Effects of light on the growth and clonal reproduction of Ligularia virgaurea

Ligularia virgaurea is a perennial herb that is widely distributed in the alpine meadow on the eastern Qinghai-Tibet plateau.We investigated the patterns of growth and reproduction of L.virgaurea under two contrasting levels of light conditions for two continuous growing seasons.Our results showed that the light affects on the maximum relative growth rate,the shoot weight ratio and the root weight ratio differed between the two growing seasons.L.virgaurea reproduced initially through rhizome in the second growing season,rather than sexual reproduction.The proportion of genets with clonal reproduction decreased under shaded conditions.A minimum genet size should be attained for clonal reproduction to begin under the shaded conditions.There was a positive linear relationship between clonal reproduction and genet size.Light level affected the allocation of total biomass to clonal structures,with less allocation under the full natural irradiance than under the shaded conditions.There seemed to be a trade-off between vegetative growth and clonal reproduction under the full natural irradiance,in terms of smaller relative growth rates of genets with clonal reproduction than those without clonal reproduction.L.virgaurea emphasized clonal reproduction under the full natural irradiance,while the plant emphasized vegetative growth under the shaded conditions.     

Sexual reproduction and clonal growth in Reinhardtia gracilis (Palmae), an understory tropical palm

Patterns of sexual reproduction and clonal growth were investigated in the understory palm Reinhardtia gracilis var. gracilior over a 3-yr period. R. gracilis is a very abundant clonal palm in the tropical rain forest of Los Tuxtlas, Veracruz, México. Because ramets form clumps, genets are easily identified in the field. Genets were monitored in a 0.5-ha area, and classified by size according to the number of ramets they possessed. In contrast to clonal growth, sexual reproduction was highly dependent on genet size. The probability of reproduction, the number of inflorescences, and the number of fruits produced were positively correlated with genet size. However, neither the probability of producing a ramet, nor the number of ramets produced per genet were correlated with genet size. Over the 3 yr of study, 55% of the genet population had at least one ramet with reproductive structures, while <1% (a single genet in one year) had six ramets with flowers. Thirty-two percent of the mature genets reproduced during each of three consecutive years. In contrast, 58% of the genets produced no new ramets during these 3 yr. No evidence was found of a trade-off between clonal growth and sexual reproduction. Ramet production increases genet size and this in turn increases genet reproductive performance. Clonal growth in this species may be viewed as a growth strategy that tends to maximize genet fitness.     

Density-dependent reproductive and vegetative allocation in the aquatic plant Pistia stratiotes (Araceae)

Pistia stratiotes is an aquatic macrophyte that grows in temporary-ponds in the southern Pantanal, Brazil. It reproduces both sexually and asexually and is usually observed forming dense mats on the water surface, a condition favored by the plant's vegetative reproduction coupled with an ability for rapid growth. In this study we examined the effect of densely crowded conditions on the production of reproductive and vegetative structures. In addition, we verified whether there is a trade-off between clonal growth and investment in sexual reproductive structures, and whether there is an allocation pattern with plant size. Individual plant biomass and the number of the rosettes producing sexual reproductive structures and vegetative growth structures both increased with density. Increase in plant size resulted in increased proportional allocation to sexual reproductive structures and vegetative growth structures. Allocation of biomass to reproduction did not occur at the expense of clonal growth. Thus, the density response appears as a increase of rosettes producing sexual reproductive structures and vegetative growth structures. Therefore, long leaves and stolons may be adaptive under densely crowded conditions where competition for light is intense. An important aspect in the study of trade-offs is the size-dependency of the allocation patterns .Usually, larger plants produce more biomass. Therefore, larger plants can allocate more biomass to both vegetative and sexual reproduction than smaller plants and thus show a positive correlation between both traits rather than the expected negative one.     

Reproductive demography of ramets and genets in a rhizomatous clonal plant <Emphasis Type="Italic">Convallaria keiskei</Emphasis>

Clonal growth occurring below the ground makes it difficult to identify individuals and demonstrate the demographic features of a focal plant species. In this study, genotypically identified ramets of a rhizomatous clonal herb, Convallaria keiskei Miq., were monitored for their growth, survival, and reproduction from 2003 to 2006. After the monitoring period, their subterranean organs were excavated to explore the underground connections of established ramets and the direction of clonal growth. We then combined data on the fate of the monitored ramets with the information of rhizome connections, clarifying reproductive demography at both the ramet and genet levels. Although each ramet initiated both sexual reproduction (via flowering) and clonal growth, clonal growth tended to precede sexual reproduction. In a surveyed genet, 51.0% of ramets produced flowers and 29.6% generated clonal offspring during the study period. Consequently, we clarified the reproductive demography of C. keiskei: clonal growth tended to precede flowering in a ramet, and a genet can keep reproducing every season at the genet level, despite a ramet not having inflorescence every year.     

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.     

根茎型植物扁秆荆三棱对光照强度和养分水平的生长响应及资源分配策略

自然界中光照和养分因子常存在时空变化,对植物造成选择压力。克隆植物可通过克隆生长和生物量分配的可塑性来适应环境变化。尽管一些研究关注了克隆植物对光照和养分因子的生长响应,但尚未深入全面了解克隆植物对光照和养分资源投资的分配策略。以根茎型草本克隆植物扁秆荆三棱(Bolboschoenus planiculmis)为研究对象,在温室实验中,将其独立分株种植于由2种光照强度(光照和遮阴)和4种养分水平(对照、低养分、中养分和高养分)交叉组成的8种处理组合中,研究了光照和养分对其生长繁殖及资源贮存策略的影响。结果表明,扁秆荆三棱的生长、无性繁殖及资源贮存性状均受到光照强度的显著影响,在遮阴条件下各生长繁殖性状指标被抑制。且构件的数目、长度等特征对养分差异的可塑性响应先于其生物量积累特征。在光照条件下,高养分处理的总生物量、叶片数、总根茎分株数、长根茎分株数、总根茎长、芽长度、芽数量等指标大于其他养分处理,而在遮阴条件下,其在不同养分处理间无显著差异,表明光照条件可影响养分对扁秆荆三棱可塑性的作用,且高营养水平不能补偿由于光照不足而导致的生长能力下降。光照强度显著影响了总根茎、总球茎及大、中、小球茎的生物量分配,遮阴条件下,总生物量减少了对地下部分根茎和球茎的分配,并将有限的生物量优先分配给小球茎。总根茎的生物量分配未对养分发生可塑性反应,而随着养分增加,总球茎分配下降,说明在养分受限的环境中球茎的贮存功能可缓冲资源缺乏对植物生长的影响。在相同条件下,根茎生物量对长根茎的分配显著大于短根茎,以保持较高的繁殖能力;而总球茎对有分株球茎的生物量分配小于无分株球茎,表明扁秆荆三棱总球茎对贮存功能的分配优先于繁殖功能。研究为进一步理解根茎型克隆植物对光强及基质养分环境变化的生态适应提供了依据。     

Fine‐scale genetic structure and flowering output of the seagrass Enhalus acoroides undergoing disturbance

Seagrass are under great stress in the tropical coast of Asia, where Enhalus acoroides is frequently the dominant species with a large food web. Here, we investigate the question of the fine‐scale genetic structure of this ecologically important foundation species, subject to severe anthropogenic disturbance in China. The genetic structure will illuminate potential mechanisms for population dynamics and sustainability, which are critical for preservation of biodiversity and for decision‐making in management and restoration. We evaluated the fine‐scale spatial genetic structure (SGS) and flowering output of E. acoroides, and indirectly estimated the relative importance of sexual versus asexual reproduction for population persistence using spatial autocorrelation analysis. Results reveal high clonal diversity for this species, as predicted from its high sexual reproduction output. The stronger Sp statistic at the ramet‐level compared with genet‐level indicates that clonality increases the SGS pattern for E. acoroides. Significant SGS at the genet‐level may be explained by the aggregated dispersal of seed/pollen cohorts. The estimated gene dispersal variance suggests that dispersal mediated by sexual reproduction is more important than clonal growth in this study area. The ongoing anthropogenic disturbance will negatively affect the mating pattern and the SGS patterns in the future due to massive death of shoots, and less frequency of sexual reproduction.     

Density and the commitment of apical meristems to clonal growth and reproduction in Hieracium pilosella

Summary We examined responses to population density in the commitment of apical meristems to reproduction and clonal growth in a rosette-forming, stoloniferous herb (Hieracium pilosella). Despite close physiological coupling between the evocation of the terminal inflorescence bud and the development of one or more axillary buds into stolons, the allocation of meristems was extremely plastic.Genets at the higher sowing densities showed density-dependent mortality consistent with self-thinning along a-3/2 trajectory. The probability of inflorescence evocation and associated stolon development was negatively dependent on surviving density. The proportinal distribution of primary stolons amongst genets became strikingly more unequal (expressed as the Gini coefficient) with increasing density. Clonal growth was resolved into the number of primary stolons per stoloniferous genet and the extent of stolon branching (i.e. number of apices per primary stolon); both showed strongly negative density-dependence. Reproduction, expressed as the mean number of flowering capitula per stoloniferous genet, declined 15-fold with increasing density; although theoretically expected to be unity, greater values resulted from capitulum production by attached secondary rosettes and lower values reflected the increasing abortion rate of inflorescence buds with increasing density.Both the total number of apices produced per unit area and the corresponding number of reproductive apices were maximal at intermediate surviving densities (700–1,000 m^-2). The balance between reproductive and clonal growth may be expressed as the probability of an apical meristem producing a capitulum, that also peaked sharply at intermediate density. This finding does not conform with linear models that predict a shift from vegetative growth to sexual reproduction with increasing population density.     

Reproductive modes of three Ligularia weeds (Asteraceae) in grasslands in Qinghai-Tibet Plateau and their implications for grassland management

Ligularia virgaurea, L. sagitta and L. przewalskii are noxious weeds that are widely distributed in psychro-grasslands in the east of Qinghai-Tibet Plateau of China. However, little is known about reproductive modes and genetic diversity in the populations of L. virgaurea and L. sagitta under different environmental conditions, although such information is available for L. przewalskii. In this study, two experiments were conducted to examine the ex situ clonal growth of these weeds, and to determine the effects of reproductive modes on the diversity of their populations, using random amplified polymorphic DNA (RAPD) analysis. Transplanted seedlings of L. sagitta and L. virgaurea showed considerable clonal growth by forming rhizomes and ramets. The RAPD analysis showed that all individuals collected from natural populations were genetically different, indicating that no individuals originated from clonal growth. It is concluded that L. virgaurea and L. sagitta have the capacity for clonal growth, but the expression of their clonality is constrained by their growing conditions in natural grasslands. Sexual reproduction plays the major role in their population diversity in natural habitats, which requires that the control measures for these weeds should be targeted at the sexually reproducing populations.     

Seasonal patterns of carbon assimilation and allocation of a summer-green forest herb, <Emphasis Type="Italic">Parasenecio</Emphasis> <Emphasis Type="Italic">auriculata</Emphasis> (Senecioneae; Asteraceae)

Summer-green herbs inhabiting deciduous forests often put out aerial shoots under bright conditions before tree-canopy closure and grow until late summer under the closed canopy. Some of them produce leaves continuously even after the initiation of canopy closure, indicating an exploitation of the low light period. The manner of carbon assimilation during bright and shade periods within a growth season should reflect the seasonal patterns of vegetative growth and reproductive allocation of individual species. We examined the seasonal patterns of assimilation, partitioning of photosynthate between reproduction and storage, and the budget of reproduction of a perennial understory herb, Parasenecio auriculata. Although photosynthetic rates per unit leaf area decreased with the seasonal reduction in light level, net assimilation at the whole-plant level was maintained at a high level even after canopy closure owing to the increase in the total leaf area. Stored resource in tubers contributed to the rapid development of aerial shoots in the early season, and annual tuber growth was completed before flowering. Instant photosynthetic products considerably contributed to the maintenance of flowers but not to fruit development because of low assimilation rate during fruiting. These findings indicated that carbon assimilation during flowering contributes to sexual reproduction without influencing the development of storage organs. Stable carbon assimilation over summer by shade-acclimatized leaves enabled the maintenance of high productivity associated with high sexual reproduction.     

Different responses to shade of evergreen and deciduous oak seedlings and the effect of acorn size

An evergreen oak species, Cyclobalanopsis multinervis, and a deciduous oak species, Quercus aliena var. acuteserrata were grown from acorns under two light levels (full sunlight and shade at about 18 % of full sunlight, simulating the light intensities in forest clearings and gaps, respectively) for one growing season. Three hypotheses were tested: (i) the deciduous species grows faster than the evergreen species in forest gaps and clearings; (ii) the deciduous species responds more strongly in terms of growth and morphology to variation in light climate than the evergreen species; and (iii) seedling size is positively correlated to acorn size. The results showed: (i) at both light levels, the deciduous seedlings gained significantly more growth in biomass and height than the evergreen seedlings; (ii) both species produced significantly more biomass in full sunlight than in shade, without showing any significant difference in height between treatments. Increase in light intensity improved the growth of the deciduous seedlings more strongly; (iii) at a similar age, the deciduous seedlings showed a greater response in leaf morphology and biomass allocation to variation in light levels, but when compared at a similar size, biomass allocation patterns did not differ significantly between species; (iv) bigger acorns tended to produce larger seedlings, larger leaf sizes and more leaf area, between and within species. These differences demonstrate that the deciduous species is gap-dependent and has the advantage over the evergreen species in forest gaps and clearings.     

Infertile times: response to damage in genets of the clonal sedge Carex bigelowii

Tolerance to grazing is a plant trait that can be adaptive in systems where plants are subjected to a diversity of herbivore attack types. To test the tolerance ability of the clonal sedge Carex bigelowii, which is food plant to several herbivores in alpine and arctic areas, and the potential fitness costs of this tolerance, replicated units of genets were subjected to three levels of damage throughout three consecutive seasons. The three levels of treatment were no damage, light damage and heavy damage, and the damage was conducted by tearing off all plant material at 3 and 0 cm above-ground respectively. The genets had no tolerance under damage in terms of sexual reproduction. In terms of clonal reproduction the genets showed tolerance under light damage but not under heavy damage. However, no fitness cost was found for this tolerance ability, i.e. genets had higher reproduction and growth under no damage. The average ramet weight had a similar decrease under both a low and high damage treatment. Changed partitioning of biomass between plant parts and reduced concentration of total non-structural carbohydrates (TNC) in storage organs are possible mechanisms for the ability to uphold clonal reproduction in response to damage. There were no significant indications that tolerance ability or its fitness cost differed between genets. Our results suggest that when subjected to heavy damage genets will only reproduce vegetatively. Consequently, it seems C. bigelowii has evolved to allocate resources to the survival of an already successful genet rather than to a potential new genet of unknown success.     

The effect of shading on photosynthesis,growth, and regrowth following defoliation for Bromus tectorum

Summary The effect of full sunlight, 60%, or 90% attenuated light on photosynthetic rate, growth, leaf morphology, dry weight allocation patterns, phenology, and tolerance to clipping was examined in the glasshouse for steppe populations of the introduced grass, Bromus tectorum. The net photosynthetic response to light for plants grown in shade was comparable to responses for plants grown in full sunlight. Plants grown in full sunlight produced more biomass, tillers and leaves, and allocated a larger proportion of their total production to roots than plants grown in shade. The accumulation of root and shoot biomass over the first two months of seedling growth was primarily responsible for the larger size at harvest of plants grown in full sunlight. Plants grown under 60% and 90% shade flowered an average of 2 and 6 weeks later, respectively, than plants grown in full sunlight. Regrowth after clipping was greater for plants grown in full sunlight compared to those grown in shade. Even a one-time clipping delayed flowering and seed maturation; the older the individual when leaf area was removed, the greater the delay in its phenology. Repeated removal of leaf area was more frequently fatal for plants in shade than in full sunlight. For plants originally grown in full sunlight, regrowth in the dark was greater than for shaded plants and was more closely correlated to non-flowering tiller number than to plant size. This correlation suggests that etiolated regrowth is more likely regulated by the number of functional meristems than by differences in the size of carbohydrate pools. Thus, shading reduces the rate of growth, number of tillers, and ability to replace leaf area lost to herbivory for B. tectorum. These responses, in turn, intensify the effect of competition and defoliation for this grass in forests. B. tectorum is largely restricted to forest gaps at least in part because of its inability to acclimate photosynthetically, the influence of shade on resource allocation, and the role of herbivory in exacerbating these effects.     

Differences in allocation patterns in clonal and sexual offspring in a woodland pseudo-annual

We compared the patterns of allocation to reproduction among seed-derived and clonal offspring of a woodland pseudo-annual. Pseudo-annuals are clonal plants which survive the winter only as seeds and hibernacles produced by the rhizome system. Previous studies indicate that flowering is related to the size of these hibernacles. Since seedlings do not have a hibernacle, we did not expect that these plants would reproduce sexually. Assuming a trade-off between sexual and asexual reproduction, and assuming a linear relationship between vegetative plant weight and weight of all reproductive structures (i.e., rhizomes, hibernacles, inflorescences, and seeds), we expected that seed-derived plants would have a stronger biomass allocation to rhizomes and hibernacles. Since resource supply affects plant size, and thus hibernacle and seed production, we also subjected the plants to different levels of shade. At the start of the experiment seed-derived and clonal offspring hardly differed in total fresh weight. At the final harvest in September seed-derived and clonal offspring did not differ in vegetative plant weight (i.e., leaves, stems, and roots). Only light availability significantly affected these plant structures. As predicted, seed-derived plants did not flower in either of the light treatments. Seed-derived plants allocated more biomass to rhizomes and hibernacles, but this was only significant in the highest-light treatment. This result was due only to an increase in the number of hibernacles. Dry weight of single hibernacles was not affected by plant type. The ecological implications of this allocation pattern are discussed. Received: 2 October 1997 / Accepted: 8 March 1998     

Fine-scale spatial structure of genets and sexes in the dioecious plant <Emphasis Type="Italic">Dioscorea japonica</Emphasis>, which disperses by both bulbils and seeds

To understand the evolution of clonal reproduction and the diversity of clonal plants, it is necessary to clarify the characteristics of each clonal habit. There has been little research on whether bulbils alter spatial genetic structure (SGS) because of the lack of connection to maternal ramets. We used simple-sequence-repeat (SSR) markers to determine the fine-scale SGS of the dioecious plant Dioscorea japonica, which disperses both as bulbils and as seeds. We also evaluated the contributions of sexual and clonal reproduction and tested for spatial sex segregation (SSS). We discovered 111 genets from 394 ramets in a 2.8-ha plot. Genotypic richness (R = 0.28) and clonal diversity (Simpson’s D = 0.94, Fager’s E = 0.90) were high. We did not find SSS, suggesting that the population does not suffer from a shortage of mating pairs due to clonal reproduction. The Sp values revealed moderate SGS at the genet level (Sp = 0.013–0.014), and the genets intermingled at a local scale. Significant SGS at the ramet level showed that ramets within the same genet tended to aggregate. We also found a skewed clonal spatial distribution. The spatial extent of genets was positively correlated with the number of ramets within a genet. The contribution of bulbil production to the variance of parent–offspring gene dispersal was about one–fifth the contribution from sexual reproduction. These results suggest that the dispersal via bulbils affects the SGS in D. japonica, although its contribution to gene dispersal is small compared to the contribution of sexual reproduction.     

Effects of various environmental conditions on growth and reproduction of the sea hareAplysia oculifera (Adams and Reeve, 1850)

We studied the influences of food type, food quantity, water currents, starvation and light on growth and reproduction of the sea hareaplysia oculifera (Adams and Reeve, 1850) under laboratory conditions. Out of five species of algae served as food,Enteromorpha intestinalis promoted the fastest growth ofA. oculifera, Ulva spp. slower growth,Cladophora sp. allowed maintenance spp. slower growth,Cladophora sp. allowed maintenance of steady body mass, and the brown algaeColpomenia sp. andPadina pavonia were rejected by the sea hares. When sea hares were exposed to four levels of water currents, growth rates decreased as water currents increased. Sea hares fed on 50% ration grew slower than those fed on 100% ration (ad libitum). During 10 days of starvation sea hares lost weight, but when subsequently fed 100% ration they recovered and grew at a rate similar to those fed continuously with 100% ration. Under shade and under natural sunlight sea hares grew at the same rates. Whenever growth rates decreased, sea hares began to spawn at a smaller body size.A. oculifera demonstrated physiological plasticity that adapted them to varied and unpredictable environmental conditions. At different conditions of food availability they applied different tactics of resource allocation between growth and reproduction.     

Effects of size and growth rate on vegetative reproduction in Typha

Summary The objective of this study was to separate the effects of plant biomass and growth rate on vegetative reproduction in two species of cat-tail, Typha latifolia and T. angustifolia. Replicate clones of both species were grown under conditions of 100%, 42%, 24%, and 9% full sunlight with harvests at 41, 70, and 91 days after shading. T. angustifolia produced most of its vegetative offspring before the first harvest and increased biomass over the remainder of the experiment by increasing the size of its ramets. In contrast, T. latifolia produced vegetative offspring gradually throughout the experiment adding new ramets only after existing clones were of mature size. As a result of these differences in the cloning process, T. angustifolia showed little correlation between vegetative reproduction and clone size while T. latifolia showed a strong correlation between gegetative reproduction and clone size at the three highest light intensities. Growth rates, average clone size and vegetative reproduction were all reduced by reductions in light intensity for both species. However, no effect of growth rate on the relationship between clone size and vegetative reproduction in T. latifolia could be detected. T. latifolia showed greater survivorship and more biomass production under 9% light than T. angustifolia indicating a greater shade tolerance.     

Heavy metals modify costs of reproduction and clonal growth in the stoloniferous herb Potentilla anserina

We examined costs of sexual reproduction and clonal propagation, and their consequences for resource allocation in the clonal stoloniferous herb, Potentilla anserina, a typical pioneer species in disturbed areas. We used heavy-metal treatment in soil to create unfavourable growing conditions, because costs of reproduction are more likely to be expressed when resources are limited. We also studied whether heavy metals affect the plasticity of clonal growth form that enables the plants to avoid poor growing conditions. Ramets collected from field were grown in a greenhouse under the heavy-metal treatment consisting of a control and two levels of heavy-metals added in soil. Clonal propagation was costly in terms of total biomass of flowering ramets. Also the costs of sexual reproduction were detected in flowering ramets. Contrary to our predictions, the costs of flower production were visible in the control but not in the heavy-metal contaminated plants. Only the flowering ramets were able to produce longer stolons under heavy-metal stress, but the stolon biomass was not affected by heavy metals. Results indicate that clonal propagation and sexual reproduction may be costly for P. anserina. However, the costs are modified by heavy-metal contamination.Co-ordinating editor: J. Tuomi     

Trade-offs between clonal and sexual reproduction in Sagittaria latifolia (Alismataceae) scale up to affect the fitness of entire clones

? Many plants combine sexual reproduction with vegetative propagation, but how trade-offs between these reproductive modes affect fitness is poorly understood. Although such trade-offs have been demonstrated at the level of individual shoots (ramets), there is little evidence that they scale up to affect genet fitness. For hermaphrodites, reproductive investment is further divided between female and male sexual functions. Female function should generally incur greater carbon costs than male function, which might involve greater nitrogen (N) costs. ? Using a common garden experiment with diclinous, clonal Sagittaria latifolia we manipulated investment in reproduction through female and male sex functions of 412 plants from monoecious and dioecious populations. ? We detected a 1?:?1 trade-off between biomass investment in female function and clonal reproduction. For male function, there was no apparent trade-off between clonal and sexual reproduction in terms of biomass investment. Instead, male function incurred a substantially higher N cost. ? Our results indicate that: trade-offs between investment in clonal propagation and sexual reproduction occur at the genet level in S.?latifolia; and sexual reproduction interferes with clonal expansion, with investment in female function limiting the quantity of clonal propagules produced, and investment in male function limiting the nutrient content of clonal propagules.     

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.