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 effect of pollination on resource allocation among sexual reproduction,clonal reproduction,and vegetative growth in <Emphasis Type="Italic">Sagittaria potamogetifolia</Emphasis> (Alismataceae)

In order to evaluate the effects of pollination on resource allocation in the marsh herb Sagittaria potamogetifolia, experimental manipulation of pollination efficiency on the pattern of resource allocation was accessed by the proportion of dry weight measurements of sexual, vegetative, and clonal organs. In trials where half of the flowers were pollinated, a significant increase of resource allocated to sexual production and decrease to vegetative production resulted compared to plants that received no pollination. In trials where pollination was 100%, these two reproductive components showed the same trend, but less dramatically. This may support the idea that the trade-offs would be more pronounced when the resource was scarce. Besides, a higher inflorescence production with a lower fruit reproduction occurred as a consequence of decreased pollination level. This increased inflorescence production may be a mechanism to promote outcrossing by enhancing floral attraction or by synchronizing reproductive activity with insect pollinators. Examination for possible trade-offs in resource allocation revealed that there was also a trade-off caused by pollination between fruits plus flowers and bulbils production, which might have detrimental effects on the survival of individuals and populations, but promotes outcrossing and genetic variability.     

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。     

Gender-specific costs of reproduction on vegetative growth and physiological performance in the dioecious shrub Corema album

Background and Aims

Reproductive costs imply trade-offs in resource distribution at the physiological level, expressed as changes in future growth and/or reproduction. In dioecious species, females generally endure higher reproductive effort, although this is not necessarily expressed through higher somatic costs, as compensatory mechanisms may foster resource uptake during reproduction.

Methods

To assess effects of reproductive allocation on vegetative growth and physiological response in terms of costs and compensation mechanisms, a manipulative experiment of inflorescence bud removal was carried out in the sexually dimorphic species Corema album. Over two consecutive growing seasons, vegetative growth patterns, water status and photochemical efficiency were measured to evaluate gender-related differences.

Key Results

Suppression of reproductive allocation resulted in a direct reduction in somatic costs of reproduction, expressed through changes in growth variables and plant physiological status. Inflorescence bud removal was related to an increase in shoot elongation and water potential in male and female plants. The response to inflorescence bud removal showed gender-related differences that were related to the moment of maximum reproductive effort in each sexual form: flowering in males and fruiting in females. Delayed costs of reproduction were found in both water status and growth variables, showing gender-related differences in resource storage and use.

Conclusions

Results are consistent with the existence of a trade-off between reproductive and vegetative biomass, indicating that reproduction and growth depend on the same resource pool. Gender-related morphological and physiological differences arise as a response to different reproductive resource requirements. Delayed somatic costs provide evidence of gender-related differences in resource allocation and storage. Adaptive differences between genders in C. album may arise through the development of mechanisms which compensate for the cost of reproduction.     

三种同域分布喉毛花的繁殖分配

以青藏高原高寒草甸中三种同域分布的喉毛花为研究对象,通过比较三个种的植株性状和繁殖分配,探讨繁殖分配的种间差异及其与植株个体大小的关系。结果表明:(1)三个种的植株高度、顶花大小和单株花数目、繁殖分配均存在种间差异,这可能与其各自的交配系统和具体的生境以及相应的生活史对策有关;(2)在三种喉毛花中,投入到营养器官和繁殖器官的绝对资源量均呈显著正相关,未检测到植株生长和繁殖间的权衡关系;(3)三个种的个体大小与繁殖器官生物量均呈显著正相关,而与繁殖分配均呈显著负相关,这表明个体越大,繁殖投入越高,而繁殖分配越低,与以往研究结果一致,这可能是由于繁殖分配与个体大小之间存在异速关系。     

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.     

Size-dependent allocation to sexual and vegetative reproduction in four clonal composites

Summary Populations of Silphium speciosum, Vernonia baldwinii, Solidago canadensis and Pityopsis graminifolia were studied to determine whether biomass allocation to sexual and vegetative reproduction and the balance between them were size-dependent and whether interpopulation differences in allocation patterns could be predicted from differences in population size distributions. All four species showed strong linear relationships between inflorescence mass and vegetative mass with negative y-intercepts. As a result, sexual reproductive effort (SRE) was a monotonically increasing function of ramet size. Genet size was a poor predictor of SRE. In each species, the regression parameters of these relationships differed significantly between burned and unburned habitats indicating size-independent interpopulation differences in patterns of reproductive effort as well as sizedependent effects. Interpopulation variation in vegetative reproductive effort (VRE) was greater than variation in SRE, but neither VRE nor the pattern of partitioning of VRE among daughter rhizomes showed significant relationships to plant size.     

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...     

羊草种群能量生殖分配的研究

对羊草种群能量生殖分配规律研究表明,在羊草种群中,各构件热值的大小顺序为穗〉营养枝叶鞘〉生殖枝叶鞘〉营养枝叶〉营养枝茎〉生殖枝茎〉生殖枝叶〉凋落物;营养生长和生殖生长的能量分配比例的大小顺序为营养枝〉根茎〉凋落物〉生殖枝;各构件能量分配比例的季节动态也有差异,在羊草种群的生殖枝构件中,茎和叶鞘的分配比例较大,而穗和叶的能量分配比例较小。     

EFFECTS OF LIGHT AND DENSITY ON RESOURCE ALLOCATION IN A FOREST HERB,ASTER ACUMINATUS (COMPOSITAE)

Recent studies show that forest herbs have low sexual reproductive efforts (SRE) and high vegetative reproductive efforts (VRE) and that a variety of factors may affect the levels of reproduction and the distribution patterns of forest species. Aster acuminatus is a micro-successional forest herb common in northern New England. Resource allocation patterns were determined for seven aster patches varying in density and light level. Plant density affects neither plant size nor any aspect of resource allocation, and VRE remains constant over all density and light levels. Light level is, however, significantly correlated with patch location, average plant size and average patch SRE. Only large plants produce flowers suggesting that light affects sexual reproduction indirectly through effects on plant size. The high rhizome and low seed survivorship of this species may partly explain the constant VRE and varying SRE levels, although the resource allocation patterns observed do not correspond well with the predictions of certain theoretical models.     

Reproductive trade-offs in genetically distinct clones ofVaccinium macrocarpon,the American cranberry

Patterns of resource allocation to growth, current reproduction, and potential future reproduction were quantified in six genetically distinct cultivars ofVaccinium macrocarpon. For all cultivars (genotypes), vegetative size is positively correlated with some measures of current reproduction (fruit and flower number) but negatively correlated with others (seed number per fruit, seed weight per fruit). Vegetative growth in the current year is significantly related to the production of reproductive terminal buds, a measure of the potential for reproduction in the following year. Stems with low levels of current reproduction — lower flower number, fruit number, and seed weight — were more likely to form reproductive terminal buds than stems with higher levels of current reproduction. Individual genotypes differed significantly for vegetative size, fruit number, fruit weight, seed number, and seed weight, as well as for the frequency of fruiting stems and reproductive terminal buds produced. Genotypes were segregated in principal component space, indicating overall differences between them in allocation to the suite of variables measured. These results indicate the possibility of fitness differences among cultivars due to genetically determined allocation strategy, which has implications for fitness differences among genotypes within natural populations.     

Allometric Effects of <Emphasis Type="Italic">Agriophyllum squarrosum</Emphasis> in Response to Soil Nutrients,Water, and Population Density in the Horqin Sandy Land of China

We monitored the allometric effects for greenhouse-grown Agriophyllum squarrosum plants in response to variations in population density and the availability of soil nutrients and water. Biomass allocations were size-dependent. The plasticity of roots, stems, leaves, and reproductive effort was “true” in response to changes in nutrient content. At a low level of soil minerals, plants allocated more resources to the development of roots and reproductive organs than to leaves, but data for stem allocations were consistent for tradeoffs between the effects of nutrients and plant size. The plasticities of leaf allocation and reproductive effort were “true” whereas those of root and stem allocations were “apparent” in response to fluctuations in soil water, being a function of plant size. Decreasing soil water content was associated with higher leaf allocation and lower reproductive effort. Except for this “apparent” plasticity of leaf allocation, none was detected with population density on biomass allocation. Architectural traits were determinants of the latter. For roots, the determining trait was the ratio of plant height to total biomass; for stems and reproduction, plant height; and for leaves, the ratio of branch numbers to plant height.     

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.     

Flowering does not decrease vegetative competitiveness of Lolium perenne

The theory of life-history evolution commonly assumes a trade-off between sexual and vegetative reproduction. Hence, production of flowers and fruits should have measurable costs in terms of reduced vegetative growth. This trade-off may be meaningful for breeding of forage and turf grasses as reduced flowering could free resources and increase productivity. But if so, less-flowering cultivars might be more competitive and invade natural swards. We tested for costs of sexual reproduction on vegetative propagation and competitiveness of the perennial grass Lolium perenne, one of the most important forage and turf grasses worldwide. We used the differences in vernalisation requirement between northern and southern European provenances to manipulate the degree of flowering. Over three growing seasons, we counted the number of flower stems and measured the clone diameter. The vernalisation treatments were successful in producing clones with largely differing degrees of flowering. However, we found no negative correlation between flowering and vegetative propagation and competitiveness. Early and strongly flowering southern provenances showed less clonal growth and higher mortality, but within provenances the response of clone diameter to flowering was positive or neutral. We conclude that investment of resources into flowering has no measurable costs on vegetative propagation and competitiveness of L. perenne. The apparent lack of costs of sexual reproduction could be explained by bet-hedging strategy that is focused on survival and growth rather than reproductive effort in order to maximise the life-time fitness.     

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.     

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.     

Individual resource allocation to vegetative growth and reproduction in subgenus <Emphasis Type="Italic">Cyclobalanopsis</Emphasis> (<Emphasis Type="Italic">Quercus</Emphasis>, Fagaceae) trees

The resource allocation for vegetative growth and female reproduction in three tree species of subgenus Cyclobalanopsis (Quercus, Fagaceae), i.e., Q. salicina, Q. sessilifolia, and Q. acuta, were examined on a per-individual basis in two consecutive reproductive seasons, in order to test whether these trees fit the predictions of the masting hypotheses about resource matching versus resource switching. Since the three Quercus species have a biennial fruiting habit, it takes 3 years for the observation of two reproductive events. Female flower and acorn production per tree were investigated by using a seed-trap method and a numerical analysis of seed dispersal. The net production of each individual was estimated as the sum of the annual increase in the dry mass of vegetative organs and reproductive investment per tree. In the data analyses, the three species were pooled, since all 12 sample trees of the subgenus apparently showed masting in the same year, with no exceptions. Female flower and acorn production per individual tree changed considerably between years. The net production per tree increased with tree size, but did not differ between years. Therefore, the reproductive allocation (proportion of a plant’s annual assimilated resources which are used for reproduction) differed dramatically between years. On the other hand, within a year, the reproductive allocation increased with increasing net production per tree. These results suggest that the switching of resource allocation between years within an individual are occurring in subgenus Cyclobalanopsis species, and the intensity of the switching increases with increasing tree size.     

Effects of parasitic castration on plant resource allocation

It has been proposed that host castration is a parasite strategy to reallocate host resources from reproductive to vegetative functions to increase parasite fitness. Since resource partitioning between reproduction and vegetative growth can affect host life-history traits, parasite effects on resource allocation can affect both plant fitness and host-parasite coevolution. Field and greenhouse experiments were used to investigate the effects of host castration by the fungus Atkinsonella hypoxylon on the resource allocation and architecture of the grass Danthonia spicata. The results indicate that non-infected D. spicata can reallocate resources from reproduction to vegetative growth when resource allocation to reproduction is prevented. However, I found no evidence that fungal castration causes reallocation of resources from host reproduction to vegetative growth. Instead, infection reduces host biomass and the fungus directly utilizes resources that would have been used for host reproduction for its own reproduction. Received: 25 March 1999 / Accepted: 24 October 1999     

Does the silver moss Bryum argenteum exhibit sex-specific patterns in vegetative growth rate,asexual fitness or prezygotic reproductive investment?

Background and Aims

Expected life history trade-offs associated with sex differences in reproductive investment are often undetected in seed plants, with the difficulty arising from logistical issues of conducting controlled experiments. By controlling genotype, age and resource status of individuals, a bryophyte was assessed for sex-specific and location-specific patterns of vegetative, asexual and sexual growth/reproduction across a regional scale.

Methods

Twelve genotypes (six male, six female) of the dioecious bryophyte Bryum argenteum were subcultured to remove environmental effects, regenerated asexually to replicate each genotype 16 times, and grown over a period of 92 d. Plants were assessed for growth rates, asexual and sexual reproductive traits, and allocation to above- and below-ground regenerative biomass.

Key Results

The degree of sexual versus asexual reproductive investment appears to be under genetic control, with three distinct ecotypes found in this study. Protonemal growth rate was positively correlated with asexual reproduction and sexual reproduction, whereas asexual reproduction was negatively correlated (appeared to trade-off) with vegetative growth (shoot production). No sex-specific trade-offs were detected. Female sex-expressing shoots were longer than males, but the sexes did not differ in growth traits, asexual traits, sexual induction times, or above- and below-ground biomass. Males, however, had much higher rates of inflorescence production than females, which translated into a significantly higher (24x) prezygotic investment for males relative to females.

Conclusions

Evidence for three distinct ecotypes is presented for a bryophyte based on regeneration traits. Prior to zygote production, the sexes of this bryophyte did not differ in vegetative growth traits but significantly differed in reproductive investment, with the latter differences potentially implicated in the strongly biased female sex ratio. The disparity between males and females for prezygotic reproductive investment is the highest known for bryophytes.     

EVOLUTION OF DIOECY IN BUCHLOE DACTYLOIDES (GRAMINEAE): TESTS FOR SEX-SPECIFIC VEGETATIVE CHARACTERS,ECOLOGICAL DIFFERENCES,AND SEXUAL NICHE-PARTITIONING

Buchloe dactyloides is a perennial dioecious grass in which male and female inflorescences are so strikingly dimorphic that they were originally assigned to different genera. The objective of this paper is to present the results of tests for sex-specific vegetative characters, ecological differences, and sexual niche-partitioning, combining them with prior information on the reproductive biology of Buchloe for an evaluation of the key factors leading to the evolution of dioecy and sexual dimorphism. Field and greenhouse data were collected from Oklahoma and Kansas populations on vegetative characters, allocation to reproduction, and relative growth and competitive success along resource gradients. Except for greater susceptibility to leaf rust by males, there were no significant differences between males and females in vegetative characters, total biomass, or reproductive effort. Field studies of spatial distributions of males and females failed to show any relation to soil, topography, or soil moisture. In a 45-month greenhouse experiment starting at the seedling stage, the relative growth and competitive success of randomly paired individuals showed no evidence for differential competitive success or for niche-partitioning of males and females. The “outcrossing advantage” and subsequent sexual specialization of the female inflorescence appear to be the major factors underlying this dimorphic system.