Morphological responses to nutrient availability in four clonal herbs

This paper examines morphological plasticity of clonal plants of contrasting habitats and of contrasting architectures in response to nutrient supply. The hypotheses were tested that plants from rich habitats possess greater plasticity in response to variation in resource supply than species from poor habitats, and that rhizomatous species are less plastic in their response than stoloniferous species. Two sympodial rhizomatous herbs (Carex flacca, C. hirta) and two monopodial stoloniferous herbs (Trifolium fragiferum, T. repens) were subjected to four levels of nutrient supply in a garden experiment. One of the two species of each genus (C. hirta, T. repens) is from fertile and the other from infertile habitats. We measured 1) whole plant characters: total plant dry weight, number of modules (product of a single apical meristem) and number of ramets; 2) ramet characters: ramet leaf area and ramet height; and 3) spacer characters: branches per module, length per module and length per module internode.All measured characters in the Trifolium species significantly responded to treatment: the values for all measured characters increased with higher levels of fertilization. The differences in plant characters between fertilization levels were larger in Trifolium repens than in T. fragiferum in terms of whole plant characters, ramet characters and stolon internode length. The two Carex species did not differ in their responses to treatment in terms of most characters measured. In ramet characters and in some whole plant characters the species from fertile habitats were more plastic than those from infertile habitats. In spacer characters this pattern was not found. Foraging could not be demonstrated unequivocally.Morphological plasticity in the stoloniferous (Trifolium) species was much larger than in the rhizomatous (Carex) species. This seems in accordance with a foremost storage function of rhizomes, as against a foremost explorative function of stolons.     

Costs of plasticity in foraging characteristics of the clonal plant Ranunculus reptans

In clonal plants, evolution of plastic foraging by increased lengths of leaves and internodes under unfavourable conditions may be constrained by costs and limits of plasticity. We studied costs and limits of plasticity in foraging characteristics in 102 genotypes of the stoloniferous herb Ranunculus reptans. We grew three replicates of each genotype with and three without competition by the naturally co-occuring grass Agrostis stolonifera. We used regression and correlation analyses to investigate potential costs of plasticity in lengths of leaves and stolon internodes, developmental instability costs of these traits, and a developmental range limit of these traits. We used randomization procedures to control for spurious correlations between parameters calculated from the same data. Under competition the number of rosettes, rooted rosettes, and flowers was 58%, 40%, and 61% lower, respectively, than in the absence of competition. Under competition lengths of leaves and stolon internodes were 14% and 6% smaller, respectively, than in the absence of competition. We detected significant costs of plasticity in stolon internode length in the presence of competition when fitness was measured in terms of the number of rosettes and the number of flowers (selection gradients against plasticity were 0.250 and 0.214, respectively). Within-environment variation (SD) in both foraging traits was not positively correlated with the corresponding plasticity, which indicates that there were no developmental instability costs. More plastic genotypes did not have less extreme trait values than less plastic genotypes for both foraging traits, which indicates that there was no developmental range limit. We conclude that in R. reptans costs of plasticity more strongly constrain evolution of foraging in the horizontal plane (i.e., stolon internode length) than in the vertical plane (i.e., leaf length).     

林缘和荒草坡不同草本层盖度小生境中积雪草的等级可塑性

为研究克隆植物在不同生境中对小生境的利用能力和对不同尺度异质性生境的等级可塑性及其格局差异,比较了林缘和荒草坡不同草本层盖度小生境中积雪草(Centella asiatica)分株种群、克隆片段和叶水平的形态和生理特征。在分株种群水平,林缘和荒草坡积雪草被测的分株种群特征差异均不显著,不同草本层盖度小生境间积雪草根冠比差异显著,即在盖度小的小生境中其根冠比显著大于盖度大的小生境。在克隆片段水平,林缘和荒草坡生境显著影响积雪草的根干重、花果干重、一级匍匐茎节间长和总匍匐茎节间长,而不同草本层盖度小生境则显著影响花果干重、根冠比、一级匍匐茎数、二级匍匐茎节间长和总匍匐茎节间长。荒草坡积雪草拥有更大的根干重和花果干重,但一级匍匐茎节间长和总匍匐茎节间长较短。草本层盖度小的小生境中积雪草的根冠比和一级匍匐茎数显著高于盖度大的小生境,而花果干重、二级匍匐茎节间长和总匍匐茎节间长则相反。在叶水平,荒草坡和草本层盖度大的小生境中,积雪草的叶片长、叶片宽和叶面积都较大,而草本层盖度大的小生境中积雪草叶柄长和比叶柄长都显著大于盖度小的小生境;林缘和荒草坡生境与不同草本层盖度小生境对4个叶绿素指标影响均不显著,林缘和荒草坡生境与不同草本层盖度小生境的交互作用对所有测量特征均无显著影响。上述结果表明,积雪草对不同生境和有草本层盖度差异的小生境的反应存在等级可塑性,但是二者的等级差异格局不同;等级反应格局的差异反映出积雪草对不同生境条件有不同的生态适应对策,且对生境小尺度异质性的可塑性强于对大尺度异质性的可塑性。     

How internode length,position and presence of leaves affect survival and growth of <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis> after fragmentation?

Disturbance is common in nature and disturbance-caused fragmentation of clones happens frequently in stoloniferous plants. After fragmentation storage in stolon internodes and leaves may enhance survival and growth of stoloniferous plants. We hypothesize that (1) increasing length of the internode attached to the ramet and (2) presence of leaves will increase ramet survival and growth, and that (3) internode positions (before or after the ramet or both) will also play a role. We tested these hypotheses with the stoloniferous, invasive herb Alternanthera philoxeroides. In one experiment, we measured survival and growth of the ramets either without stolon internode (0 cm in length) or attached with internodes of 2, 4, 6 and 8 cm and either with or without leaves. In the other experiment, we measured survival and growth of the ramets attached with a proximal internode (before the ramet), a distal internode (after the ramet) or both. Increasing internode length and presence of leaves significantly increased the survival rate and growth (biomass, leaf area, number of ramets, stolon length and number of leaves) of the A. philoxeroides plants. All growth measures of A. philoxeroides at harvest were larger when the ramets were attached with a distal internode than when they were attached with a proximal internode, but the survival rate was lower. These results support the hypotheses and suggest that storage in stolons and leaves may be of great significance for clonal plants in frequently disturbed habitats and may contribute greatly to the invasiveness of A. philoxeroides.     

The role of stolon internodes for ramet survival after clone fragmentation in Potentilla anserina

Stoloniferous herbs are common in disturbed habitats, where interconnected ramets frequently become separated by water, grazing animals or human activity. Stolon internodes serve as spacers and transport channels for resources, but they may also fulfil a storage function. We simulated physical disturbance and breakage of stolon connections in the stoloniferous herb Potentilla anserina, to test the hypothesis that juvenile ramets can survive clone fragmentation better if they remain attached to an internode after being severed from the rest of the clone. Juvenile, unrooted ramets were cut off from stolons with and without the preceding (basal) internode and placed on wet soil in a greenhouse. All ramets with an attached internode survived, while survival rates dropped to 37% for ramets from which internodes had been removed. The dry weight per unit volume of internodes decreased by about 20% during the experiment. We conclude that stolons are likely to fulfil a storage function that may be of ecological importance in disturbed habitats.     

Variation in petiole and internode length affects plant performance in <Emphasis Type="Italic">Trifolium repens</Emphasis> under opposing selection regimes

We studied the effects of genotypic and plastic variation in vertical and horizontal spacer lengths on plant performance in a stoloniferous herb subjected to opposing selection regimes. We hypothesized that longer vertical structures are beneficial if plants are subjected to competition, but they should negatively affect plant performance if plants are exposed to aboveground disturbance. To test these hypotheses we subjected 34 genotypes of Trifolium repens to competition and disturbance treatments. Competition was imposed by a grass canopy consisting of Lolium perenne, and disturbance was simulated by regularly clipping the target plants and all the surrounding vegetation at 1 cm above soil level. Conform to our hypothesis, genotypes with longer vertical structures (petioles) produced fewer ramets than genotypes with shorter petioles in the disturbance treatment. However, genotypes with longer petioles did not perform better under competition than genotypes with shorter petioles. Genotypes with highly plastic vertical structures tended to produce more shoot mass under competition, and they produced fewer ramets if subjected to disturbance. Unexpectedly, horizontal structures (stolon internodes) expanded in response to competition which, furthermore, was associated with enhanced plant performance. However, producing longer internodes is inherently associated with costs in terms of increased resource allocation to the longer structures, but not to benefits in terms of increased resource capture. Positive correlations among the length and plasticity of vertical and horizontal structures may explain the apparent positive effect of producing longer internodes on plant performance. Our data thus support the notion that trait correlations may weaken selective forces acting on a focal trait in a specific environment if opposing selection pressures act on genetically correlated traits.     

光照强度和基质养分对匍匐茎草本金戴戴克隆生长和克隆形态的影响

在深度遮光(光照强度为高光条件的6.25%,约为自然光照的5.3%)或低养分条件下,金戴戴(Halerpestes ruthenica Ovcz.)生物量、初级分株叶面积、分株总数、匍匐茎总数和总长度均显著减小,而比节间长和比叶柄长显著增加.在低养分条件下,金戴戴匍匐茎平均节间长显著增加,而匍匐茎分枝强度和分株数显著减小.这些结果与克隆植物觅食模型相符合,表明当生长于异质性生境中,金戴戴可能通过以克隆生长和克隆形态的可塑性实现的觅养行为来增加对养分资源的摄取.在深度遮光条件下,金戴戴平均间隔子长度(即平均节间长和平均叶柄长)均显著减小.这一结果与以往实验中匍匐茎草本间隔子对中度和轻度遮光(光照强度为高光条件的13%～75%,>10%的自然光照)的反应不同.这表明,在深度遮光条件下匍匐茎克隆植物可能不发生通过间隔子可塑性实现的觅光行为.光照强度和基质养分条件的交互作用对许多性状如总生物量、匍匐茎总数和总长度、二级和三级分株数、分株总数、初级分株叶面积以及分枝强度均有十分显著的效应.在高光条件下,基质养分对这些性状有十分显著的影响;而在低光条件下,基质养分条件对这些性状不产生影响或影响较小.这表明,光照强度影响金戴戴对基质养分的可塑性反应.在深度遮光或低养分条件下,金戴戴可能通过减小匍匐茎节间粗度(增加比节间长)来增加或维持其相对长度,从而更有机会逃离资源丰度低的斑块.     

Physiological integration helps a clonal macrophyte spread into competitive environments and coexist with other species

Physiological integration may help clonal macrophytes invade or escape from existing communities. No studies have tested the above hypothesis in aquatic plants. In an outdoor pond experiment, we subjected clonal fragments of the submerged macrophyte Vallisneria spiralis L. to heterogeneous environments in which V. spiralis spread from bare habitats towards vegetated habitats occupied by Myriophyllum spicatum L. or V. spiralis spread from vegetated habitats towards bare habitats. V. spiralis stolons between ramets in bare habitats and in vegetated habitats were either intact or severed. We investigated the habitat selection of V. spiralis by examining the allocation of biomass and ramets to heterogeneous habitats during its vegetative spread phase. Results showed that the stolon connection had different effects on the habitat selection of V. spiralis with regard to invasion and escape. When V. spiralis spread from bare to vegetated habitats, in comparison to severing the stolon, the stolon connection eventually facilitated a 49% increase in biomass and a 27% increase in number of ramets allocated to vegetated habitats. However, when V. spiralis spread from vegetated to bare habitats, biomass and ramets allocated to bare habitats were not significantly changed by the stolon connection (only a 5% increase in biomass and a 6% increase in number of ramets). These results indicate that clonal integration facilitated V. spiralis not to escape from but invade into vegetated habitats. The study provides evidence that physiological integration is important for survival and tolerance of ramets in competitively stressful environments and can help clonal macrophytes coexist with other species.     

光照强度和基质养分对匍匐茎草本金戴戴克隆生长和克隆形态的影响

在深度遮光 (光照强度为高光条件的 6 .2 5% ,约为自然光照的 5.3% )或低养分条件下 ,金戴戴 (HalerpestesruthenicaOvcz.)生物量、初级分株叶面积、分株总数、匍匐茎总数和总长度均显著减小 ,而比节间长和比叶柄长显著增加。在低养分条件下 ,金戴戴匍匐茎平均节间长显著增加 ,而匍匐茎分枝强度和分株数显著减小。这些结果与克隆植物觅食模型相符合 ,表明当生长于异质性生境中 ,金戴戴可能通过以克隆生长和克隆形态的可塑性实现的觅养行为来增加对养分资源的摄取。在深度遮光条件下 ,金戴戴平均间隔子长度 (即平均节间长和平均叶柄长 )均显著减小。这一结果与以往实验中匍匐茎草本间隔子对中度和轻度遮光 (光照强度为高光条件的 1 3%～ 75% ,>1 0 %的自然光照 )的反应不同。这表明 ,在深度遮光条件下匍匐茎克隆植物可能不发生通过间隔子可塑性实现的觅光行为。光照强度和基质养分条件的交互作用对许多性状如总生物量、匍匐茎总数和总长度、二级和三级分株数、分株总数、初级分株叶面积以及分枝强度均有十分显著的效应。在高光条件下 ,基质养分对这些性状有十分显著的影响 ;而在低光条件下 ,基质养分条件对这些性状不产生影响或影响较小。这表明 ,光照强度影响金戴戴对基质养分的可塑性反应。在深度遮光     

Life history variation and infraspecific heterochrony inRhinanthus glacialis (Scrophulariaceae)

Comparative cultivation experiments were made with 14 populations of the annual, hemiparasitic speciesRhinanthus glacialis from seven different habitat types. Phenological data were recorded and plants from field samples and from corresponding experimental samples were morphologically analysed.There are up to ten weeks difference in the start of flowering between means of the extreme experimental populations. Flowering time variation correlates with the number of internodes and also with most of the other morphological characters. Flowering time also shows close relationships to different grassland types and the examined populations appear to be well-adapted to the environmental conditions in natural or man-influenced grassland types as far as flowering time is concerned. The number of internodes is mostly insensitive to the applied experimental influences. The rate of internode production is constant. Trade-offs are suggested for some morphological traits (i.e., few internodes with few branches and thus few flowers as trade-off for early flowering). — Previous studies of life history variation based on population data have not been incorporated into the theory of heterochrony. The pattern of life history variation of populations ofR. glacialis together with information about morphology, ecology, and phylogeny suggests that some variants have evolved from ancestors via heterochronic processes. Earlier flowering populations from montane hay meadows, litter meadows, and dry meadows belong to paedomorphic variants. These have fewer internodes and branches as well as reduced height in comparison with the suggested ancestor. Later flowering populations from subalpine limestone grassland belong to a peramorphic variant with more internodes, more branches, and taller stems in comparison with the suggested ancestor.     

Senescence and decomposition of white clover stolons in grazed upland grass/clover swards

The fate of marked sections of stolons of white clover (Trifolium repens) over a 50-week period from May 1987 was followed in grazed grass/clover swards maintained at 5-cm sward surface height with and without N fertiliser. There was little effect of N treatment on the pattern of survival of stolon sections. The proportion of live stolons recovered decreased during the experiment, and in May 1988 on average only 29% of the marked sections remained alive. At all harvests only a small percentage of stolon sections showed signs of senescence; the maximum percentage, on average 20% of those marked, occurred in autumn, 15–20 weeks after marking. Following this period, i.e. in late autumn/winter, the most rapid increase in percentage of decomposed stolons was measured. Over 50% of stolon sections were buried within the 5-week period following marking and nearly all were buried after 20 weeks; generally a much smaller proportion of stolon tips was buried. Nutrient concentrations of N, P and K fell to their lowest levels in autumn, before increasing in the following spring. Results are discussed in relation to the cycling of nutrients via stolon senescence.     

The role of vegetative spread and seed dispersal for optimal life histories of clonal plants: a simulation study

Sexual and vegetative reproduction of clonal plants phenotypically differ in dispersal distance, in the phenology of offspring production and establishment, and in the success of establishment. We applied a combination of analytical and of spatially explicit individual-based simulation modelling to calculate long-term fitness. We predicted optimal clonal plant life histories in a parameter space spanned by stolon number, stolon internode length, and relative allocation to sexual and vegetative reproduction. For a given allocation to sexual reproduction and number of stolons, fitness was optimised for rather short internode lengths under small disturbances, and for the longest possible internodes under larger disturbances. A trade-off between length and number of vegetative spacers drew parameters away from their unrestricted optima. Now, intermediate length and number of spacers led to maximum fitness under large disturbances. Simultaneous trade-offs between sexual and vegetative reproduction and between the length and number of spacers could also lead to fitness optima at intermediate parameter values, depending on the success of seedling establishment. We demonstrated that spatial habitat structure (1) selects for an efficient use of available space either by optimum internode length or by investment into seeds, which disperse farther than vegetative spacers, and (2) leads to an interaction between trade-offs. We conclude, that dispersal distance, i.e. a spatial life-history component, and trade-offs must be included in considerations on adaptive evolution of clonal life histories.     

海岸单叶蔓荆沙埋胁迫下碳水化合物变化与其耐沙埋的关系

选择烟台海岸沙地抗沙埋强的单叶蔓荆(Vitex trifolia var.simplicifolia)为试材,在自然环境条件下根据单叶蔓荆匍匐茎长度进行了轻度(1/3茎长)、中度(2/3茎长)和重度半埋以及全埋处理。在沙埋20d后,测定了不同沙埋处理下匍匐茎各段上匍匐茎长度、枝条高度、不定根长度,以及可溶性糖、淀粉、纤维素含量,以探讨单叶蔓荆碳水化合物变化和转化在其耐沙埋中作用。结果显示,在轻度、中度半埋和全埋下单叶蔓荆匍匐茎长度均显著大于对照,被沙埋匍匐茎处有大量不定根生成;同时,可溶性糖和淀粉含量增高和纤维素含量下降,尤其是生长最快的匍匐茎顶部(如轻度半埋),茎中可溶性糖较低、淀粉增加最多,纤维素最低。但是被重度半埋和全埋的匍匐茎生长较少,茎中纤维素含量较多、淀粉含量较少。研究表明,沙埋是一种胁迫,它损伤叶片、扰乱碳水化合物代谢平衡。但它又是胁迫信号使植物产生适应性反应,它使未遭沙埋的匍匐茎顶端通过加速碳水化合物转化、分解纤维素、提高淀粉和可溶性糖含量,为顶端生长提供能量和营养,以加速匍匐茎快速生长摆脱沙埋。同时沙埋部位枝叶通过分解其纤维素,产生更多的可溶性糖和淀粉为匍匐茎不定根生长提供能量。因此,沙埋后匍匐茎内碳水化合物的转化是其快速生长和摆脱沙埋的能量来源而在其适应沙埋生长中起重要作用。单叶蔓荆对沙埋的适应性反应表现了其具有表型可塑性特性,该特性是其沙埋后维护匍匐茎顶部快速生长、不定根形成、碳水化合物转化以及具有较高抗沙埋能力的关键。     

Morphological responses to local light conditions in clonal herbs from contrasting habitats,and their modification due to physiological integration

Morphological responses to light and effects of physiological integration on local morphological responses are examined for Hydrocotyle vulgaris and Lamiastrum galeobdolon, stoloniferous herbs from open fenlands and forest understoreys, respectively. An assessment was made of whether these clonal herbs of similar morphology but from contrasting habitats show different foraging behaviour for light. In a garden experiment, the plants wer subjected to four levels of light availability, and to a split treatment in which the primary stolons grew along the border of patches of the two intermediate light levels. In this treatment the plant parts on opposite sides of the primary stolons were in contrasting light environments. Petiole extension was more responsive to light conditions in Hydrocotyle than in Lamiastrum, while the opposite was true for leaf area. Both species showed similar responses in stolon internode length and specific leaf area (SLA). Integration did not significantly modify local responses in stolon internode length in either species. Local responses in petiole length, leaf area and SLA of Hydrocotyle ramets were not significantly affected by physiological integration, except for the SLA of ramets in high light which was evened out by integration. In contrast, in Lamiastrum, local responses in petiole length, leaf area and SLA of many ramets in the shaded and/or light patch were significantly evened out by integration. As a result, interconnected ramets in patches of different light supply developed very different morphologies in Hydrocotyle, but not in Lamiastrum. The results indicate that the species differed in ramet morphological responses to light intensity as well as in effects of integration on local morphological responses, and suggest that species from different habitats show different foraging behaviour for light.     

Environmental correlates of growth traits of the stoloniferous plant Potentilla palustris

Growth form is one of the important life history traits ultimately influencing plant fitness. Potentilla palustris is a stoloniferous plant growing in a range of habitats from densely vegetated wet meadows to acidic transitional fens, and its growth form varies according to habitat. In a four year multi-site comparative study, we investigated which biotic and abiotic characteristics influence most its growth traits. Vegetation composition and physiognomy, as well as numerous abiotic environmental variables, were recorded at 32 study sites located on an altitudinal gradient. Growth traits of P. palustris were best explained by the surrounding vegetation physiognomy and not by abiotic conditions, although the latter obviously represents the factors indirectly influencing its growth. Stolon length traits and branching were positively correlated with vegetation density and height, and negatively with altitude. Plants flowered more in taller vegetation, and leaf area was greater in wetter sites with lower vegetation cover. Potentilla palustris appeared to be well adapted to transitional fens, but its vegetative growth was fastest in wet meadows and alluvial habitats on highly organic humid soils. It produced more branches and larger leaves in alluvial habitats with open water, while it had enhanced generative reproduction in wet meadows. Species composition was less important than vegetation physiognomy. In less favorable habitat types, P. palustris prefers an escape strategy of linear growth. Internode length exhibited pronounced plasticity, increasing particularly in tall dense vegetation of lower altitude, whereas internode number remained fairly constant over various habitats. It is evident that both plastic low cost growth traits (internode elongation), and constant high cost traits (internode number) contribute to the P. palustris escape strategy under tall dense vegetation. Phenotypic plasticity enhances the potential of P. palustris to grow in a wide range of habitats and so increases plant fitness on regional scale.     

Aspects of the Comparative Physiology of Ranunculus bulbosus L. and Ranunculus repens L.: II. Carbon Dioxide Assimilation and Distribution of Photosynthates

Detailed analysis of the interrelationships between sourcesof photosynthate production and sites of utilization in thetaxonomically closely related species Ranunculus bulbosus L.and R. repens L. showed that leaves whether present on rosette,stem, or stolon had similar levels of ¹⁴CO₂-fixation but thepattern of distribution of radiocarbon to the rest of the plantdiffered. Fruits of R. bulbosus had a lower fixation rate thanleaves but were characterized by total retention of the fixedradiocarbon. Rosette leaves of R. bulbosus supplied the youngleaves, developing apices in the rosette, roots, and corms,whereas the labelled assimilates from cauline leaves were evenlydistributed between reproductive and vegetative parts. The cormwas the major sink both at the flowering and fruiting stages.When plants were treated with ¹⁴CO₂ in the field even higherlevels of radiocarbon moved into the corm than in comparableexperiments under greenhouse conditions. The rosette leaf ofR. repens exported mainly to actively growing stolons in plantswith many stolons bearing rooted ramets although growth of astolon was also substantially supported by photosynthates producedby its own ramets. A proportion of the radiocarbon fixed byleaves of mature ramets was exported and moved in a predominantlyacropetal direction into the stolon apex, stolon axis, and youngramets of the same stolon. The stock in R. repens had a muchlower demand for assimilates than the corm in R. bulbosus. The results are consistent with the concept that R. bulbosusoperates a conservative policy involving the replacement ofthe parent in situ by a daughter from the corm, coupled withextensive fruit production. In R. repens the emphasis is onlateral spread and exploitation of substantial areas of groundby vegetative spread and replacement of the parent by daughtersmany of which may occupy sites some distance from the parent.     

不同光照条件下聚花过路黄的克隆构型和分株种群特征

以都江堰灵岩山常绿阔叶林林下、林缘和林缘旷地3种不同光照环境中匍匐茎草本聚花过路黄(Lysimachla congestiflora)为对象,对其匍匐茎节间长、分枝强度、分枝角度、分株种群密度、分株种群生物量等指标进行了测定和分析．结果表明,聚花过路黄的匍匐茎节间长、分枝强度、分株种群密度和分株种群生物量在3种生境间差异显著．Kruskal Wallis检验表明,匍匐茎节间长度和分枝角度的频次分布在3种生境间差异显著．3种生境中匍匐茎节间长度分布偏斜度(skewness)的大小分别为：林缘旷地>林缘>林下．林缘旷地与林缘和林下生境在分枝角度、分株种群高度和分株种群根冠比差异显著,而且林缘旷地生境中分枝角度分布偏斜度最小．林缘和林下生境在分枝角度、分株种群高度和分株种群根冠比差异不显著．从林缘旷地、林缘到林下,聚花过路黄的克隆构型和分株种群特征发生了相应改变．结合克隆植物对资源的利用对策,讨论了不同生境中聚花过路黄克隆构型和分株种群特征可塑性的生态适应意义．     

Clonal growth strategies in simultaneously persistent and expanding <Emphasis Type="Italic">Trifolium repens</Emphasis> patches

Plants with clonal growth can generate patches dominated by a single species. In time, patches can change and may fragment, form a ring, dissolve or both persist and expand. For patches to maintain their original habitat and simultaneously increase in size, ramets or clonal fragments must both promote local persistence inside the patch and grow out of the patch into new habitats. This study analyses simultaneously expanding and persistent Trifolium patches in a nutrient-poor lawn that is frequently cut, and where the Trifolium is competitively superior to the grass species. Trifolium primary stolon growth strategies were analyzed in relation to their location (border, middle, and center) inside the patch, and according to patch size (small, medium, and large). It was hypothesized that different growth strategies inside a patch can explain both persistent and expanding patch of Trifolium, and that growth strategies were different between patch sizes. Primary Trifolium stolons had two different growth strategies inside and at the border of patches: (i) stolons at the border were long, grew fast, had few lateral stolons, and grew out of the patch, while (ii) stolons inside the patch were smaller, grew slowly, and had more lateral stolons and a wide range of growth directions. Growth strategies were not different between patch sizes. The directional growth and the high growth rate at the border will increase the patch size with time, while the growth strategy near the center consolidates the patch in space and time, by placing ramets inside the patch. Different growth strategies near the center and on the border result in Trifolium patches that are simultaneously persistent and on the increase. The results also indicate a division of labor among primary Trifolium stolons in a patch.     

Phosphate Uptake and Transport by Roots and Stolons of Intact White Clover (Trifolium repens L.)

The extent to which phosphate can be absorbed directly fromthe outer medium by stolon internodes and contribute to thetotal accumulation of phosphate by intact plants of white clover(Trifolium repens L. cv. Blanca) was assessed in hydroponicexperiments in a controlled environment room. The uptake ofphosphate by intact roots or stolons was measured by sealinga segment (6-0 mm long) across a flow-cell in which ³²P-labellednutrient solution was circulated for 24 h, the rest of the rootsystem receiving unlabelled nutrient solution. The rate of uptakeof phosphate (µmol g^–1 d^–1 dry wt. basis)by roots was more than 300 times that by intact stolons. Pretreatmentof stolons by gentle abrasion to remove cuticle, so as to simulatethe condition of stolons in the field, increased the uptakeof phosphate 7-fold compared with that of intact stolons. However,the potential of stolons to contribute to the P status of whitedover in the field was calculated to be small (5%). When an incision was made through the hypodermal layer of stolons,the rate of phosphate uptake greatly increased, attaining 71%of that by root segments. This increase, which was greater athigher phosphate concentrations, indicates that the suberi.zedhypodermis constitutes a major barrier to the influx of phosphatein the stolon. After withholding phosphate for different time intervals, thesubsequent rate of phosphate uptake by roots was increased 2-3-foldafter 2 d phosphate deprivation and 3-4-fold after 6 d or 13d phosphate deprivation. A higher proportion of absorbed phosphatewas transported to shoots in phosphate-deprived plants. After1 d of uptake following restoration of the phosphate supply,the concentrations of labelled phosphate in shoots were greaterthan in control plants, although the concentrations of labelin roots was less. However, the rate of uptake of phosphateby stolons, following deprivation, was not significantly increased.These results suggest that the mechanism regulating the enhancedrate of phosphate loading into the xylem, initiated by a periodof phosphate deprivation, is specific to roots and is not inducedin stolons. The results are discussed in relation to the growth and acquisitionof phosphate by white clover in the field. Key words: Nutrient deficiency, phosphate, stolons, transport (ions), Trifolium repens     

Habitat selection and web plasticity by the orb spider Argiope keyserlingi (Argiopidae): Do they compromise foraging success for predator avoidance?

Abstract Orb web spiders face a dilemma: forage in open habitats and risk predation or forage in closed habitats to minimize risk but at reduced foraging profitability. We tested whether Argiope keyserlingi opts for safer habitats at the expense of foraging success by (i) determining habitat selection indices in open and closed habitats; (ii) marking and releasing individual juvenile, subadult and adults over two 4‐week periods to determine if life‐history stage influences habitat selection; and (iii) determining the biotic and abiotic environmental parameters that relate to A. keyserlingi abundance. We found that A. keyserlingi selected closed habitats. Sedge and anthropogenic structures were selected and trees were avoided. Juveniles were never found in open habitats, most likely because of high postdispersal mortality. Subadults and adults may shift from closed to open habitats while juveniles never shifted habitat. Foliage density, plant height, potential prey abundance, and mantid and bird abundance were correlated with A. keyserlingi abundance, with only bird abundance explaining habitat selection. We measured web capture area, spiral distance (distance between spiral threads) and the number of decoration arms (0, 1, 2, 3 or 4) in the field and did laboratory experiments to test the influence of (i) space and vegetation; (ii) prey abundance; and (iii) web damage, on web architecture. Argiope keyserlingi webs exhibited geometric plasticity by having larger prey capture areas and spiral distances in open habitats. Decoration design did not differ between habitats however. Variation in space availability, air temperature, prey abundance and web damage explained the variations in web architecture. Potential prey size and diversity differed between habitats but prey abundance did not. As large prey may be important for spider survivorship, foraging success appears to be compromised by occupying closed habitats.