Sink strength manipulation in branches of a Mediterranean woody plant suggests sink-driven allocation of biomass in fruits but not of nutrients in seeds

The relative autonomy of branches within an individual plant may favor different resource allocation responses after reproductive losses. The assessment of these reproductive strategies at the branch level before their integration at the plant level would provide more insight into how plants deal with reproductive losses. Here, we present a field experiment to assess changes in the allocation strategies at the branch level after sink strength manipulation in a Mediterranean woody plant (Cistus ladanifer). We applied three levels of removal of developing fruits (75, 25 and 0 %) on branches of the same plants, and measured their effects on resource allocation (biomass, nitrogen and phosphorus) and seed production after controlling for the effects of branch diameter and leaf weight per branch. Our results suggest that after experimental fruit thinning, C. ladanifer branches presented a sink-driven allocation of biomass to fruits but this was not the case for the allocation of nutrients to seeds, which could be driven by competition with leaf biomass. Reductions in biomass per fruit resulted in a reduction in seed output since the average weight per seed remained constant. From these results, it could be suggested that an heterogeneous distribution of fruit losses among the branches within a crown would produce a higher impact on reproductive output than a more equitable distribution.     

Trade-offs between vegetative growth and acorn production in Quercus lobata during a mast year: the relevance of crop size and hierarchical level within the canopy

The concept of trade-offs between reproduction and other fitness traits is a fundamental principle of life history theory. For many plant species, the cost of sexual reproduction affects vegetative growth in years of high seed production through the allocation of resources to reproduction at different hierarchical levels of canopy organization. We have examined these tradeoffs at the shoot and branch level in an endemic California oak, Quercus lobata, during a mast year. To determine whether acorn production caused a reduction in vegetative growth, we studied trees that were high and low acorn producers, respectively. We observed that in both low and high acorn producers, shoots without acorns located adjacent to reproductive shoots showed reduced vegetative growth but that reduced branch-level growth on acorn-bearing branches occurred only in low acorn producers. The availability of local resources, measured as previous year growth, was the main factor determining acorn biomass. These findings show that the costs of reproduction varied among hierarchical levels, suggesting some degree of physiological autonomy of shoots in terms of acorn production. Costs also differed among trees with different acorn crops, suggesting that trees with large acorn crops had more available resources to allocate for growth and acorn production and to compensate for immediate local costs of seed production. These findings provide new insight into the proximate mechanisms for mast-seeding as a reproductive strategy.     

Patterns of resource allocation in the dioecious alpine herb Aciphylla simplicifolia (Apiaceae)

Abstract Patterns of reproductive and vegetative biomass allocation were compared in male and female plants of the alpine herb Aciphylla simplicifolia. Male and female plants had similar vegetative biomass but differed in the pattern of resource allocation. Inflorescences of males and females were similar in weight at the time of flowering, but differed in biomass allocation to some structures within the inflorescences, particularly those associated with ovule production and pollinator attraction (number and size of flowers). At the time of fruit production, female inflorescences were 2.6 times heavier than at flowering with developing fruit six times heavier than flowers. In addition to the increase in biomass allocated to structures associated with the provisioning and dissemination of seed, support structures (main and side stalks) were also heavier. As a result of this additional investment of resources at the time of fruit production, the reproductive effort (RE) of female plants was much higher than that of males: 37% of above ground biomass compared with 21% for males. Differences in RE did not change with plant size; however, allocation to reproduction appeared to be a constant proportion of biomass over nearly all plant sizes sampled. These results show that sex‐specific resource allocation can be a complex of temporal and morphological patterns.     

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。     

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.     

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...     

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.     

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.     

Interactions in the patterns of vegetative growth and reproduction in woody dioecious plants

Summary Interactions between vegetative growth and reproduction were evaluated in Peumus boldus, Lithraea caustica and Laretia acaulis, three woody dioecious species in central Chile. Phenological observations were made periodically on marked branches of male and female plants, and biomass allocation (dry weight) to vegetative and reproductive tissues was measured. The magnitude of flowering was evaluated in groups of plants in three successive seasons. The patterns of activities are species- and sex-dependent, and cycles of 2–4 years have been established. Branches that produce flowers either do not grow or grow less than branches without flowers, and males and females have differential resource allocation: male branches attain higher biomass values. Groups of plants show seasonal behavior that suggest synchrony in their reproductive activities.     

Effects of defoliation and shading on the physiological cost of reproduction in silky locoweed Oxytropis sericea

Background

The production of flowers, fruits and seeds demands considerable energy and nutrients, which can limit the allocation of these resources to other plant functions and, thereby, influence survival and future reproduction. The magnitude of the physiological costs of reproduction depends on both the factors limiting seed production (pollen, ovules or resources) and the capacity of plants to compensate for high resource demand.

Methods

To assess the magnitude and consequences of reproductive costs, we used shading and defoliation to reduce photosynthate production by fully pollinated plants of a perennial legume, Oxytropis sericea (Fabaceae), and examined the resulting impact on photosynthate allocation, and nectar, fruit and seed production.

Key Results

Although these leaf manipulations reduced photosynthesis and nectar production, they did not alter photosynthate allocation, as revealed by ¹³C tracing, or fruit or seed production. That photosynthate allocation to reproductive organs increased >190 % and taproot mass declined by 29 % between flowering and fruiting indicates that reproduction was physiologically costly.

Conclusions

The insensitivity of fruit and seed production to leaf manipulation is consistent with either compensatory mobilization of stored resources or ovule limitation. Seed production differed considerably between the two years of the study in association with contrasting precipitation prior to flowering, perhaps reflecting contrasting limits on reproductive performance.     

Cost of reproduction in the perennial herb Asphodelus albus (Liliaceae)

The cost of reproduction has been studied in two populations of the polycarpic herb Asphodelus albus under natural conditions The percentage of plants with flowers was determined in four sites and varied markedly among them The occurrence of reproduction was size-dependent, increasing flowering probability with plant size The cost of reproduction was assessed in terms of modular growth in reproductive plants relative to modular growth in vegetative ones I compared the modular growth of vegetative and reproductive plants considering two different densities m each of two populations Neither incidence of flowering nor modular growth were affected by density Flowering plants exhibited a withinramet demographic cost (in terms of modular growth) relative to non-flowering ramets in one population but not in the other This cost was greater in larger plants These results were concordant with the occurrence of flowering at both sites Both populations exhibited size-dependent patterns of allocation to reproduction, but no significant relationships were found between allocation to reproduction and cost of reproduction The data presented demonstrate differences in the cost of reproduction within a species This cost might determine whether a plant begins the reproduction, but probably have no effect on the reproductive allocation since the weight of the reproductive structures was not related to modular growth     

Reproductive correlates of mating system variation in Eichhornia paniculata (Spreng.) Solms (Pontederiaceae)

Comparative studies of related plant species indicate that evolutionary shifts in mating systems are accompanied by changes in reproductive attributes such as flower size, floral morphology, and pollen/ovule ratio. Recent theoretical work suggests that patterns of investment in reproduction should also change with the mating system. In a glasshouse study, we investigated the extent to which mating system differences among populations of Eichhornia paniculata (Pontederiaceae) were correlated with changes in allocation to male and female function, floral display, and the regulation of investment in reproduction through fruit and ovule abortion. Significant differences in the amount of biomass allocated to reproductive structures were evident among six populations of E. paniculata. As predicted by sex allocation theory, the proportion of dry weight allocated to male function decreased with the outcrossing rate of populations. Six of the eight attributes used to characterize floral display also differed significantly among populations. However, with the exception of two attributes describing the number of flowers produced by inflorescences, these were not correlated with outcrossing rate. Levels of fruit and ovule abortion were determined in two populations with contrasting mating systems under different nutrient and pollination treatments. Virtually all fruits initiated by plants from a self-fertilizing population were matured, while the amount of fruit abortion in an outcrossing population increased with flower production. Ovule abortion was low in both populations. Our results demonstrate that the evolution of self-fertilization in E. paniculata is associated with changes in investment to reproduction that normally distinguish selfing and outcrossing species.     

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.     

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.     

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.     

濒危植物长柄双花木自然种群生殖构件的时空动态

根据野外观测数据,研究了濒危植物长柄双花木(Disanthus cercidifolius var.longipes)在不同生境条件下自然种群开花物候变异以及生殖构件的时空动态.结果表明,不同群落类型中长柄双花木开花物候存在一定差异,在纯林中个体的平均花期最长,达75d,而个体间开花同步性最低,仅为0.717;竹林中的个体花期最短,只有47d,而同步指数最高,达0.968.生殖枝数量及花序数均随个体年龄增长而增加,在年龄为30～35年时达最高,此后又呈下降趋势;在海拔为810m左右处,个体生殖枝数及着生的花序数均达最大值,且冠层间差异明显.纯林中个体着生的生殖枝数及每生殖枝上着生的花序数均最高,分别为411.39和7.857;竹林中个体每生殖枝着生的花序数显著低于其它群落.年龄小的个体,各时期生殖构件败育率均高于年龄大的个体;另外,生殖构件的败育与海拔高度间无相关性,而与构件发育时间、分布的树冠层次以及所处的群落类型相关.影响生殖构件败育的主要因子是光照条件.     

薇甘菊不同生长方式下的繁殖特征比较

利用实验生态学方法, 对攀缘生长与伏地生长的薇甘菊(Mikania micrantha)的形态特征、生物量分配和繁殖特征进行了比较研究。结果表明: 在潮湿生境条件下, 攀缘生长的植株与伏地生长的植株相比对主茎的生物量投资较大, 而对分枝茎的生物量投资则相对较小; 但在干旱生境条件下, 攀缘生长的植株对主茎和分枝茎的生物量投资均较伏地生长的植株小。在潮湿与干旱生境中, 攀缘植株的繁殖投资分别为0.429 g·g^–1和0.342 g·g^–1, 显著高于相同生境中的伏地植株。在薇甘菊种群的补充与更新过程中, 攀缘生长的植株以有性生殖占主导地位, 在潮湿与干旱生境中产生的个体分别占种群数量的84.7%和62.6%; 伏地生长的植株则主要以无性(克隆)繁殖为主, 在潮湿与干旱生境中有性生殖产生的个体仅占种群数量的40.4%和35.9%。生长方式与土壤水分二因子互作效应对薇甘菊的生物量分配和繁殖指标影响均达到显著水平(P<0.05)。通过研究得出以下结论: 不同生长方式的薇甘菊对环境条件具有不同的生长与繁殖适应对策。     

Effects of Local Plant Density and Incomplete Dichogamy on the Reproductive Success of the Rare Neotropical Palm Geonoma epetiolata

Density dependent processes are known to influence reproduction and establishment of plant populations. In this study, we evaluated the effects of local density and sexual expression on the reproductive success of the rare palm species Geonoma epetiolata in Costa Rica. We classified individuals in two density categories based on distances to the two nearest neighbors and recorded the occurrence of sexual overlap on each individual. Overlap between pistillate and staminate flowers in monoecious plants allows geitonogamous pollination, potentially reducing dependence on mates for reproduction. We measured plant size and light availability, and evaluated the influence of these variables on pollination success, fruit production, fruit abortion and seed mass. Pollination success significantly increased with density but there was no effect of sexual overlap. In contrast, there was no effect of density on the probability of initiated fruit, and a negative effect on fruit set. Fruit abortion was lowest in isolated plants with no sexual overlap. Plant size and light did not contribute to variation in reproductive success. In conclusion, the pollination advantage of plants in dense neighborhoods does not compensate reproductive losses incurred by fruit abortion, possibly due to mating among genetically related individuals. Geonoma epetiolata is threatened by habitat loss and poaching of seeds for the horticultural market. High fruit abortion rates associated with density in G. epetiolata suggest that seed collections from the remnant wild populations of this palm species may cause bottlenecks that further threaten population viability.     

Patterns of vegetative growth and reproduction in relation to branch orders: the plant as a spatially structured population

The patterns of vegetative growth and reproduction in relation to orders of terminal branches were examined in the evergreen woody plant, Eurya japonica. The branch order number was determined centrifugally. The trunk was given order number 1; branches issuing directly from the trunk were order 2; branches growing on order 2 branches were order 3, and so on. The results of this study show the differential patterns of vegetative growth and reproduction in relation to the branch orders. Lower-order shoots of terminal branches grew more, but produced few flowers. On the other hand, for the higher-order terminal branches, shoot growth was very limited but flowering was more intense. The results show that a tree can be interpreted not as a mere population of equivalent modules but as a spatially structured population. Thus, it is essential to consider the position of modules within the branch system when patterns of vegetative growth and reproduction are examined. It is hypothesized that the difference in the opportunity cost in relation to the branch orders is the main cause of the spatial structure for patterns of vegetative growth and reproduction. Furthermore, for same-order terminal branches, current-year shoot elongation was independent of flowering intensity. These results suggest that this species only invests resources in reproduction that are surplus to its requirements for the functions associated with survival, such as growth and/or storage. Received: 22 July 1999 / Accepted: 12 January 2000     

Interpreting reproductive allometry: Individual strategies of allocation explain size-dependent reproduction in plant populations

Size-dependent or allometric relationships between reproductive and vegetative size are extremely common in plant populations. Reproductive allometry where plant size differences are due to environmental variability has been interpreted both as an adaptive strategy of plant growth and allocation, and as the product of fixed developmental constraints. Patterns of development are crucial in defining reproductive allometry but development is not fixed across individuals. For example, environmental adversity (e.g. resource impoverishment) tends to favor reproduction at relatively small sizes – an adaptive response to environmental adversity. While small individuals may have lower reproductive output than large individuals, all plants should maximize their reproductive output and relative allocation to reproduction may be constant across sizes. Thus, where individual plants within a population initiate reproduction at different sizes, no significant reproductive allometry is an appropriate null expectation. Reproductive allometry occurs in plant populations where initiating reproduction at small sizes yields relatively high or low reproductive size at final development. Both of these outcomes are common in plant populations. Our interpretation of reproductive allometry combines previous adaptive and developmental constraint interpretations, and is the first to successfully explain the range of relationships in plant populations where relative allocation has been observed to increase, decrease or remain constant will increasing plant size.