Regulatory Mechanisms of Reproductive Effort in Plants 1. Plasticity in Reproductive Energy Allocation and Propagule Output of Helianthus annuus L. (Compositae) Cultivated at Varying Densities and Nitrogen Levels

Abstract Plasticity in growth, reproductive energy allocation (RA), and reproductive output were investigated in Helianthus annuus L. cultivar. Russia (Compositae) grown under varying densities and soil nitrogen levels. The role and behaviour of pollinators in seed production was also examined.
Exceedingly marked plastic responses were detected in individual biomass, the patterns of resource allocation to total reproductive structures (RA) and also to propagules, fecundity, reproductive outputs, and propagule size and weight under changing densities and soil nitrogen levels. Plants cultivated at higher densities exhibited proportionately lower individual biomass, lower RA, lower fecundity, lower seed output, and smaller seed size in response to increasing density and decreasing soil nitrogen levels. However, differences due to different N-levels were not as great as those to changing density.
One of the most significant findings was that seed production under limited resource availability, i.e., lack of ample solar radiation and soil nutrients, due to strong interference at higher density plots, is exceedingly costlty , This was most clearly demonstrated by a very sharp increase in relative energy partitioning to a single propagule in response to the increased density and decreasing nitrogen levels, the relative energy cost to a single achcne (R_A) increasing from one to twenty-fourfold. Reproduction was also affected by pollinator-limitations, and seed size showed a marked position effect.     

Reproductive allocation patterns in different density populations of spring wheat

The effects of increased intraspecific competition on size hierarchies （size inequality） and reproductive allocation were investigated in populations of the annual plant, spring wheat （Triticurn aestivurn）. A series of densities （100, 300, 1 000, 3 000 and 10 000 plants/m^2） along a gradient of competition intensity were designed in this experiment. The results showed that average shoot biomass decreased with increased density. Reproductive allocation was negatively correlated to Gini coefficient （R^2 = 0.927）, which suggested that reproductive allocation is inclined to decrease as size inequality increases. These results suggest that both vegetative and reproductive structures were significantly affected by intensive competition. However, results also indicated that there were different relationships between plant size and reproductive allocation pattern in different densities. In the lowest density population, lacking competition （100 plants/m^2）, individual reproductive allocation was size independent but, in high density populations （300, 1 000, 3 000 and 10 000 plants/m^2）, where competition occurred, individual reproductive allocation was size dependent： the small proportion of larger individuals were winners in competition and got higher reproductive allocation （lower marginal reproductive allocation; MRA）, and the larger proportion of smaller individuals were suppressed and got lower reproductive allocation （higher MRA）. In conclusion, our results support the prediction that elevated intraspecific competition would result in higher levels of size inequality and decreased reproductive allocation （with a negative relationship between them）. However, deeper analysis indicated that these frequency- and size-dependent reproductive strategies were not evolutionarily stable strategies.     

天津沿海滩涂互花米草种群生殖分株数量特征及生殖分配研究

通过对天津沿海滩涂单优群落的取样调查,研究了互花米草种群生殖分株的数量特征和生殖分配规律。结果表明,在籽实成熟期,互花米草种群单个生殖分株重为15.62±9.26 g,穗和种子重分别为2.68±2.08和1.39±1.12 g,小穗和种子数分别为537.7±362.2和490.2±376.3个,生殖分配Ⅰ和生殖分配Ⅱ分别为15.02±5.83和7.62±3.8%,结实率为81.9 ±28.6%。穗和种子的形成分别需要分株积累3.2和3.6 g以上的生物量。穗长、穗重、种子数、小穗数、生殖分配Ⅰ和生殖分配Ⅱ分别与分株高度呈极显著(p<0.01)的正相关关系。穗重、小穗数、种子重、种子数、生殖分配Ⅰ和生殖分配Ⅱ分别与分株重呈极显著(p<0.01)的线性正相关关系。种子数、小穗数、生殖分配Ⅰ和生殖分配Ⅱ分别与茎和叶鞘生物量分配呈极显著(p<0.01)的负相关关系。穗各构件在空间上的分布也存在较强的规律性。     

Reproductive Biology of an Amphicarpic Annual, Polygonum thunbergii (Polygonaceae): Spatio-temporal Changes in Growth, Structure and Reproductive Components of a Population over an Environmental Gradient

Abstract The process and mechanisms of spatio-temporal changes in growth, population structure, as well as various yield and reproductive components of a population of an amphicarpic annual, Polygonum thunbergii (Polygonaceae), with two reproductive systems (aerial chasmogamous flowers and subterranean cleistogamous flowers) along an environmental gradient (light, moisture and soil nitrogen levels) were investigated in the field.
The results clearly demonstrate that growth and allocation patterns, population structure, and reproductive output of individuals changed sharply along the environmental gradient in response to seasonal and spatial changes in resource availability. Models predicted that light conditions bring about one-sided competition, whereas nutrient conditions in the soil engender two-sided competition. As expected, the degree of one-sided competition was prominent in the case of a planophile, Polygonum thunbergii. Allocation patterns, seed outputs, individual seed size as well as relative energy costs of chasmogamous and cleistogamous seed as affected by light and nutrient levels were also critically analyzed. The most noteworthy finding was that the size of chasmogamous seeds sharply decreased in response to a decrease in the light regime, while cleistogamous seed size remained constant along the gradient. However, relative cost of both chasmogamous and cleistogamous seeds sharply increased with decrease in the light level, reflecting different degrees of environmental stress, biotic interference, or both.     

Optimal Reproductive Strategy of Plants, with Special Reference to the Modes of Reproductive Resource Allocation

Abstract A growth model for reproductive energy allocation pattern and schedule is proposed. Assumptions are as follows: (1) the assimilation rate for an individual is given by a logistic curve of vegetative dry weight; (2) size variability is expressed by the parameter W of the logistic curve (asymptotic value of vegetative dry weight); (3) a plant controls allocation of the assimilate to vegetative and reproductive structures so as to maximize the reproductive energy investment at the end of the growth period. The models were analyzed in comparison with field and experimental observations and gave reasonable explanations for the reproductive allocation pattern of individuals which reflects ecological preferences and life history characteristics, such as environmental conditions of habitats (stable or changing), length of life span (annual, biennial or perennial) and growth form (erectophile or planophile). Decreasing RA (reproductive allocation) with individual size and delayed switchover time from vegetative to reproductive growth were found in plants which occur in stable environments and have a more or less fixed growth period; in those which occur in changing environments where growth period depends on individual size, RAs that remain constant or increase with variations in individual size and early switchover time were detected. Most perennials conform to the former case, but annuals and biennials conform to the latter case. Under extremely overcrowded conditions, planophiles, which are much more subject to crowding effect than erectophiles, tend to have increasing RA with increasing size, while erectophiles tend to have almost constant RA irrespective of size. These trends are discussed in the light of the life history characteristics and ecological distribution of plant species studied.     

Reproductive allocation of biomass and nitrogen in annual and perennial Lesquerella crops

* BACKGROUND AND AIMS: The use of perennial crops could contribute to increase agricultural sustainability. However, almost all of the major grain crops are herbaceous annuals and opportunities to replace them with more long-lived perennials have been poorly explored. This follows the presumption that the perennial life cycle is associated with a lower potential yield, due to a reduced allocation of biomass to grains. The hypothesis was tested that allocation to perpetuation organs in the perennial L. mendocina would not be directly related to a lower allocation to seeds. * METHODS: Two field experiments were carried on with the annual Lesquerella fendleri and the iteroparous perennial L. mendocina, two promising oil-seed crops for low-productivity environments, subjected to different water and nitrogen availability. * KEY RESULTS: Seed biomass allocation was similar for both species, and unresponsive to water and nitrogen availability. Greater root and vegetative shoot allocation in the perennial was counterbalanced by a lower allocation to other reproductive structures compared with the annual Lesquerella. Allometric relationships revealed that allocation differences between the annual and the perennial increased linearly with plant size. The general allocation patterns for nitrogen did not differ from those of biomass. However, nitrogen concentrations were higher in the vegetative shoot and root of L. mendocina than of L. fendleri but remained stable in seeds of both species. * CONCLUSIONS: It is concluded that vegetative organs are more hierarchically important sinks in L. mendocina than in the annual L. fendleri, but without disadvantages in seed hierarchy.     

Linking flowering and reproductive allocation in response to nitrogen addition in an alpine meadow

Aims Plants can change in phenology and biomass allocation in response to environmental change. It has been demonstrated that nitrogen is the most limiting resource for plants in many terrestrial ecosystems. Previous studies have usually focused on either flowering phenology or biomass allocation of plants in response to nitrogen addition; however, attempts to link flowering phenology and biomass allocation are still rare. In this study, we tested the effects of nitrogen addition on both flowering phenology and reproductive allocation in 34 common species. We also examined the potential linkage between flowering time and reproductive allocation in response to nitrogen addition.Methods We conducted a 3-year nitrogen addition experiment in Tibetan alpine meadow. We measured first flowering date and the reproductive allocation for 34 common plant species in control, low and high nitrogen added plots, respectively. One-way analysis of variance was used to examine differences of first flowering date and reproductive allocation among treatments. The relationships between the change in species first flowering date and change in reproductive allocation in response to nitrogen addition were examined by calculating Pearson correlation coefficients.Important findings For most species, both first flowering date and reproductive allocation significantly responded to nitrogen addition. Nitrogen addition significantly delayed the first flowering date and reduced the reproductive allocation for all graminoid species, but accelerated flowering and increased reproductive allocation for most forb species. We found that changes in first flowering date significantly negatively correlated with the changes in reproductive allocation over species in response to nitrogen, which indicated a positive relationship between flowering response and plant performance in reproductive allocation. Species that advanced their flowering time with nitrogen addition increased their reproductive allocation, whereas those that delayed flowering time tended to decline in reproductive allocation with nitrogen addition. Our results suggest that species-specific switch from vegetative growth to reproductive growth could influence species performance.     

谷子的亲缘识别能力及其与种植密度和土壤养分水平的关系

为探明作物是否具有识别邻株身份的能力以及这种能力是否受到环境因子的调控,通过大田试验,研究邻株身份(亲缘株、非亲缘株和陌生株)、种植密度和土壤养分水平的交互效应对谷子(Setaria italica)地上部分生物量分配的影响。结果表明,谷子与亲缘株为邻时的净繁殖生物量分配和种子生物量分配,比与非亲缘株为邻时显著提高,且营养生物量分配显著降低(P<0.05)。在高种植密度条件下,亲缘组谷子的穗长、净繁殖生物量分配和种子生物量分配显著大于非亲缘组,而营养生物量分配显著小于非亲缘组(P<0.05)。随着土壤养分水平提高,亲缘组谷子的种子生物量分配显著增加,营养生物量分配显著减少(P<0.05)。由此推断,谷子具有对亲缘邻株的识别能力,且这种能力受种植密度和土壤养分水平的调控,在高种植密度和高土壤养分水平条件下,谷子的亲缘邻株识别能力较强。     

Life History Studies on the Genus Trillium (Liliaceae) I. Reproductive Biology of Four Japanese Species

Abstract Four Japanese Trillium (Liliaceae) species which are representative perennial herbs of the temperate deciduous forests— Trillium kamtschaticum (2x), T. apetalon (4x), T. tschonoskii (4x) and T. smallii (6x)—were studied for their reproductive characteristics, e.g., patterns of reproductive resource allocation and reproductive output.
In spite of the differences in ploidy levels (from 2x to 6x), all four species showed very similar reproductive traits. It became evident that in response to the increase in reproductive allocation to total reproductive organs (RA), the number of seeds produced per plant (P_N) clearly increased. This trend is well in accord with the relationship found in four North American species and also several temperate woodland perennial herbs which occur in closed, stable and predictable environments, and possess typical xenogamous breeding systems. Although there is no conspicuous trend between ploidy levels and P_N, one of the significant differences noted in this study was in seed weight, which ranged from 2.93 mg in diploids, to 3.42–3.45 mg in tetraploids, and to 4.47 mg in hexaploids.     

Reproductive efficiency and growth of Emex australis in relation to stress

The reaction of Emex australis to different degrees of stress was examined in a pot experiment by varying pot size and in a field experiment by varying density. In pots, the proportion of energy in seeds was more stable to the imposition of stress than in roots and stems; that in roots increasing with increasing stress and that in stems declining. Similarly in the field, there was little difference between densities of 1–32 plants m^?2 in the proportion of seeds in biomass. There were no significant differences in mean individual seed weights in either experiment and all plants set seed. Again in both experiments, high stress plants were more precocious than low stress ones. In pots, this took the form of earlier attainment of maxima in calorific values, ‘reproductive effort’, root, stem and seed weights and in the field, a shorter life cycle and earlier senescence. In pots, estimates of the overall mean reproductive effort were 62% for the low and medium stress plants and 36% for the high stress plants. The results are discussed in terms of the high values for reproductive effort, the plasticity and precocity of E. australis under stress and the implications for weed control.     

SEED AND VEGETATIVE REPRODUCTION IN RELATION TO DENSITY IN FRAGARIA VIRGINIANA (ROSACEAE)

Transplants of Fragaria virginiana Duchesne, the wild strawberry, from field and woods sites, were studied under controlled laboratory conditions to determine their seed and vegetative reproduction responses under different density conditions. The following results were obtained: (1) Increased competition among high density grown plants resulted in lower total biomass when compared to low density grown plants; (2) Increased shading in high density plots resulted in a greater percentage of biomass in leaves; (3) The percentage of total biomass in reproductive organs (seed and vegetative) was higher in low density grown plants; (4) Vegetative “reproductive effort” was higher in low density plots than in high density plots while seed “reproductive effort” remained constant between the two density treatments; (5) No significant difference between plants from habitats of different successional maturity was observed under the experimental treatments. These results are discussed in relation to reproductive density response models.     

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...     

Effects of resource availability and propagule supply on native species recruitment in sagebrush ecosystems invaded by Bromus tectorum

Resource availability and propagule supply are major factors influencing establishment and persistence of both native and invasive species. Increased soil nitrogen (N) availability and high propagule inputs contribute to the ability of annual invasive grasses to dominate disturbed ecosystems. Nitrogen reduction through carbon (C) additions can potentially immobilize soil N and reduce the competitiveness of annual invasive grasses. Native perennial species are more tolerant of resource limiting conditions and may benefit if N reduction decreases the competitive advantage of annual invaders and if sufficient propagules are available for their establishment. Bromus tectorum, an exotic annual grass in the sagebrush steppe of western North America, is rapidly displacing native plant species and causing widespread changes in ecosystem processes. We tested whether nitrogen reduction would negatively affect B. tectorum while creating an opportunity for establishment of native perennial species. A C source, sucrose, was added to the soil, and then plots were seeded with different densities of both B. tectorum (0, 150, 300, 600, and 1,200 viable seeds m⁻²) and native species (0, 150, 300, and 600 viable seeds m⁻²). Adding sucrose had short-term (1 year) negative effects on available nitrogen and B. tectorum density, biomass and seed numbers, but did not increase establishment of native species. Increasing propagule availability increased both B. tectorum and native species establishment. Effects of B. tectorum on native species were density dependent and native establishment increased as B. tectorum propagule availability decreased. Survival of native seedlings was low indicating that recruitment is governed by the seedling stage.     

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。     

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。     

Allometry of Salsola collina in response to soil nutrients,water supply and population density

The allometry of greenhouse‐grown Salsola collina Pall. in response to variation in soil nutrient content, water supply and population density has been compared. The results showed that the biomass allocation was size‐dependent. Root, stem, leaf and reproductive allocation showed a ‘true’ plasticity in response to soil nutrient variation. At low soil nutrient content, plants tended to allocate more biomass to the development of reproductive organs than to stem and leaf, but root allocation was consistent due to a tradeoff between the effects of plant size and soil nutrient content. The plasticity of stem allocation and reproductive effort was ‘true’, while the plasticity of root allocation was ‘apparent’, but there was no plasticity for leaf allocation in response to soil water variation. At lower soil water content, plants tended to allocate more biomass to the stem than to development of reproductive organs. With the exception of ‘apparent’ plasticity of root allocation, no plasticity was detected in biomass allocation when population density was varied.     

The interplay between shifts in biomass allocation and costs of reproduction in four grassland perennials under simulated successional change

When perennial herbs face the risk of being outcompeted in the course of succession, they are hypothesized to either increase their biomass allocation to flowers and seeds or to invest more in vegetative growth. We tested these hypotheses in a 3-year garden experiment with four perennials (Hypochaeris radicata, Cirsium dissectum, Succisa pratensis and Centaurea jacea) by growing them in the midst of a tall tussock-forming grass (Molinia caerulea) that may successionally replace them in their natural habitat. In all species except for the short-lived H. radicata, costs of sexual reproduction were significant over the 3 years, since continuous bud removal enhanced total biomass or rosette number. To mimic succession we added nutrients, which resulted in a tripled grass biomass and higher death rates in the shorter-lived species. The simulated succession resulted also in a number of coupled growth responses in the survivors: enhanced plant size as well as elevated seed production. The latter was partly due to larger plant sizes, but mostly due to higher reproductive allocation, which in turn could be partly explained by lower relative somatic costs and by lower root–shoot ratios in the high-nutrient plots. Our results suggest that perennial plants can increase both their persistence and their colonization ability by simultaneously increasing their vegetative size and reproductive allocation in response to enhanced competition and nutrient influxes. These responses can be very important for the survival of a species in a metapopulation context. Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

狮牙草状风毛菊果期资源分配对海拔的响应

植物资源分配是目前植物生态学研究的热点问题, 主要集中在性分配和繁殖分配两个方面。该研究以分布在青藏高原的狮牙草状风毛菊(Saussurea leontodontoides)作为研究材料, 研究了6个海拔高度上果期植株的繁殖特征及资源分配的差异, 并用异速模型分析了繁殖性状及资源分配与个体大小的关系。结果显示: 1)狮牙草状风毛菊的个体大小、繁殖器官生物量、营养器官生物量、种子数和营养分配均与海拔存在极显著负相关关系, 百粒质量和繁殖分配与海拔存在极显著正相关关系。2)在不同海拔高度下, 百粒质量、种子数、繁殖器官生物量及营养分配与植株个体大小呈正相关关系; 繁殖分配与植株个体大小呈负相关关系; 而营养器官生物量与植株个体大小呈极显著正相关关系。这表明海拔和个体大小对狮牙草状风毛菊的繁殖对策有不同程度的影响, 狮牙草状风毛菊通过增加繁殖部分的生物量和百粒质量来适应高海拔的胁迫环境。     

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.     

放牧影响下羊草种群生殖生态学的研究

在种群水平上系统研究了放牧对松嫩草原关草种群无性繁殖、有性生殖和生物量的生殖分配的影响。结果表明,放牧制约了羊草种群无性繁殖和有性生殖的更新途径,表现在随放牧强度增加,无性繁殖体营养技密度、根茎牙密度和有性生殖体种子生物量、结实数和生殖枝分化率等指标均显著下降,是生牧后下降迅速。随放牧强度增加,关草种群根茎生物量的分配比例显著增加,而同化器官、非同化器官和种子生物量分配比例都显著下降,极牧阶段几科