率及根茎形态和形态发生的某些特点。通过人工受精,检测了其繁育系统。结果表明:长花丝植株不座果,短花丝植株座果,但不同年份、不同生境中座果率不同,中花丝植株也座果但饱满种子率教低。在正常年份,短花丝植株的座果率接近40％,并且林缘的座果率稍高于皆伐迹地和次生林下的。考虑到中花丝植株在所研究地区居群数很少,我们认为绝大多数刺五加种子是异花传粉的产物,其有性生殖与营养繁殖相配合,在无人为破坏的前提下,该物种不应该濒危。     

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.     

Cottonwood hybrids gain fitness traits of both parents: a mechanism for theirlong-term persistence?

Using surveys of natural populations, experimental crosses, and common garden trials, we tested the hypothesis that hybrid cottonwoods (Populus fremontii × P. angustifolia) from the Weber River in northern Utah would produce as many viable offspring as their parental species. We found that both F(1) generations and backcross generations can be just as fit as the parent taxa. First, F(1) hybrids produced as many viable seed as P. angustifolia (but less than P. fremontii), and backcross genotypes produced as many viable seeds as both parent taxa. Second, hybrids produced nearly two times as many ramets from root sprouts as P. angustifolia and four times as many ramets as P. fremontii. Third, the high mortality of germinated seedlings of all tree types (i.e., >90%) and very low mortality of asexually derived ramets provide hybrids with equal sexual reproduction and enhanced asexual reproduction, especially since backcross hybrids exhibit transgressive segregation in ramet production. Our findings suggest that the introgression of P. fremontii seed traits into the hybrid genome is responsible for their equivalent performance (at least to one parent) in sexual reproduction, while the contributions of asexual traits from P. angustifola results in hybrids having equal or greater fitness.     

Conditions for seedling establishment and probable function of seed in the clonal sedge Carex rugulosa Kük. in riverside marshes

Carex rugulosa Kük. forms large meadows in moist zones along estuarine riversides. These meadows are usually maintained by rhizomatous ramet production (clonal growth), and the appearance of seedlings is uncommon. We studied the conditions required for seedling establishment. In areas in which clonal ramets were dense (906–1050/m²), and the relative light intensity at ground level was low (0.8–3.8%), seed germination was entirely suppressed. In contrast, many seedlings (288–513/m²) were observed in areas in which clonal ramets had been considerably reduced (13–269/m²) by continuous inundation in the previous year. In these areas, the relative light intensity had increased (20.5–79.3%). It was inferred that seeds resisted the prolonged inundation that killed the ramets, and that germination was induced under these open conditions. These results suggest that the main functions of sexual reproduction in C. rugulosa are recovery after severe damage to vegetation using seeds from the seed bank and the dispersal of seeds to areas without dense vegetation. Most seedlings died when the marshy meadow was temporarily inundated by heavy rain. However, transplanted seedlings survived better at artificially elevated sites that were not submerged. This shows that inundation during the seedling stage impedes seedling survival. Therefore, seeds can contribute to recovery after ramet decline or colonization to open area when (i) the rainfall is not heavy to cause seedling submergence or (ii) seeds are dispersed to higher area which is not inundated.     

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。     

Somatic and demographic costs under different temperature regimes in the late-flowering alpine perennial herb Saxifraga stellaris (Saxifragaceae)

If reproduction involves costs, preventing reproduction one year should result in increased growth and/or reproduction the next year. Costs of reproduction in the late-flowering perennial alpine herb Saxifraga stellaris were studied in an experimental field study. To determine whether cost of reproduction differed between two contrasting temperature regimes, I examined plants in Open Top Chambers (OTCs) and control plots. One year before measurements all flower buds on the experiment plants were removed. There was no impact of the flower bud removal or the interaction between flower removal and temperature on prefloration time or seed maturation time. However, cost of reproduction influenced growth (somatic cost), vegetative propagation, flowering frequency, number of stem per ramet, number of fruits, and seed mass (demographic cost). However, significant interaction effects of flower removal and temperature on growth and fruit production revealed that the cost of reproduction differed between the two temperature regimes. The warmed plants showed reduced cost of reproduction compared to plants growing under natural temperature.     

克隆乔木黄牛奶树两种繁殖方式的适合度分析

对黄牛奶树两种生殖方式(克隆生殖和有性生殖)的适合度研究结果表明,不同生境下,黄牛奶树繁殖及占据空间的方式有一定差异.在水肥资源丰富、郁闭度较大(常绿阔叶林和竹林)条件下,克隆生殖和有性生殖幼苗存活率均较高,占据空间的能力较强,但克隆苗在数量和占据空间的能力上占较大优势,主要以克隆方式进行繁殖;水肥资源贫乏、光照较强的条件下,两类苗存活率和占据空间能力均较低,实生苗在数量和占据空间能力上占优势.黄牛奶树两种繁殖方式的瓶颈期不同,有性生殖的瓶颈期在种子到幼苗阶段,而克隆苗在幼苗到成株阶段.黄牛奶树的入侵过程是先以实生苗侵入一个地点定居,在郁闭度较大、水肥良好的条件下,以克隆方式迅速占领空间.生长初期克隆苗表现出极大的优势,后期(15龄以上)则优势不复存在.