松嫩平原异质生境羊草种群营养繁殖特征

多年生无性系禾草主要依靠营养繁殖实现种群的持续更新,而各类芽和由芽形成的苗为其种群进行营养繁殖的潜在种群。通过单位面积挖掘取样,利用营养繁殖世代数划分分株、根茎和各类型芽和苗的龄级,对松嫩平原封育草甸、长期割草草甸、封育积沙草甸和林间草地的羊草潜在种群组成和大小进行了比较研究。结果表明: 在生长季末期,羊草的潜在种群均由根茎芽和各龄级分蘖节向上生长的芽以及根茎苗和分蘖节苗组成。在4种生境条件下,羊草分株潜在种群由3～4个龄级组成,1～4龄级根茎均可形成潜在种群,其中,根茎潜在种群占整个羊草潜在种群数量比例为68.3%,占绝对优势。1龄分株和根茎产生的潜在种群占总体的60.2%,是潜在种群组成的主要部分,是种群更新的主要来源。分株潜在种群数量以长期割草草甸最高,根茎潜在种群数量以封育积沙草甸最高。营养繁殖力均以年轻龄级分株和根茎最高,根茎的营养繁殖力是分株的10.0倍,在羊草种群营养繁殖过程中始终占据优势。羊草的潜在种群构成在不同生境条件下变异较大,但体现出对生境差异的趋异适应。     

松嫩平原两个生态型羊草种群生长机制

对松嫩平原两个生态型羊草种群生长机制进行了研究.结果表明,两个生态型羊草种群分株重与密度之间关系呈幂函数变化,方程中b值（-0.098～-0.272）大于-1.两个生态型羊草种群生物量、分株重随着密度、高度的增加而逐渐增加,它们之间的关系符合幂函数或直线变化规律,达到显著或极显著相关水平.两个生态型羊草种群生物量和分株重随密度和高度的变化速率在营养生长期内均最小.灰绿生态型的变化速率b（0.7872、0.0134）在营养生长期小于黄绿生态型（0.8793、0.0222）;到果后营养期,灰绿生态型（0.8048、0.0303）逐渐超过黄绿生态型（0.7796、0.0258）.两个生态型羊草种群在研究样地环境中未达到环境容纳量,两个生态型羊草种群在整个生育期内不断生长,具有相同的生长规律,且灰绿生态型羊草种群具有更强的生长潜力.     

光强梯度对羊草无性系分化与生长的影响

在温室内用透光率分别为 60 %、45%、30 %的遮阴网盖在花盆上遮阴 ,研究不同光照强度对羊草无性系分化与生长的影响 .结果表明 ,低光强影响羊草基株的生物量积累 ,抑制羊草无性系的分化和生长 .减少羊草地下根茎的数目 ( A=61 4个 ,D=2 31个 )和无性系分株数 ( A=70 4个 ,D=1 52个 ) .羊草根茎长度和节数、无性系分株干重等性状在不同的光强梯度之间存在显著差异 ( P<0 .0 1 ) ,这种差别在春季尤其显著 .     

松嫩平原两个趋异类型羊草无性系种群特征的比较研究

松嫩平原上羊草(Leymus chinensis(Tzvel.)Tzvel.)有两个趋异类型:灰绿型和黄绿型。两个类型羊草的分蘖节一般均存活2～4年,最多可存活5年;根茎一般存活2～3年,最多可存活4年。两个类型无性系种群的分蘖株均为增长型的年龄结构类型。种群根茎的累积长度,灰绿型为18035cm/m~2,黄绿型为21218cm/m~2,其中,均以1、2龄占绝对比重。两个类型均以1龄分蘖株生产力最大,至3龄分蘖株明显减小;各龄根茎的生物量随着年龄的增加呈直线下降;1龄根茎的营养繁殖力甚强,至8月中旬所形成的芽数均已远远多于地上全部分蘖株数;2龄根茎尚存在较小的营养繁殖潜力,3、4龄根茎均已丧失了营养繁殖力。两个类型羊草无性系种群都是通过根茎芽补充更新。     

松嫩平原栽培条件下羊草无性系构件的结构

羊草是长根茎型禾草,是典型的无性系植物,在松嫩平原的生长季末期,栽培条件下羊草无性系分株由分蘖株和分蘖苗组成,在具有充分生长空间而又没有种间竞争的风沙土上,羊草分株的分蘖节在一个生长季内可以繁殖4个世代,按分蘖节的繁殖世代划分龄级,现实与潜在无性系构件的年龄谱均以1龄级比重最大,随着龄级的增加明显减少,呈增长型的年龄结构,羊草无性系分株的生产力主要与分株形成及生长的时间长短有关,形成时间越早、生长时间越长的分蘖株对无性系的物质生产和营养繁殖的贡献越大,羊草无性系在空间扩展与物质贮存上具有一定的可调节性。     

内蒙古锡林河流域不同生境中羊草的克隆构型和分株种群特征

 克隆植物的形态可塑性在基株和种群水平上分别表现为克隆构型和分株种群特征的变化。研究对象为内蒙古锡林河流域草地、林地、沙地3种生境下的羊草(Leymus chinensis)种群,通过对羊草根茎节间长度、间隔子长度、分枝强度、分枝角度、株高和分株密度等指标的测定和分析,对这3种不同生境中羊草的克隆构型及分株种群特征进行了研究。结果表明羊草克隆构型相关特征,如,根茎节间长度,根茎节间长度频次分布格局、间隔子长度、间隔子长度频次分布格局,在不同生境差异较大。同时,羊草的分枝角度在不同生境间差异显著。而每间隔子所     

水分胁迫对羊草光合产物分配及其气体交换特征的影响

 通过对典型草原优势植物种羊草(Leymus chinensis)的盆栽实验,模拟5个土壤水分梯度（分别为土壤持水量的75%～80%(对照)、60%～65%、50%～55%、35%～40%和25%～30% ）对羊草叶片相对含水量、光合速率、光合产物分配和种群CO2交换速率的影响。结果表明：随着土壤水分胁迫的增加,羊草叶片相对含水量呈先增加而后下降的单峰型变化,且在50%～55%处理下达到最大;叶片光合速率随着水分胁迫的增加而减小,且75%～80%、60%～65%、50%～55%的水分处理与35%～40%、25%～30%的水分处理的叶片光合速度日动态规律不同。羊草总生物量及根、鞘、叶生物量均随着水分胁迫的增加呈下降趋势。干旱促进早期羊草根的分配和根冠比增加, 但到后期却使它们降低, 表明羊草在受到较长期的持续干旱后通过增加根部的比重来提高抗旱性的能力逐渐降低。羊草根茎的生物量和分配随着土壤水分含量降低均呈现出先增加而后下降的趋势,羊草根茎的生物量在50%～55%处理下达最大（1.28 g·株-1）,而羊草根茎的分配在35%～40%处理下达最大（48.5%）。羊草种群CO2的净交换速率随着水分胁迫的增加而减小,其日交换量随着水分胁迫的增加而增加,且在60%～65%处理下达到最高,而后呈下降趋势,并在25%～30% 处理下为负值。研究结果表明,土壤持水量的40%可能是羊草对于水分变化响应的阈值。     

季节放牧下内蒙古温带草原羊草根茎叶功能性状的权衡

放牧是草地主要利用方式之一,不同季节放牧通过影响草地功能性状间的权衡从而影响牧后再生及补偿性生长。通过测定内蒙古温带草原优势种羊草(Leymus chinensis)的株高、节间距和分蘖数等软性状及气体交换、抗氧化酶系统和根茎叶渗透调节物质的含量等硬性状,分析了不同季节放牧处理下羊草功能性状的变化及其权衡关系。结果表明,3年短期放牧处理下,类连续放牧(T1)比春季放牧样地(T2)羊草表现出更强的避牧性与耐牧性。类连续放牧与春季放牧样地羊草软性状及光合特性表现出一致性,6月放牧干扰降低了羊草的净光合速率(P_n),8月放牧干扰通过增加电子传递速率(ETR)及光系统Ⅱ(PSⅡ)分配于光化学反应(P)的比值等增大P_n。但春季放牧样地羊草株高较高,且光合产物较多分配于叶片,导致大量有机物质被啃食,不利于牧草再生。而类连续放牧羊草将较多的有机物质分配于根茎,有利于牧草根系吸水及牧后再生。因此,3年短期放牧处理下,类连续放牧更有利于牧草再生及草原的可持续利用。     

根茎在羊草响应短期NaCl胁迫过程中的作用

根茎是根状茎型克隆植物的特有结构,在养分储存、运输和分蘖茎的形成等方面起关键性作用。然而有关根茎生理学方面的研究却十分匮乏。为了探讨根茎在植物感应环境胁迫中的作用, 本文以羊草(Leymus chinensis)为实验材料,研究和比较了短期NaCl胁迫根、根茎、根和根茎3种处理方式下羊草对盐胁迫的响应。试验结果表明:200 mmol·L^-1 NaCl处理羊草根、根茎、根和根茎24 h,显著(p<0.05)降低羊草叶片净光合速率和蒸腾速率,增加叶片渗透浓度与脯氨酸含量;其中同时处理根和根茎叶片,蒸腾速率和净光合速率的降低程度显著高于分别处理根和根茎。在分别处理根与根茎的情况下,叶片含水量、脯氨酸含量、净光合速率、蒸腾速率均无显著性差异。不论单独胁迫根、根茎还是同时胁迫根和根茎,羊草根、根茎和叶片内Na⁺含量都显著高于对照,而羊草根、根茎和叶片内K⁺含量都显著低于对照。这些结果显示:1)根茎在羊草响应盐胁迫的生理过程中与根系具有类似的功能;2)羊草根茎在盐胁迫条件下能够有效地吸收Na⁺;3)鉴于根茎的生物量和表面积都明显地低于根系,在盐胁迫下羊草根茎吸收Na⁺的效率高于根系。     

新疆伊犁河谷平原狗牙根无性系构件的生长

采用随机挖掘整个无性系的方法,在新疆伊犁河谷平原2年无人工管理的葡萄园撂荒地,对狗牙根无性系进行了大样本取样,定量分析了构件生长的规律.结果表明：在30个狗牙根无性系中,分株数平均为（272.6±186.6）个,其中营养分株占总分株数的82.3%,是生殖分株的4.3倍;无性系总生物量为(45.4±40.0) g,其中根茎占总生物量的54.4%,营养分株占21.0%,匍匐茎占14.8%,生殖分株占9.4%;根茎与匍匐茎的累计长度分别为(5.1±4.7) m和(3.3±3.4) m、着生的芽数分别为(291.5±246.8)和(78.8±87.4)个.根茎和匍匐茎芽数与无性系的营养分株、生殖分株、根茎和匍匐茎的数量性状之间均呈极显著的正相关关系(P<0.01),表明在新疆伊犁河谷平原,狗牙根无性系主要以根茎芽实现其持续更新.     

Roles of succession,light, nutrients and disturbance on population vigor and maintenance of the rare plant Solidago shortii (Asteraceae)

Results of field and glasshouse experiments on Solidago shortii, and our observations on this species over many years, were used to construct a conceptual model of the roles of succession, light, soil nutrients and disturbance on population vigor and maintenance of this federal-endangered species. As cover of woody vegetation increased at a population site between 1986 and 1992, number of flowering ramets of S. shortii significantly decreased but number of vegetative ramets remained nearly constant. Adult plants transplanted into a redcedar thicket and those shaded in a glasshouse produced many fewer flowering ramets and capitula per flowering ramet and less biomass and had higher mortality than those in the open. Seedlings/juveniles shaded in a glasshouse had significantly less dry biomass; lower RGR, NAR, leaf area and root/shoot ratio and higher LAR, SLA and LWR than nonshaded ones. In a field site and glasshouse, fertilized plants (NPK) consistently had more flowering ramets and capitula per flowering ramet than nonfertilized ones. Hierarchy of dry weight of plants grown in a glasshouse in soils derived from five types of bedrock was phosphatic limestone > calcareous shale > sandstone > black shale = dolomite. Flowering and biomass production in the field-fertilizer and soil-type experiments were associated closely with levels of P. Number of flowering ramets significantly increased in plants transferred from shaded to nonshaded glasshouse conditions, but no such increase occurred after opening the canopy above plants in a thicket. Both high light and high nutrient levels apparently are necessary to maintain high vigor of S. shortii. In areas subject to invasion by woody plants, periodic high intensity disturbance may be required to prevent population extirpation.     

Nitrogen fertilizer and gender effects on the secondary metabolism of yaupon,a caffeine-containing North American holly

Females and males of sexually dimorphic species have distinct resource demands due to differential allocation to reproduction. Sexual allocation theory predicts that functional traits will diverge between sexes to support these demands. However, such dimorphism may be masked by the impact of current reproduction on source-sink interactions between vegetative and reproductive organs. We ask whether natural selection has led to genetic dimorphism in homologous physiological traits between sexes of the dioecious willow shrub, Salix glauca. In a common garden experiment we compared physiological responses to drought stress by male and female ramets in the absence of confounding demands from reproductive structures. Ramets experienced similar pre-dawn leaf water status (Ψ_l) as parental genets in flower within the natural population, indicating that experimental dry-down mirrored environmental conditions in nature. Male and female ramets achieved similar instantaneous water use efficiency, based on the ratio of carbon gain to water loss, under wet and dry conditions. However, female ramets experienced greater water stress (i.e., more negative Ψ_l) than males under dry conditions. Lower Ψ_l for female ramets may partly reflect the maintenance of conductance under drought; males, in contrast, maintain Ψ_l under drought by reducing conductance. Differences between sexes in terms of conductance and leaf water status of the vegetative ramets were absent in a concomitant comparison of parental flowering plants. Our results show (1) genetic divergence in physiology between sexes of S. glauca occurs in the absence of gender-specific reproductive sinks, (2) males are the more physiologically plastic sex with respect to water use, and (3) paradoxically, divergence in water relations between sexes is not detectable at sexual maturity under natural conditions.     

Dynamics of distribution and performance of ramets constructing genets: a demographic�Cgenetic study in a clonal plant, Convallaria keiskei

Background and Aims

In clonal plants producing vegetative offspring, performance at the genet level as well as at the ramet level should be investigated in order to understand the entire picture of the population dynamics and the life history characteristics. In this study, demography, including reproduction and survival, the growth patterns and the spatial distributions of ramets within genets of the clonal herb Convallaria keiskei were explored.

Methods

Vegetative growth, flowering and survival of shoots whose genets were identified using microsatellite markers were monitored in four study plots for 3 years (2003–2005). The size structures of ramets in genets and their temporal shifts were then analysed. Their spatial distributions were also examined.

Key Results

During the census, 274 and 149 ramets were mapped in two 1 × 2 m plots, and 83 and 94 ramets in two 2 × 2 m quadrats. Thirty-eight genotypes were identified from 580 samples. Each plot included 5–18 genets, and most ramets belonged to the predominant genet(s) in each plot. Shoots foliated yearly for several years, but flowering ramets did not have an inflorescence the next year. A considerable number of new clonal offspring persistently appeared, forming a bell-shaped curve of the size structure of ramets in each genet. Comparing the structures modelled by the normal distributions suggested variation among ramets belonging to a single genet and variation among genets. Furthermore, spatial analyses revealed clumped and distant distributions of ramet pairs in a genet, in which the distant patterns corresponded to the linearly elongating clonal growth pattern of this species.

Conclusion

Characteristics of ramet performances such as flowering and recruitment of clonal offspring, in addition to growth, played a large part in the regulation of genet dynamics and distribution, which were different among the studied genets. These might be characteristics particularly relevant to clonal life histories.Key words: Clonal plant, Convallaria keiskei, demography, genet, genetic identification, growth pattern, life history, ramet, spatial distribution     

Partial mechanical stimulation facilitates the growth of the rhizomatous plant Leymus secalinus: modulation by clonal integration

Background and Aims

Mechanical stimulation (MS) often induces plants to undergo thigmomorphogenesis and to synthesize an array of signalling substances. In clonal plants, connected ramets often share resources and hormones. However, little is known about whether and how clonal integration influences the ability of clonal plants to withstand MS. We hypothesized that the effects of MS may be modulated by clonal integration.

Methods

We conducted an experiment in which ramet pairs of Leymus secalinus were subjected to three treatments: (1) connected ramet pairs under a homogeneous condition [i.e. the proximal (relatively old) and distal (relatively young) ramets were not mechanically stressed]; (2) connected ramet pairs under a heterogeneous condition (i.e. the proximal ramet was mechanically stressed but the distal ramet was not); and (3) disconnected ramet pairs under the same condition as in treatment 2. At the end of the experiment, we harvested all plants and determined their biomass and allocation.

Key Results

Clonal integration had no significant influence on measured traits of distal L. secalinus ramets without MS. However, under MS, plants with distal ramets that were connected to a mother ramet produced more total plant biomass, below-ground biomass, ramets and total rhizome length than those that were not connected. Partial MS exerted local effects on stimulated ramets and remote effects on connected unstimulated ramets. Partial MS increased total biomass, root/shoot ratio, number of ramets and total rhizome length of stimulated proximal ramets, and increased total biomass, root weight ratio, number of ramets and total rhizome length of connected unstimulated ramets due to clonal integration.

Conclusions

These findings suggest that thigmomorphogenesis may protect plants from the stresses caused by high winds or trampling and that thigmomorphogenesis can be strongly modulated by the degree of clonal integration.     

Life history and population dynamics of Atriplex triangularis

The dynamics of a population of the stoloniferous Potentilla anserina was studied during four years. The mortality of established ramets followed a log-linear pattern superimposed on a seasonal fluctuation with an increased mortality during the period of vegetative spread. The major external mortality factors were unfavourable weather conditions such as drought and freezing. The turnover rate of the population was slow (31.6 yr). Vegetative spread dominated the recruitment to the population. Seedlings occurred but were exposed to a much higher mortality than vegetative daughter ramets. The juvenile period of seedlings is at least 4 yr in the field. This contrast to vegetative daughter ramets which are potentially flowering and stoloniferous the year after they are established, even if age affects both reproduction and vegetative spread. The density of most of the population is below the level where intraspecific regulative effects operate. However, in very dense stands both stolon and flower production were negatively density dependent.Nomenclature follows Lid (1974), Norsk og Svensk Flora, 2nd ed. Oslo.I am indebted to Lenn Jerling, Peter Torstensson, Anders Telenius and Gudrún Jónsdóttir for their help, and inspiring criticism throughout this study.     

Sex Ratios and Resource Allocation among Sexually Reproducing Plants of Rubus chamaemorus

Sex ratios and patterns of size variation and resource allocationwere investigated in the dioecious species Rubus chamaemorus.Sex ratios among flowering ramets varied from 6% to 40% of females.Female ramets were slightly, although not significantly, tallerthan males. It appeared that population effects (including bothgenetic population and environmental site effects) on plantsize and allocation patterns at flowering are considerably greaterthan sex effects. If both flowering and fruit production areconsidered, then female allocation to reproduction clearly exceedsmale allocation. In females, no significant relationship wasdetected between the mass of reproductive and vegetative tissues,while males did exhibit such a relationship. Reproductive effortwas less for tall males than for small males. Despite the occurrence of sexual reproduction, the main modeof reproduction in R. chamaemorus is vegetative propagation,which is the best strategy for reproduction in the unpredictableclimate of high latitudes but which leads to skewed sex ratios.As a consequence of vigorous vegetative reproduction, individualclones can grow to be large. The results of electrophoreticstudies show that the numbers of clones per population are low.Copyright1994, 1999 Academic Press Rubus chamaemorus, cloudberry, sex ratios, resource allocation, clonal structure, electrophoresis     

Population and community-level compositional patterns shape the realized niche of the rare arctic-alpine species Carex lachenalii Schkuhr

The ability for vegetative growth and development of generative organs often reflects an adaptation to the environment and may be a suitable proxy for understanding population dynamics of rare relict species. An example of such a plant is Carex lachenalii Schkuhr, an arctic-alpine species, in the temperate zone of Europe only occurring in isolated localities of high-elevation mountain ranges. We aimed to assess whether there were relationships between flower production and clonal growth of C. lachenalii, both at the tuft and plot level, and how co-occurring vegetation could modify this relationship. In the study we focused on population-level traits of C. lachenalii, vegetation traits and components of functional diversity. At the tuft level we found that the proportion of flowering ramets of C. lachenalii decreased with increasing diameter of the tuft. At the plot level, in snowbed vegetation C. lachenalii produced more flowering ramets. We suggest this is due to higher environmental stress, expressed by high importance of habitat filtering (low functional dispersion) in shaping species composition of co-occurring vegetation. In granite grasslands and milder environment (expressed by higher functional dispersion), C. lachenalii produced more vegetative ramets, which we suggest is a result of a more competitive environment. While in snowbeds investment in flowering ramets could promote successful persistence of C. lachenalii, survival of subpopulations occurring in the highly competitive conditions of granite grasslands may be uncertain due to potentially weak adaptation to competition with graminoids and dwarf shrubs.     

Genets of dwarf bamboo do not die after one flowering event: evidence from genetic structure and flowering pattern

Dwarf bamboos in the genus Sasa are believed to be long-lived, synchronously flowering, and monocarpic plants. However, the monocarpy of dwarf bamboo has not been confirmed, because whether all ramets within one genet flower at the same time cannot be determined without differentiating the genetic structure among ramets. This study aims to evaluate the reproductive traits of Sasa pubiculmis by verifying the monocarpy and physiological integration between flowering ramets and non-flowering ramets during a 4-year flowering period. One genotypically identified genet, which covered an area of approximately 3 ha, had both flowering and non-flowering patches of ramets during the 4-year flowering period (2004–2007). A fraction of the flowering genet remained non-flowering during the 4 years of observation, and did not die after mass flowering. Flowering ramets were physically connected to non-flowering ramets via rhizomes, and assimilated ¹³C was allocated from non-flowering ramets to flowering ramets. Consequently, we clarified that this dwarf bamboo potentially has polycarpic reproductive traits rather than monocarpic, and a genet can keep rhizomes and non-flowering patches alive to sustain the organism after mass flowering.     

The Control of Apical Bud Growth and Senescence by Auxin and Gibberellin in Genetic Lines of Peas

Pea (Pisum sativum L.) lines G2 (dwarf) and NGB1769 (tall) (Sn Hr) produce flowers and fruit under long (LD) or short (SD) days, but senesce only under LD. Endogenous gibberellin (GA) levels were inversely correlated with photoperiod (over 9-18 h) and senescence: GA20 was 3-fold and GA1 was 10- to 11-fold higher in flowering SD G2 shoots, and the vegetative tissues within the SD apical bud contained 4-fold higher levels of GA20, as compared with the LD tissues. Prefloral G2 plants under both photoperiods had GA1 and GA20 levels similar to the flowering plants under LD. Levels of indole-3-acetic acid (IAA) were similar in G2 shoots in LD or SD; SD apical bud vegetative tissues had a slightly higher IAA content. Young floral buds from LD plants had twice as much IAA as under SD. In NGB1769 shoots GA1 decreased after flower initiation only under LD, which correlated with the decreased growth potential. We suggest that the higher GA1 content of G2 and NGB1769 plants under SD conditions is responsible for the extended vegetative growth and continued meristematic activity in the shoot apex. This and the increased IAA level of LD floral buds may play a role in the regulation of nutrient partitioning, since more photosynthate partitions of reproductive tissue under LD conditions, and the rate of reproductive development in LD peas is faster than under SD.     

Functional specialization of ramets in Scirpus maritimus – Splitting the tasks of sexual reproduction, vegetative growth, and resource storage

Clonal plant species can be considered as populations of interconnected ramets which are basically identical in form and function, and potentially independent from each other. Experimental studies and field observations suggest that an intra-clonal specialization of ramets with different roles (division of labour) can increase the performance of clonal systems under heterogeneous conditions. This paper explores structural and functional variation in the emergent macrophyte Scirpus maritimus, which forms ramets that specialize in three main activities: sexual reproduction, photosynthetic assimilation and vegetative growth, and reserve storage. The main question asked in this study is whether such specialization is a developmentally programmed syndrome in this species, and whether environmental conditions can alter the pattern of ramet differentiation along rhizome systems.We analyzed clonal fragments collected from a population in the field, and grew clones individually in pots of two sizes to simulate different degrees of crowding and shoot density. Specialization of ramets was largely predictable from their position along the rhizome system, indicating that specialization is an inherent feature (developmentally programmed) of clone ontogeny in S. maritimus. In the field, sexual ramets were always situated at the base of rhizome systems, vegetative ramets were in first and intermediate positions, and shoot-less storage ramets were almost always formed distally on rhizomes (terminal ramets). In the pot experiment flowering ramets were observed in all positions along rhizome systems, suggesting that specialization for sexual reproduction shows a plastic response to environmental conditions.S. maritimus can adjust the relative numbers of ramets with and without above-ground shoots when grown in different shoot densities, i.e. the frequency of individual ramets responsible for a certain functional or developmental process can be adjusted to environmental conditions and internal needs. In S. maritimus, the density-dependant regulation of storage versus vegetative growth and sexual reproduction may represent a mechanism to limit shoot competition in crowded populations.