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。     

Altitudinal Variations in Reproductive Allocation of Bergenia purpurascens (Saxifragaceae)

The strategy of resource allocation between vegetative and reproductive functions, quantitative relationship between size and reproductive output are central aspects of plant life history. To test the tactics of resource allocation and its altitudinal trend, we examined the reproductive allocation (RA) of Bergenia purpurascens (Saxifragaceae), in six populations along a shady slope in Sejila Mountain of southeast Tibet, at an altitude gradient from 4 200 m to 4 640 m. Our results showed that (1) with increasing altitude, vegetative biomass, reproductive biomass, total aboveground biomass, flower number per plant and length of flower stalk decreased significantly, but the number of leaves did not change greatly. However, the change of RA did not show a monotonic trend when altitude increased, shifting from significantly decreasing below the tree line to slightly increasing above it; (2) vegetative biomass was positively correlated with reproductive biomass, but negatively correlated with RA in all populations, but the level of significance was different among the populations; (3) RA decreased with individual size in all populations, whereas the relationship between absolute resource allocated to reproduction and individual size was allometric; (4) reproductive allometry and a size threshold for reproduction did exist in this alpine perennial, but the obvious altitudinal trend was only found along the populations below the tree line, not above it. We then concluded the altitude could not fully explain the change of resource allocation strategy of this alpine perennial, and different effects of size and habitat on RA may result from various environmental constraints along the altitudinal gradient or genetic background. Therefore, each individual within a population will follow its own developmental trajectory shaped by its genotype and the habitats. The most innovative finding was plant adaptation and resource trade off might be sharply altered at the tree line, which is a sensitive area in alpine mountains. Further investigations are needed to better understand the relationship between the reproductive allocation and changing environmental conditions.     

川西风毛菊果期资源分配对海拔的响应

以分布于青藏高原东缘的川西风毛菊为试验材料,研究了其不同海拔高度16个居群的果期资源分配。结果显示:1)随着海拔的升高,个体大小、繁殖器官生物量、营养器官生物量、根系质量、茎叶质量以及每株植物种子总数量均不断减小,但种子百粒重不断增加;2)繁殖分配和根系分配与海拔呈正相关关系,营养分配和茎叶分配与海拔呈负相关关系;3)果期繁殖分配和营养分配、根系分配和茎叶分配以及种子数量与百粒重之间均存在资源分配上的权衡。研究结论:1)海拔作为外界因子对川西风毛菊果期各生物量及资源分配有显著的影响;2)随着海拔的升高,川西风毛菊通过增加繁殖分配,根系分配以及种子百粒重来适应胁迫环境,提高自身的适合度。     

Reproductive allocation in hermaphrodite and female plants of Sidalcea oregana SSP. spicata (Malvaceae) using four currencies

Reproductive allocation was investigated in female and hermaphrodite plants of gynodioecious Sidalcea oregana ssp. spicata. Total reproductive investment and partitioning of that investment was documented at the level of whole plants in terms of four ecologically relevant currencies: biomass, nitrogen, phosphorus, and potassium. Nutrient augmentations in the field confirmed that nutrients were limiting plant vegetative growth and propensity to flower; thus the use of these nutrients as currency was appropriate. Once the effects of plant size were removed, the sex morphs allocated similar total amounts of biomass, nitrogen, phosphorus, and potassium to reproduction, but partitioned those differentially. For any given individual size, females allocated larger proportions of their reproductive resource budgets to seeds. Hermaphrodites' reproductive investment in pollen and flowers was allocated at the expense of allocation to seeds. These data are relevant to the evolution of gynodioecy from hermaphroditism and support the hypothesis that females reallocate resources not spent on pollen to seeds.     

岩白菜（虎耳草科）不同海拔居群的繁殖分配

资源分配策略是植物生活史研究的重要内容之一,植物用于繁殖的相对资源比例（即繁殖分配）与植株的生活史特征、个体大小及植株的生境密切相关。本文研究了藏东南色季拉山一个阴坡上海拔4200m～4640m范围内6个不同居群的虎耳草科多年生草本植物岩白菜（Bergenia purpurascens）的繁殖分配特征,结果发现：（1）繁殖器官生物量、营养器官生物量、地上部分总生物量、花数目、花序轴长度均随海拔的升高而显著降低,而叶数目随海拔变化不大,繁殖分配值则先降低后升高,转折点在林线过渡带（海拔4400m）处;（2）各居群（海拔4300m居群除外）营养器官生物量与繁殖器官生物量均显著正相关,而营养器官生物量与繁殖分配则负相关,但各居群的显著性不同;（3）各居群繁殖器官生物量与植株个体大小（营养器官生物量）呈不同程度的异速增长,而繁殖分配则与植株个体大小负相关;（4）各居群植株都存在一个繁殖所需的个体大小阈值,而且这一阈值在林线以下区域随海拔的升高而显著增大,在林线以上区域变化不显著。研究结果表明,海拔并不是影响岩白菜繁殖分配策略的唯一生态因子,不同居群的生境状况和植株个体大小都与其资源分配策略密切相关,高山地区林线的存在对植物资源的权衡方式会产生巨大影响。     

Size-dependent resource allocation among vegetative propagules and male and female functions in the forest herb Laportea bulbifera

Reproductive resource investment among vegetative propagules and male and female sexual function and their size-dependence were investigated in a perennial forest herb, Laportea bulbifera . A theoretical model based on fitness gain curves predicts that optimal investments in three reproductive modes will increase with plant size if fitness returns in all three modes increase but become saturated with investment. In a field population, large plants of L. bulbifera produced both male and female inflorescences with propagules, while small plants produced only vegetative propagules. Biomass of propagules, male inflorescences, and infructescences with achenes were all positively correlated with plant size. The increase in investment with plant size was larger for propagule production than for sexual reproduction. The relationship between propagule biomass and plant size was constant irrespective of year, while the relationship between the biomass of sexual reproductive organs and plant size differed between two successive years. Annual change of individual sex expression was investigated for 25 transplanted plants. Although each plant changed its sex expression variously among male, female and bisexual from year to year, 23 out of 25 plants produced both male and female inflorescences in at least one year. The number of viable (germinated and survived) offspring from seeds was not significantly different from the number from propagules. The production cost of a propagule was higher than that of a seed. Resource allocation theory does not seem to be applicable to size-dependent resource allocation, especially the allocation between seeds and propagules in this species.     

Regional and local effects on reproductive allocation in epicormic and lignotuberous populations of <Emphasis Type="Italic">Banksia menziesii</Emphasis>

Reproductive allocation (RA) is a measure of how resources (biomass, nutrients) are partitioned between reproductive structures and the rest of the plant. For plants that resprout after fire, the percentage of resources allocated to reproduction may vary depending on their resprouting ability. Our study examines the percentage RA (biomass, N, P, K) and nutrient content of current season’s growth in southern (Swan Coastal Plain) epicormic and northern (Eneabba Plain) lignotuberous resprouter populations of Banksia menziesii (Proteaceae), a species endemic to nutrient-impoverished sandplains of southwestern Australia. Within each population, plants along road edges were compared with plants not associated with road edges. There was no difference in total nutrient content of current year’s growth between both resprouting types, except that total K in the shoots of lignotuberous populations was >2 times that in the epicormic populations. Non-road lignotuberous plants allocated twice the biomass, N and P, and 13.5 times the K, to reproduction as non-road epicormic plants. Lignotuberous populations had the highest RA (17–34% of biomass, N, P, K), with non-road epicormic populations the lowest RA (3–15%). This can be viewed as an adaptive (ultimate) response to the poorer postfire survival and recruitment conditions where the lignotuberous populations occur. Total biomass and nutrient content of road-edge plants was 2–3 times that of non-edge plants. Lignotuberous populations in both road positions allocated the same fraction of biomass, N and P to reproduction, whereas road-edge populations allocated 10% less K than non-road. Road-edge epicormic populations allocated 5–10% more biomass, N, P and K to reproduction than non-road populations. This can be viewed as an ecophysiological (proximate) response to the better growing conditions created by the roadways that may also ultimately have an adaptive explanation.     

Production of male flowers does not decrease with plant size in insect-pollinated Sagittaria trifolia, contrary to predictions of size-dependent sex allocation

It has been proposed that relative allocation to female function increases with plant size in animalpollinated species.Previous investigations in several monoecious Sagittaria species seem to run contrary to the prediction of size-dependent sex allocation (SDS),throwing doubt on the generalization of SDS.Plant size,phenotypic gender,and flower production were measured in experimental populations of an aquatic,insect-pollinated herb Sagittaria trifolia (Alismataceae) under highly different densities.The comparison of ramets produced clonally can reduce confounding effects from genetic and environmental factors.In the high-density population,48％ of ramets were male without female flowers,but in the low-density population all ramets were monoecious.We observed allometric growth in reproductive allocation with ramet size,as evident in biomass of reproductive structures and number of flowers.However,within both populations female and male flower production were isometric with ramet size,in contrast to an allometric growth in femaleness as predicted by SDS.Phenotypic gender was not related to ramet size in either population.The results indicated that large plants may increase both female and male function even in animal-pollinated plants,pointing towards further studies to test the hypothesis of size-dependent sex allocation using different allocation currencies.     

Plasticity of female reproductive resource allocation depends on the presence or absence of prior environmental sex determination in Ceratopteris richardii

Resource allocation plasticity enables individuals to alter patterns of nutrient use between reproductive and vegetative output to better fit their current environment. In sexually labile plant species, abiotic environmental factors can influence expression of dimorphic gender, resulting in environmental sex determination (ESD), which potentially reduces the need for plasticity of resource allocation by preemptively matching an individual’s future nutrient demands to resource availability in its location. Ceratopteris richardii gametophytes exhibit gender‐dependent differences in relative carbon and nitrogen content, and ESD in certain nutrient environments. This study examined whether prior ESD in C. richardii gametophyte populations reduced subsequent plasticity of reproductive allocation compared to instances where no ESD occurred, by quantifying phenotypic responses to reduced P, N, or CO₂ availabilities. All three nutrient‐limited environments resulted in decreased size of egg‐bearing (meristic) gametophytes compared to nonlimited environments, but gametophytes failed to respond to N and CO₂ limitation at the time of sex determination, resulting in no ESD. N limitation resulted in a predictable allometric re‐allocation of resources based on small gametophyte size, whereas CO₂ limitation caused a change in reproductive output consistent with true plasticity. Withholding exogenous P caused ESD and had no effect on relative reproductive output of resultant meristic gametophytes because the size decrease was minor. Under P limitation, ESD matched the resource demands of gender phenotypes to their environment before the onset of developmental dimorphism, reducing the need for large allocation adjustments after sex determination.     

How reproductive allocation and flowering probability of individuals in plant populations are affected by position in stand size hierarchy,plant size and CO2 regime

Aims We investigate the effect of position within a size-structured population on the reproductive allocation (RA) and flowering probability of individual plants of Sinapis arvensis. We also assess the effects of plant size and changing level of CO₂ on both responses.Methods Sinapis arvensis L., (field mustard), an annual agricultural weed, was grown in monoculture at six densities under ambient and elevated CO₂ in a study with 84 stands. Individual aboveground biomass and reproductive biomass were measured. Varying density produced a wide range of mean plant sizes across stands and size hierarchies within stands. Many (~40%) individuals had zero reproductive biomass. Employing a novel modelling approach, we analysed the joint effects of position in stand size hierarchy, plant size and CO₂ on RA and flowering probability of individuals.Important findings We found a strong effect of position within the size hierarchy of individuals in a population: for an individual of a given size, greater size relative to neighbours substantially increased RA and flowering probability at a single harvest time. There was no other effect of plant size on RA. We found a positive effect of elevated CO₂ on RA regardless of position within the size hierarchy. These observed patterns could impact doubly on the reproductive biomass (R) of small individuals. First, because RA is not affected by size, smaller plants will have smaller R than larger plants; and second, for smaller plants lower down in a population size hierarchy, their RA and hence R will be further reduced. These results suggest that size relative to neighbours may be independent of and more important than direct abiotic effects in determining RA. Further studies are required to evaluate how these observed patterns generalize to other populations in non-experimental conditions.     

青藏高原特有植物露蕊乌头 (毛茛科) 从冰期避难所扩张后繁殖资源分配的变化

物种分布范围的形成是进化生态学研究的基本问题之一,但植物的资源分配策略是否与物种边界形成有关一直没有相关研究。青藏高原特有植物露蕊乌头在末次最大冰期时有4个避难所,但冰期后只有一个避难所的种群发生了扩张并最终形成了现代分布格局。以露蕊乌头的避难所种群（同仁种群）和扩张后邻近分布区边缘的两个种群（兴海种群和海北种群）为研究对象,通过比较避难所种群和边缘种群的资源分配方式,探讨露蕊乌头的资源分配与该植物分布区及边界形成的关系。结果发现：1）兴海和海北种群的营养结构（包括根、植株高度和茎叶生物量）均显著低于同仁种群,海北种群的繁殖结构（花数量和花生物量）显著低于同仁和兴海种群,但海北和兴海的繁殖分配均显著高于同仁种群;2）3个种群的繁殖资源与个体大小呈现显著的正相关关系,投入到繁殖资源的比例（繁殖分配）与个体大小呈显著的负相关关系。对露蕊乌头的研究结果一方面进一步证明了个体大小依赖的繁殖分配,但不符合“植物开始繁殖必须达到一定的大小（阈值）”这一结论,这可能与露蕊乌头的生活史特征有关;而另一方面,露蕊乌头在扩张过程中逐渐增加了对繁殖资源投资的比例,说明胁迫生境中有性繁殖对该植物具有更为重要的意义,且露蕊乌头在扩张过程中可能逐渐实现繁殖产出最大化,并可能在边缘种群实现最优繁殖分配进而最终形成该物种分布区的边界,但这一结论仍需在更多的植物类群中验证。     

Changes in Reproductive Allocation after Expansion from the Glacial Refugium and Implications for the Distribution Range in the Qinghai Tibet Plateau Native Herb,Gymnaconitum gymnandrum (Ranunculaceae)

A fundamental goal of ecology and evolution is to explain patterns of species distribution and abundance. However, the way in which stable distribution ranges are shaped by natural selection is still poorly understood, especially whether patterns of resource allocation have contributed to the range size and the formation of range boundary received little attention. For annual herb, the maximum reproductive allocation is predicted to be 50%, and thus we predicted that reproductive allocation might contribute to the formation of range boundary since plant will enhance allocations to reproduction in stressful environments. In this study, we presented our data on resource allocation between population from the glacial refegium and those from the marginal populations in Gymnaconitum gymnandrum, an alpine biennial native to the Qinghai Tibet Plateau, aiming to find the contribution of resource allocation to the formation of range boundary. Our results showed that resource allocations to vegetative organs, including roots, plant height and stem leaf biomass, were significantly higher in the refugium population that in the two marginal populations, and allocations to reproductive organs, including flower number and flower biomass, were significantly lower in one marginal population (Haibei population) than in the other marginal population (Xinghai population) and the refugium population (Tongren population). However, reproductive allocation was significantly higher in the marginal populations than in the refugium population. In addition, in each of the three populations, we found a positive relationship between the plant size and flower biomass but a negative relationship between the plant size and reproductive allocation. Our results indicated a size dependent reproductive allocation in Ggymnandrum, but we did not find a size threshold for reproduction in each of the three populations of this plant, which might be attributed to the life history of this biennial herb. We also suggested that reproductive allocation was increased during the process of range expansion and may rise to the optimal reproductive allocation in the marginal populations, which suggested the important role of sexual reproduction for plants in more stressful environments and the formation of range boundary. However, these conclusions need to be further proved in other plant species.     

古尔班通古特沙漠三种生境下梭梭种群的生殖分配特征

对古尔班通古特沙漠梭梭种群进行样地调查,从各构件生物量分配特征、开花和结果特征、不同发育阶段的生殖分配、同化枝与生殖生物量之间关系等方面研究了3种生境下梭梭种群的生殖分配特征。结果表明:3种生境中各构件生物量分配格局均具有显著差异,且变化趋势不一致,梭梭平均生殖分配在0.64%～1.5%;3种生境下,梭梭开花和结果特征均具有显著差异,表现出砾石生境>沙质生境>土质生境。不同生境梭梭生殖分配均随径级的增加变化不同,砾石生境和沙质生境样地随着径级的增加而降低,而土质生境样地随着径级的增加而增加;不同生境类型下梭梭同化枝、生殖生物量与径级的关系不同;梭梭的生殖分配和同化枝生物量之间呈线性或多项式正相关。     

Between-population variation in size-dependent reproduction and reproductive allocation in Pinguiculavulgaris (Lentibulariaceae) and its environmental correlates

Several important fitness components in herbaceous perennial plants are commonly related to plant size: flowering probability, reproductive allocation and fecundity. However, evidence for such size-dependence of fitness components is mostly anecdotal and unconnected to other life history traits. Here we report size-dependence for flowering probability and reproductive allocation in 11 populations of Pinguicula vulgaris and relate it to environmental factors. Flowering probability was size-dependent in all populations of P. vulgaris , and indicated the existence of a threshold size for reproduction. Populations at low altitudes and in wet soils showed a significantly higher threshold size for reproduction. Reproductive mass was also size-dependent in all populations. We found considerable between-population differences in the slope and the intercept of the regression between plant size and reproductive mass. This variation was weakly related to the environmental factors measured. In general, relationships between different size-dependent fitness components were low. Instead of showing a covariation of traits, in line with interpretations in terms of life history "tactics", P. vulgaris seemed to independently vary each size-dependent fitness component in each locality. In particular, no significant relationship was found between threshold size for reproduction and the slope of size-dependent reproductive allocation, as predicted by previous authors. Neither we found a significant influence of somatic cost of reproduction on size-dependent fitness components.     

Sexual differences in biomass and nutrient allocation of first-year Silene dioica plants

Reproductive and somatic biomass, nitrogen (N), and phosphorus (P) pools were compared between females and males in 1st-year plants of Silene dioica. We estimated irretrievable resources allocated to seeds, pollen, flowers, and unrecovered summer leaf investment by collecting plant parts at abscission throughout the season. At the end of the season, we determined resources lost through senescent stems and autumn leaf turnover and resources stored in perennial roots and overwintering buds. Sexual differences in allocation patterns depended on the resource used for comparison, and whether absolute or proportional resource pools were assessed. Total resource pools in terms of biomass and N were similar for females and males. However, male plants acquired relatively more P. The proportional reproductive investment, i.e., reproductive effort, was similar for males and females in terms of biomass and N. In terms of P, male reproductive effort was higher. There was no difference between sexes in the proportional and relative biomass allocated to perennial roots and overwintering buds. However, in terms of absolute and relative N allocation to below-ground parts, females had larger reserves than males. Females, moreover, had a larger proportion of their P in below-ground parts. However, as male total P pools were larger, absolute P reserves did not differ between sexes. The high reproductive effort and N depletion of below-ground parts in males resulted largely from higher flower production compared to females. In females, seeds were the major component of reproductive effort. These results show that if biomass and nutrient allocation are assessed in parallel for dioecious plants, we obtain a more complete view of their sexual differences. Received: 07 May 1998 / Accepted: 30 October 1998     

Reproductive allocation strategies in desert and Mediterranean populations of annual plants grown with and without water stress

Reproductive effort (relative allocation of biomass to diaspore production) was compared in matched pairs of Mediterranean and desert populations of three unrelated annual species, Erucaria hispanica (L.) Druce, Bromus fasciculatus C. Presl. and Brachypodium distachyon (L.) Beauv., grown under high and low levels of water availability in a common-environment experiment. Desert populations in all three species showed higher reproductive effort than corresponding Mediterranean populations, as expressed by both a reproductive index (RI= reproductive biomass/vegetative biomass), and a reproductive efficiency index (REI=number of diaspores/total plant biomass). Moreover, in E. hispanica and Brachypodium distachyon, inter-populational differences in reproductive effort were greater under water stress, the main limiting factor for plant growth in the desert. These results indicate that variability in reproductive effort in response to drought is a critical and dynamic component of life history strategies in annual species in heterogeneous, unpredictable xeric environments. When subjected to water stress the Mediterranean populations of E. hispanica and B. distachyon showed greater plasticity (e.g. had a greater reduction) in reproductive effort than the desert populations, while in Bromus fasciculatus both populations showed similar amounts of plasticity.     

Sex allocation of a cosexual wind-pollinated tree, Quercus dentata, in terms of four currencies

Sex allocation of a cosexual wind-pollinated species, Quercus dentata (Fagaceae), was analyzed using biomass, carbon, nitrogen and phosphorus as currencies based on data accumulated for 61 individuals from 1997–2004. Strongly female-biased sex allocation was indicated when measured in terms of biomass and carbon, but no significant bias was detected when measured in terms of nitrogen or phosphorus. From an adaptive viewpoint, there is little support for strong female-biased sex allocation, suggesting that sex allocation in terms of nitrogen or phosphorus is closer to the real picture. The relative sex allocation considerably varied from year to year, but the relative femaleness of individuals in the population was rather constant across years. No significant correlation was observed between relative sex allocation and fecundity or tree height, but individuals that showed very low fecundity tended to produce only acorns.     

青藏高原东缘星状风毛菊生殖分配对海拔的响应

通过采样调查法和烘干称量法, 检测了青藏高原东缘不同海拔20个居群星状风毛菊(Saussurea stella)的花部特征, 研究其花部特征对海拔及个体大小的响应。结果表明: 1)繁殖分配是依赖个体大小的, 个体越大, 繁殖分配越小; 2)花期繁殖器官和营养器官的生物量、头状花序的数量及大小、雌雄功能以及雄性内部花丝与花粉之间均存在资源分配上的权衡; 3)星状风毛菊通过花部特征的变异响应海拔的变化。研究发现, 星状风毛菊通过花部特征的变异, 特别是花柱和花丝的伸长及投入更多资源于吸引结构花瓣, 以此来补偿该种在高海拔地区因传粉者减少、活动频率降低和花粉数减少等因素对传粉带来的影响, 使其能够在高海拔的胁迫环境下有效地传粉, 完成有性繁殖。     

Male reproductive allocation in fireflies (Photinus spp.)

Abstract. Resource allocation between somatic and reproductive structures has important fitness consequences for individuals, and optimal trade-offs are expected to depend not only on mating system differences among species but also on levels of resource stress within species. We tested the prediction that polyandry (associated with increased sperm competition) will increase male reproductive allocation in bioluminescent fireflies in Photinus spp. by comparing the relative mass of testes, seminal vesicles, and reproductive accessory glands among a monandrous and several polyandrous species. In addition, we examined a single population of a polyandrous species, Photinus greeni , to see how reproductive allocation might shift between years in response to different levels of larval resource stress. As predicted, males of P. collustrans , a monandrous species, showed a fivefold lower allocation to sperm production and a 100-fold lower allocation to reproductive accessory glands compared with males from polyandrous species. We also found evidence within P. greeni of a trade-off between allocation either to reproduction or to somatic tissue; following larval resource stress, males eclosed at significantly shorter body lengths, yet showed a 35% increase in their reproductive allocation. These results demonstrate that mating systems strongly influence male allocation to reproductive accessory glands as well as to sperm production. Furthermore, these results suggest that under larval resource stress males of Photinus spp. increase their allocation to reproduction at the expense of somatic tissue, thus maximizing their ability to produce nuptial gifts required for reproductive success.