Small-scale heterogeneity in soil quality influences photosynthetic efficiency and habitat selection in a clonal plant

BACKGROUND AND AIMS: In clonal plants, internode connections allow translocation of photosynthates, water, nutrients and other substances among ramets. Clonal plants form large systems that are likely to experience small-scale spatial heterogeneity. Physiological and morphological responses of Fragaria vesca to small-scale heterogeneity in soil quality were investigated, together with how such heterogeneity influences the placement of ramets. As a result of their own activities plants may modify the suitability of their habitats over time. However, most experiments on habitat selection by clonal plants have not generally considered time as an important variable. In the present study, how the foraging behaviour of clonal plants may change over time was also investigated. METHODS: In a complex of environments with different heterogeneity, plant performance was determined in terms of biomass, ramet production and photosynthetic activity. To identify habitat selection, the number of ramets produced and patch where they rooted were monitored. KEY RESULTS: Parent ramets in heterogeneous environments showed significantly higher maximum and effective quantum yields of photosystem II than parents in homogeneous environments. Parents in heterogeneous environments also showed significantly higher investment in photosynthetic biomass and stolon/total biomass, produced longer stolons, and had higher mean leaf size than parents in homogeneous environments. Total biomass and number of offspring ramets were similar in both environments. However, plants in homogeneous environments showed random allocation of offspring ramets to surrounding patches, whereas plants in heterogeneous environments showed preferential allocation of offspring to higher-quality patches. CONCLUSIONS: The results suggest that F. vesca employs physiological and morphological strategies to enable efficient resource foraging in heterogeneous environments and demonstrate the benefits of physiological integration in terms of photosynthetic efficiency. The findings indicate that short-term responses cannot be directly extrapolated to the longer term principally because preferential colonization of high-quality patches means that these patches eventually show reduced quality. This highlights the importance of considering the time factor in experiments examining responses of clonal plants to heterogeneity.     

水分异质性影响两种根茎型克隆植物赖草和假苇拂子茅的水分存储能力

水分在自然系统中呈异质性分布。有关水分异质性对克隆植物生长、形态和生理影响的研究已有大量的工作, 但是水分异质性对克隆植物存储能力, 尤其是水分存储能力影响的研究却十分缺乏。该文将两种根茎型克隆植物赖草(Leymus secalinus)和假苇拂子茅(Calamagrostis pseudophragmites)进行水分异质性和同质性实验处理, 探讨水分异质性对克隆植物水分存储能力、生长和形态的影响。在异质性水分处理下, 两种克隆植物的间隔子、枝和根的含水量均显著增加。两种克隆植物对水分异质性分布的适应策略有所不同, 赖草通过降低单个克隆分株的生长、提高芽的数量以应对水分异质性, 而假苇拂子茅通过增强整个分株种群的地下部分(根状茎、根和芽)生长来应对水分资源的异质性分布。水分储存能力的增强可以提高克隆植物适应水分异质性的能力。     

Do Physiological Integration and Soil Heterogeneity Influence the Clonal Growth and Foraging of <Emphasis Type="Italic">Schoenoplectus pungens</Emphasis>?

Physiological integration and foraging behavior have both been proposed as advantages for clonal growth in heterogeneous environments. We tested three predictions concerning their short- and long-term effects on the growth of the clonal perennial sedge Schoenoplectus pungens (Pers.) Volk. ex Schinz and R. Keller: (1) growth would be greatest for clones with connected rhizomes and on heterogeneous soil, (2) clones would preferentially place biomass in the nutrient-rich patches of a spatially heterogeneous environment, and (3) physiological integration would decrease a clone’s ability to forage. We tested our predictions by growing S. pungens clones for 2 years in an experimental garden with two severing treatments (connected and severed rhizomes) crossed with two soil treatments (homogeneous and heterogeneous nutrient distribution). Severing treatments were only carried out in the first year. As predicted, severing significantly decreased total biomass and per capita growth rate in year one and individual ramet biomass both in year one and the year after severing stopped. This reduction in growth was most likely caused by severing damage, because the total biomass and growth rate in severed treatments did not vary with soil heterogeneity. Contrary to our prediction, total biomass and number of ramets were highest on homogeneous soil at the end of year two, regardless of severing treatment, possibly because ramets in heterogeneous treatments were initially planted in a nutrient-poor patch. Finally, as predicted, S. pungens concentrated ramets in the nutrient-rich patches of the heterogeneous soil treatment. This foraging behavior seemed enhanced by physiological integration in the first year, but any possible enhancement disappeared the year after severing stopped. It seems that over time, individual ramets become independent, and parent ramets respond independently to the conditions of their local microsite when producing offspring, a life-history pattern that may be the rule for clonal species with the spreading “guerrilla” growth form.     

Foraging responses of clonal plants to multi-patch environmental heterogeneity: spatial preference and temporal reversibility

Background and aims

Plant root placement is highly plastic in order to acquire patchily distributed nutrients and to ensure their survival, growth and reproduction. Considering the spatial extension of clonal organs, we selected two clonal plants (Leymus chinensis (Trin.) Tzvel. and Hierochloe glabra Trin.) to determine the spatio-temporal effects of environmental heterogeneity on belowground organs and newly-born ramets.

Methods

Small-scale and multi-patch heterogeneous environments were manipulated by creating four patches filled with different types of soil in a same pot. The four patches were composed of sandy soil, sandy loam, loam soil and humus soil, respectively. Ramet number, bud number, mean spacer length, rhizome length, and biomass allocation within each patch were measured to identify plant foraging responses.

Results

The preferential patch of L. chinensis was humus soil patch which was the highest in nutrient availability, whereas H. glabra preferred to place ramets in sandy loam and loam soil patches. When growing in homogeneous environments, both species randomly rooted their offspring ramets in the four compartments. In heterogeneous environments, foraging responses were detected in ramet placement, aboveground biomass and total rhizome length. However, there were no differences in bud number or belowground biomass among four types of patches in heterogeneous environments, which might suggest that there would be no inter-patch differences in seedling establishment in the next year.

Conclusions

Plants show selective allocation of offspring ramets to preferential patches in the presence of multi-patch environmental heterogeneity. Responses of H. glabra to multi-patch heterogeneity were faster than those of L. chinensis, demonstrating that the foraging patterns are species-specific. Clonal plants can rapidly respond to environmental heterogeneity, whereas foraging responses are potentially reversible over a longer temporal scale.     

根状茎型植物扁秆荆三棱对土壤养分异质性尺度和对比度的生长响应

土壤养分的空间异质性在自然界普遍存在, 而克隆植物被认为能很好地适应和利用土壤养分异质性。尽管尺度和对比度是异质性的两个重要属性, 但有关土壤养分异质性的尺度和对比度及其交互作用对克隆植物生长和分株分布格局影响的研究仍比较缺乏。在一个温室实验中, 根状茎型草本克隆植物扁秆荆三棱(Bolboschoenus planiculmis) (异名扁秆藨草(Scirpus planiculumis))被种植在由高养分斑块和低养分斑块组成的异质性环境中。实验为两种尺度处理(大斑块和小斑块)和两种对比度处理(高对比度和低对比度)交叉组成的4种处理组合。在每个处理中, 高养分和低养分斑块的总面积相同; 在所有4种处理中, 土壤养分的总量也完全相同。无论在整个克隆(植株)水平, 还是在斑块水平, 尺度、对比度及其交互作用对扁秆荆三棱的生物量、分株数、根状茎长和块茎数的影响均不显著。然而, 在斑块水平, 扁秆荆三棱在高养分斑块中的生物量、分株数、根状茎长和块茎数均显著高于低养分斑块, 而在高养分斑块中相邻分株间的距离(间隔物长)小于低养分斑块, 并且这种效应均不依赖于斑块尺度的大小和对比度的高低。因此, 在土壤养分异质性环境中, 扁秆荆三棱可以通过缩短间隔物长, 并可能通过提高根状茎的分枝强度, 把较多的分株和潜在分株放置在养分条件好的斑块中。这种响应格局体现出克隆植物的觅食行为, 有利于整个克隆对异质性资源的吸收和利用。然而, 该实验中的尺度和对比度对扁秆荆三棱分株的放置格局均没有显著效应。作者推测, 在一个更大的斑块尺度和(或)对比度范围内, 扁秆荆三棱对土壤养分异质性的响应可能不同。因此, 下一步的研究应涉及更广泛的尺度和对比度。     

Soil heterogeneity affects ramet placement of Hydrocotyle vulgaris

Aims Soil heterogeneity is common in natural habitats. It may trigger foraging responses (placing more ramets and/or roots in nutrient-rich patches than in nutrient-poor patches) and further affect the growth of plants. However, the impact of soil heterogeneity on competitive interactions has been little tested.Methods We conducted a greenhouse experiment to investigate the effects of soil heterogeneity on intraspecific competition with a stoloniferous herb Hydrocotyle vulgaris. We grew one (without competition) or nine ramets (with competition) of H. vulgaris under a homogeneous environment and two heterogeneous environments differing in patch size (large or small patches). In the heterogeneous treatment, the soil consisted of the same number of nutrient-rich and nutrient-poor patches arranged in a chessboard manner, and in the homogeneous treatment, the soil was an even mixture of the same amount of the nutrient-rich and the nutrient-poor soil.Important findings Irrespective of intraspecific competition, H. vulgaris showed foraging responses to soil heterogeneity in the large patch treatment, e.g. it produced significantly more biomass, ramets, aboveground mass and root mass in the nutrient-rich patches than in the nutrient-poor patches. In the small patch treatment, foraging responses were observed when intraspecific competition was present, but responses were not observed when there was no competition. However, we find a significant effect of soil heterogeneity on neither overall growth nor competitive intensity of H. vulgaris. Our results suggest that foraging responses to soil heterogeneity may not necessarily be adaptive and intraspecific competition may not be influenced by soil heterogeneity.     

Invasive clonal plant species have a greater root-foraging plasticity than non-invasive ones

Clonality is frequently positively correlated with plant invasiveness, but which aspects of clonality make some clonal species more invasive than others is not known. Due to their spreading growth form, clonal plants are likely to experience spatial heterogeneity in nutrient availability. Plasticity in allocation of biomass to clonal growth organs and roots may allow these plants to forage for high-nutrient patches. We investigated whether this foraging response is stronger in species that have become invasive than in species that have not. We used six confamilial pairs of native European clonal plant species differing in invasion success in the USA. We grew all species in large pots under homogeneous or heterogeneous nutrient conditions in a greenhouse, and compared their nutrient-foraging response and performance. Neither invasive nor non-invasive species showed significant foraging responses to heterogeneity in clonal growth organ biomass or in aboveground biomass of clonal offspring. Invasive species had, however, a greater positive foraging response in terms of root and belowground biomass than non-invasive species. Invasive species also produced more total biomass. Our results suggest that the ability for strong root foraging is among the characteristics promoting invasiveness in clonal plants.     

Effects of Spatial Scale of Soil Heterogeneity on the Growth of a Clonal Plant Producing Both Spreading and Clumping Ramets

Soil nutrients are commonly heterogeneously distributed at different spatial scales. Although numerous studies have tested the effects of soil nutrient heterogeneity on growth of clonal plants producing either spreading ramets or clumping ramets, no study has examined the effects on the growth of clonal plants producing both spreading and clumping ramets and how spatial scale affects such effects. To test these effects, clones of Buchloe dactyloides, a stoloniferous clonal plant that produces both clumping and spreading ramets, were grown in six heterogeneous environments with different patch sizes and one homogeneous environment containing the same quantity of nutrients. Total biomass, total number of ramets, number of clumping ramets, number of spreading ramets, spacer length, or root:shoot ratio of the whole plants did not differ significantly among the seven treatments. However, at the patch level there were significant effects of patch size by nutrient level on biomass, number of ramets, number of spreading ramets, and number of clumping ramets, and these four variables were significantly larger in the nutrient-rich patches than in the nutrient-poor patches in the heterogeneous treatment with the largest patch size, but not in the other five heterogeneous treatments with smaller patch sizes. Neither nutrient level nor patch size significantly affected spacer length or root:shoot ratio. Based on our results, we propose that B. dactyloides can efficiently exploit nutrient-rich patches by a plastic response of clumping ramets and spreading ramets at larger spatial scales of soil heterogeneity but not at smaller ones.     

Phenotypic Responses of a Stoloniferous Clonal Plant Buchloe dactyloides to Scale-Dependent Nutrient Heterogeneity

Clonal plants could modify phenotypic responses to nutrients heterogeneously distributed both in space and time by physiological integration. It will take times to do phenotypic responses to modifications which are various in different growth periods. An optimal phenotype is reached when there is a match between nutrient conditions and foraging ability. A single plantlet of Buchloe dactyloides with two stolons was transplanted into heterogeneous nutrient conditions. One stolon grew in homogeneous nutrient patch, while the other cultured in different scales of heterogeneous nutrient patches. As compared to the other nutrient treatment, heterogeneous nutrient treatments with small scale of 25×25 cm resulted in a higher biomass, and larger number of ramets, clumps and stolons in B. dactyloides at both genet and clonal fragment levels. Significant differences of number of ramets, clumps and stolons were detected at the rapid growth stage, but not in the early stage of the experiment. Foraging ability was more efficient in heterogeneous than in homogeneous nutrient conditions as assessed by higher root mass and root to shoot ratio. Different nutrient treatments did not prompt significant differences in internode and root length. Physiological integration significantly increased biomass, but did not influence other growth or morphological characters. These results suggest that physiological integration modifies phenotypic plasticity of B. dactyloides for efficient foraging of nutrients in heterogeneous nutrient conditions. These effects are more pronounced at genet and clonal fragment levels when the patch scale is 25×25 cm. Time is a key factor when phenotypic plasticity of B. dactyloides in heterogeneous nutrient conditions is examined.     

Buffalograss decreases ramet propagation in infertile patches to enhance interconnected ramet proliferation in fertile patches

Buffalograss (Buchloë dactyloides) is known for its low-nutrient tolerance. However, in natural habitats, nutrients are usually patchily distributed. For clonal plants like buffalograss, physiological integration is an important strategy to cope with adverse environmental conditions. In order to examine how integration helps buffalograss to survive in patchy conditions, a greenhouse experiment was conducted for 91 days. Interconnected ramet pairs of stoloniferous buffalograss were planted in two partitioned same-sized containers, and subjected to identical or contrasting nutrient supply. In contrast to normally perceived resource-sharing concepts, results showed that buffalograss genets reduced production of new ramets in nutrient-poor patches promoting at the same time propagation of interconnected ramets in nutrient-rich patches. Ramets in nutrient-rich patches gained significant benefit from heterogeneous treatments, whereas nutrient-poor ramets performed even worse than in uniform low-nutrient treatment. Younger ramets developed more biomass than elder ramets with the same amounts of nutrient supply under homogeneous treatment, while elder ramets were more tolerant when nutrients were scarce. Heterogeneity had a particular strong effect on stolons and new ramet production in nutrient-rich patches. Rooted ramets in nutrient-poor patches suffered from a by-pass of nutrients to interconnected ramets on nutrient-rich substrate that probably resulted from different transpiration rates. We conclude that this resource-sharing strategy is advantageous for buffalograss to concentrate more ramets in fertile patches, and facilitate the survivorship of more buffalograss ramets in adverse environments with uneven nutrient supply.     

土壤养分水平影响绢毛匍匐委陵菜匍匐茎生物量投资

为研究匍匐茎草本植物对基质养分供应水平的生物量分配格局的可塑性,在一盆栽实验中对绢毛匍匐委陵菜(Potentilla reptans var. sericophylla)进行了8种不同的养分处理。绢毛匍匐委陵菜植株生物量、匍匐茎数、分株数以及匍匐茎节间长在中等养分条件下最大。随土壤养分的降低,绢毛匍匐委陵菜对叶片和叶柄的生物量投资减小,而对根系的生物量投资增加。在中等养分条件下,绢毛匍匐委陵菜对匍匐茎的生物量投资倾向于最大,而在更高或更低的养分条件下倾向于减少。此生物量分配格局与de Kroon和Schieving的模型模拟结果相符合,结果表明在中等资源水平下增加对匍匐茎的生物量投资是克隆植物增加资源获取的对策之一。     

Intermediate growth forms as a model for the study of plant clonality functioning: an example with root sprouters

In this contribution we want to show that growth forms intermediate between non-clonal and clonal plants can be used to ask questions about the functional ecology of clonality. We discuss this idea on plants sprouting adventitiously from roots and accomplishing clonal growth via root spacers. Based on extensive literature dealing with growth forms of root sprouting plants, we characterise forms functionally intermediate between clonal root-sprouters and non-clonal plants. We delimit them according to their potential ability to form adventitious shoots and horizontal roots. By reviewing experimental work with root sprouters, we identify the most important triggering factors and developmental constraints influencing these intermediate forms plant age, life-history mode and life-history stage. Using this information we ask questions about the importance of root sprouting in (1) conditions of unpredictable disturbance, where root-sprouting ability may be viewed as a tool for vegetative regeneration, and in (2) temporarily and spatially heterogeneous environment, where foraging by roots may serve as a way of exploiting patchy resources.     

沙丘坡面异质性小生境中准噶尔无叶豆对水分条件变化的响应

以多年生克隆植物准噶尔无叶豆(Eremosparton songoricum(Litv.)Vass.)为材料,选择河边(A种群)和沙漠腹地(B种群)两个沙丘,研究从沙丘底部至顶部,沿着水分条件连续变化的梯度,准噶尔无叶豆在分株种群和克隆片段水平的形态变化特征,以期能揭示其在异质性小生境内利用水分资源的对策,并为准噶尔无叶豆的资源保护、培育和利用提供有意义的参考.研究发现:①在分株种群水平,A种群分株高度及地上部生物量显著高于B种群,而B种群地下部(根)的生物量则显著高于A种群;②在克隆片段水平,随着沙丘底部至顶部,A种群与B种群克隆片段高度和地上生物量都减小,而分株密度都增加,但升高或降低的强度不同;A种群根的生物量和长度增加,主要是水平的位于地下0～10 cm层面的直径10mm以下的根长度增加,而B种群根的生物量减小,但长度却在增加,主要是水平的位于地下0～10 cm层面的直径6mm以下的细根长度增加.水平细根的长度增加,更利于无性系进行广泛觅食,同时促进无性系尽快越过不利生境斑块和提高分株在有利生境中的生长概率.结果表明,准噶尔无叶豆对沙丘坡面水分条件连续变化的异质性小生境存在分株种群及克隆片段两个等级的可塑性响应,并通过可塑性变化适应了沙丘坡面水分条件的分异.     

Effects of Heterogeneous Competitor Distribution and Ramet Aggregation on the Growth and Size Structure of a Clonal Plant

Spatially heterogeneous distribution of interspecific competitors and intraspecific aggregation of offspring ramets may affect the growth and size structure of clonal plant populations, but these have been rarely studied. We conducted a greenhouse experiment in which we grew a population of eight offspring ramets (plants) of the stoloniferous clonal plant Hydrocotyle vulgaris aggregately or segregately in two homogeneous treatments with or without a competing grass Festuca elata and a heterogeneous treatment with a patchy distribution of the grass. In patchy grass treatments, H. vulgaris produced markedly more biomass, ramets and stolons in open patches (without grasses) than in grass patches, but displayed lower size variations as measured by coefficient of variation of biomass, ramets and stolons among the eight plants. In open areas, H. vulgaris produced statistically the same amounts of biomass and even more stolons and showed higher size variations in patchy grass treatments than in open (no grass) treatments. In grass areas, H. vulgaris grew much worse and displayed higher size variations in patchy grass treatments than in full grass treatments. Ramet aggregation decreased the growth of H. vulgaris in open treatments and in both open and grass patches in patchy grass treatments, but had little effect in full grass treatments. Ramet aggregation had little effect on size variations. Therefore, heterogeneous distribution of competitors can affect the growth and size structure of clonal plant populations, and ramet aggregation may decrease population growth when they grow in open environments or heterogeneous environments with a patchy distribution of interspecific competitors.     

Differential foraging for resources, and structural plasticity in plants

Although ecologists have spent much effort in analysing the foraging behaviour of animals, the study of plants as foraging organisms is a relatively unexplored subject. There is often, however, much greater potential for analysis of foraging behaviour in plants than in animals. Unlike most animals, many plant species leave permanent or semi-permanent records of their foraging activities because their resource-acquiring structures (primarily leaves and roots), persist for a considerable time, as also do the structures (trunks, branches, stolons, runners or rhizomes) which enable leaves or roots to be projected into particular positions in the habitat. In addition, plant ecologists are not burdened with the difficulties associated with determining how changes in foraging behaviour affect fitness in animals(1), because plant mass (or, in the case of clonal species, number of ramets produced), is usually closely correlated with fitness.     

Effects of moisture availability on clonal growth in bamboo Pleioblastus maculata

The effects of moisture availability on clonal growth and biomass investment in the bamboo Pleioblastus maculata were investigated over a four-year period by transplanting Pleioblastus maculata clones into soils with different levels of moisture availability in the field. The results showed that: (1) The higher the moisture availability, the greater the total biomass of P. maculata clones. Although fewer culms are produced at the higher moisture levels, mean tiller biomass is greater. (2) Under different levels of moisture availability, obvious differences in the total rhizome length (p < 0.01), spacer length (p < 0.05) and the sizes of bamboo culms (height, p < 0.01; diameter, p < 0.01) were observed. Thus, the higher the moisture availability, the shorter the rhizomes and the larger ramets. (3) In microhabitats with low moisture availability, bamboo allocated more biomass to underground organs, which promotes elongation of rhizomes and increases root production, thereby helping to capture underground resources essential to growth. In microhabitats of high moisture availability, the biomass is primarily allocated to the aboveground growth of ramets. (4) We suggest that soil moisture availability effects the foraging strategies of bamboo, that bamboo plants growing with low moisture availability produce longer rhizomes (that is, more, although shorter, spacers) with more biomass allocation than plants in high moisture and have a better ability to forage to increase the probability of locating adequate moisture patches. Also, longer length distance between shoots (that is, longer spacers) in high soil moisture than in low is adapted to avoid intense competition from faster growing aboveground growth in high moisture patches.     

Root growth and plant biomass in <Emphasis Type="Italic">Lolium perenne</Emphasis> exploring a nutrient-rich patch in soil

We investigated soil exploration by roots and plant growth in a heterogeneous environment to determine whether roots can selectively explore a nutrient-rich patch, and how nutrient heterogeneity affects biomass allocation and total biomass before a patch is reached. Lolium perenne L. plants were grown in a factorial experiment with combinations of fertilization (heterogeneous and homogeneous) and day of harvest (14, 28, 42, or 56 days after transplanting). The plant in the heterogeneous treatment was smaller in its mean total biomass, and allocated more biomass to roots. The distributions of root length and root biomass in the heterogeneous treatment did not favor the nutrient-rich patch, and did not correspond to the patchy distribution of inorganic nitrogen. Specific root length (length/biomass) was higher and root elongation was more extensive both laterally and vertically in the heterogeneous treatment. These characteristics may enable plants to acquire nutrients efficiently and increase the probability of encountering nutrient-rich patches in a heterogeneous soil. However, heterogeneity of soil nutrients would hold back plant growth before a patch was reached. Therefore, although no significant selective root placement in the nutrient-rich patch was observed, plant growth before reaching nutrient-rich patches differed between heterogeneous and homogeneous environments.     

Influence of sediment fertility on morphological variability of Vallisneria spiralis L.

One homogeneous and three heterogeneous nutrient enrichment treatments were imposed to investigate the growth responses of Vallisneria spiralis L. Morphological features of V. spiralis differed significantly between different nutrient patches. Roots elongated in nutrient-poor patches, and the specific root length (SRL) also increased significantly. Stolon length, diameter and leaf length and width increased significantly in nutrient-rich patches. Total plant biomass of V. spiralis grown in the homogeneous and three heterogeneous treatments on average were 2.9, 3.0, 3.9 and 2.3 fold higher than that grown in the control treatment. Number of ramets per clone was significantly higher in the heterogeneous treatments than in the homogeneous treatment. In three varying heterogeneous treatments, ramet biomass in nutrient-rich patches was 2.7, 4.3 and 3.0 fold higher than in nutrient-poor patches; however, ramet number was not affected by sediment nutrients, resulting in bigger ramets in nutrient-rich patches. The biomass allocation established adaptive plasticity to heterogeneous environments. The maximum value of biomass allocation to underground parts reached 16% in nutrient-rich patches, whereas the minimum value of underground parts reached 20% in nutrient-poor patches. Results demonstrate that clonal V. spiralis can maintain itself preferentially in favourable nutrient-rich sediments, whereas nutrient-poor conditions could be escaped by plastic biomass allocation.     

Biomass allocation in a clonal vine: Effects of intraspecific competition and nutrient availability

Biomass allocation to roots, rhizomes, runners and climbing stems (i.e. twining axis and attached leaves) was studied inCalystegia sepium L., a clonal vine. In an experiment which took 2 months, nutrient availability (low and high) and intraspecific competition (none, shoot root and both shoot and root) were manipulated. Under low nutrients the highest biomass of climbing stems was found in plants with shoot competition; the lowest was found in plants with both shoot and root competition. Total biomass under high nutrients was also greatest in plants with shoot competition. Thus, plants benefited from climbing up a shared stake rather than separate stakes. Larger plants allocated a higher proportion of biomass to runners in the nutrient-poor environment than in the nutrient-rich environment. This behaviour may increase the chance of finding nutrient-rich patches in the neighbourhood of the mother plant in a heterogeneous environment.     

The effects of physiological integration on biomass partitioning in plant modules: an experimental study with the stoloniferous herb Glechoma hederacea

Morphological and physiological plasticity are crucial attributes enabling plants to acquire resources from heterogeneous habitats. Although physiological integration can modify biomass partitioning in modules, especially when connected modules experience different conditions, its ecological importance has been largely overlooked. This experiment examined its effects on above- and belowground biomass partitioning by modules in the stoloniferous herb Glechoma hederacea. We studied how biomass allocation to roots by younger ramets was affected by connection to older ramets, and by nutrient conditions. A lower proportion of biomass was allocated to roots by younger ramets growing under low nutrient (LN) conditions when connected to older ramets in high nutrient (HN) conditions than when they were isolated, demonstrating localised modification of biomass partitioning due to physiological integration. The proportion of biomass allocated to roots by younger ramets was also lower when connected to older ramets in HN conditions than when connected to older ramets in LN conditions. Thus, the effect of integration on biomass partitioning depended on the nutrient conditions experienced by connected ramets. Such changes in biomass partitioning would result in more extensive stolon growth, and greater lateral displacement of new ramets. Understanding the ecological implications of phenotypic plasticity in plants will require further examination of the effects of physiological integration when connected modules experience contrasting growing conditions. This study demonstrates that such integration affects the biomass allocation strategy of connected ramets, enhancing resource acquisition in heterogeneous habitats. The widespread success of clonality in many communities is likely to be strongly promoted by this characteristic.