水曲柳落叶松根系之间的相互作用研究

 通过水曲柳(Fraxinus mandshurica)和落叶松(Larix gmelinii)两个树种苗木纯栽和混栽试验,在温室条件下研究两树种根系之间的相互作用。结果表明,水曲柳与落叶松混栽使水曲柳细根和粗根生物量比纯栽分别增加47％和46％,叶和茎生物量分别增加30％和48％,同时改变了水曲柳地下部分与地上部分比率,促进了水曲柳细根和叶协调生长。而混栽的落叶松细根和粗根生物量比纯栽分别减少19％和35％左右,叶和茎生物量减少22％和34％,但是落叶松地下部分和地上部分各生物量组分变化不存在显著差异。混栽改变了水曲柳根系的空间分布和根系的形态,导致根系密度和根长度分别增加47％和34％,有利于水曲柳吸收较多的养分和水分,提高其竞争效率。落叶松起到改变土壤环境作用。     

不同灌溉量对绿洲-荒漠过渡带多枝柽柳幼苗根系生长和分布的影响

为了探讨绿洲-荒漠过渡带上受损柽柳群落幼苗适宜生长的土壤水分条件,在塔南策勒绿洲外围设置A（不灌溉）、B（适度灌溉）、C（充分灌溉）3个处理的田间试验,于7～10月份考察了各处理柽柳幼苗粗根（>2 mm）和细根（<2 mm）生物量积累、空间分布等变化特征。结果显示:（1）随着灌溉量的增加,柽柳根系生物量积累不断增加,且在生长季末（10月份）增速最大,同时细根占总根重的比例从20.5%上升到29.8%,显著提高了幼苗吸收养分和水分的能力。（2）干旱胁迫（处理A）下柽柳幼苗根系的下扎深度大,但水平根幅的扩展却有限;适度灌溉（处理B）的生物量较处理A大,但小于处理C;根系扎根深度表现为处理A>处理B>处理C,各指标均处于中间状态;水分条件最好（处理C）时虽然垂直扎根深度小,但水平根幅最大,可以更有效利用浅层土壤资源。（3）柽柳幼苗大量根系集中在0～40 cm的土壤表层,而且水分条件越好,这种集中趋势越明显;根系生物量随土壤深度的增加呈递减变化,细根的根长密度也有相似变化,但细根的比根长变化规律不明显,这可能与各层土壤微观环境的变化有关。研究表明,灌溉对柽柳幼苗根系的生长和分布有显著影响,充分灌溉下柽柳幼苗根系生物量积累最多且空间分布最大,该灌溉量有利于柽柳幼苗根系的正常生长和合理分布。     

水曲柳和落叶松细根形态及母根与子根比例关系

细根（直径〈2mm）的分枝是根系重要的结构特征,不同根序等级的细根在养分和水分吸收、C的消耗和寿命方面具有较大的差异,定量研究各根序等级之间的比例关系对认识细根死亡的顺序具有重要的理论意义。根据Pregitzer等2002年提供的方法,研究了17年生水曲柳（Fraxinus mandshurica Rupr．）和落叶松（Larix gmelinii Rupr．）人工纯林1-5级细根的直径、长度、比根长、生物量和数量。结果表明,两树种细根中1级根序的数量占总根系数量80％-90％,它们直径小、长度短、比根长高。随着根序等级（1级-5级）的增加细根直径增粗和长度增加、比根长减小。细根的数量和生物量在上下土层的分布受土壤资源有效性的影响。水曲柳5级根序-2级根序之间母根与子根的数量关系是1：3,落叶松是1：2-3。2级根序与1级根序之间母根与子根的数量关系,水曲柳是1：10—12,落叶松是1：8。如果当年生长的1级细根当中保持1：3的比例,将有65％-75％的1级细根死亡,占根系总数的55％～65％,总长度的40％-50％,以及总生物量的20％-30％。     

水曲柳根系生物量、比根长和根长密度的分布格局

采用连续钻取土芯法在生长季内对东北林业大学帽儿山实验林场17年生水曲柳人工林根系取样,研究水曲柳不同直径根系现存生物量、比根长和根长密度及垂直分布状况.结果表明,水曲柳人工林根系总生物量为1 637.6 g·m^-2,其中活根生物量占85%,死根占15%.在活根生物量当中,粗根(直径5～30 mm)占的比例最高（69.95%）,其次为活细根（直径<1 mm,13.53%）,小根(1～2 mm)和中等直径的根(2～5 mm)比例较小（分别为7.21%和9.31%）.直径<1 mm活细根的比根长为32.20 m·g^-1,直径5～30 mm粗根的比根长为0.08 m·g^-1.单位面积上活根的总长度为6 602.54 m·m^-2,其中直径<1 mm的细根占92.43%,其它直径等级则不到活根总长度的8%.直径<1 mm的细根生物量与根长密度具显著线性关系（R²=0.923）,但与比根长无显著相关关系（R²=0.134）.     

水曲柳根系生物量、比根长和根长密度的分布格局

采用连续钻取土芯法在生长季内对东北林业大学帽儿山实验林场17年生水曲柳人工林根系取样,研究水曲柳不同直径根系现存生物量、比根长和根长密度及垂直分布状况.结果表明,水曲柳人工林根系总生物量为1 637.6 g·m^-2,其中活根生物量占85%,死根占15%.在活根生物量当中,粗根(直径5～30 mm)占的比例最高（69.95%）,其次为活细根（直径<1 mm,13.53%）,小根(1～2 mm)和中等直径的根(2～5 mm)比例较小（分别为7.21%和9.31%）.直径<1 mm活细根的比根长为32.20 m·g^-1,直径5～30 mm粗根的比根长为0.08 m·g^-1.单位面积上活根的总长度为6 602.54 m·m^-2,其中直径<1 mm的细根占92.43%,其它直径等级则不到活根总长度的8%.直径<1 mm的细根生物量与根长密度具显著线性关系（R²=0.923）,但与比根长无显著相关关系（R²=0.134）.     

科尔沁沙地赤松和樟子松根系生物量分配与构型特征

沙地赤松(Pinus densiflora)在科尔沁沙地南缘区已有50年的引种历史,但其生长表现和根系生物量分配与构型特征还很少被报道。本研究以同龄(40 a)樟子松为对照,在生长指标测定基础上,采用分层分段全挖法采集根系,对沙地赤松不同径级根生物量分配规律进行研究,测定根长、连接数量、平均连接长度等指标,同时计算分形维数和分形丰度。结果表明:与樟子松相比,沙地赤松具有较大生长量和生物量,其根生物量显著高于樟子松,是其1.96倍,细根(直径≤0.2 cm)生物量更显著高于樟子松,是其4.76倍;沙地赤松根系生物量占总生物量的29.0%,细根生物量占总根系生物量的1.1%,细根长度占根总长度的44.3%;樟子松根系生物量占总生物量的25.6%,细根生物量占总根系生物量的0.4%,细根长度占根总长度的28.8%;从根系垂直分布看,沙地赤松地下0~180 cm均有细根分布,且40~180 cm范围内细根生物量占总细根生物量的65.2%;樟子松几乎全部细根分布于0~100 cm范围内,此范围细根生物量占总细根生物量的99.2%,且0~40 cm土层细根生物量占63.4%;虽然两树种根系平均连接长度没有显著差异,但沙地赤松细根及部分中根(0.2~2.0 cm)连接数量显著高于樟子松;沙地赤松根系分形维数为1.548±0.251,是樟子松(1.293±0.190)的1.2倍,并且分形丰度是樟子松的1.3倍;与樟子松相比,沙地赤松根系具有较强的吸收能力,能够利用较大范围的深层水分和养分,根系分支多,拓扑结构更加复杂。     

异质养分环境中一年生分蘖草本黍根系的生长特征

为揭示黍（Panicum miliaceum L.)根系对异质养分环境的生长反应,作研究了黍根系从起始斑块向目标斑块水平生长时,时始斑块和目标斑块养分水平根生长的影响,就低养分起始珏块而言,粗根生物量,粗根长度,粗根表面积和细极长度在高养分目标斑块中的分配比例均小于其在低养分目标斑块中的分配比例,而细根长度及其密度,细根表面积指及其密度的变化恰好相反,就高养分起始斑块而言,高养分目标斑块的细根长度,细根长度密度,细根表面积指数和细根表面积密均不于低养分目标斑块,而粗根对目标斑块中养分状的反应不明显。当黍根系从桢的起始斑块进入不同的目标斑块后,目标斑块的养分状况对细根生物量及其分配无影响,而显影响细根长度和表现积,这指示细根是通过长度和表面积可塑性而不是生物量变化响应目标斑块中的养分差异。     

红砂幼苗根系形态特征和水分利用效率对土壤水分变化的响应

为探讨干旱与半干旱区受损红砂种群幼苗适宜生长的土壤水分条件,采用盆栽方法,研究了红砂幼苗在充分灌溉(FI)、适度灌溉(MI)、干旱处理(DT)3个水分处理下根系形态和水分利用效率的变化特征。结果表明:(1)红砂幼苗根系形态因水分条件和根序的不同而各异;随灌溉量的减少红砂幼苗根系直径和根体积均表现为FIMIDT,但干旱处理促进了根系的伸长生长和比表面积和比根长增加,根系形态的可塑性是红砂幼苗获取水分适应干旱环境的重要策略之一。(2)随根序的升高,各处理水平下红砂幼苗根长、比根长均显著减少,而其根直径和体积却显著增加,表明红砂幼苗根系内部具有高度的形态异质性。(3)与FI处理相比,MI和DT处理下红砂幼苗根系总生物量分别增加了50.00%、19.23%,但MI和DT处理却显著降低了红砂幼苗地上生物量,特别是叶片生物量下降幅度最大,分别降低了62.15%、83.28%,导致根冠比随灌溉量的减少而逐渐增加。(4)干旱处理显著提高了红砂幼苗的水分利用效率。研究认为,在灌溉量减少的情况下,红砂幼苗可通过根长、根系表面积和体积、直径等形态变化来优化其空间分布构型,以调节植株对水分的利用,提高水分利用效率。     

桢楠种源幼苗细根形态和生物量研究

为了解桢楠(Phoebe zhennan)不同种源细根形态和生物量分配的差异,采用全根调查的方法,对桢楠自然分布区13个种源2.5年生幼苗的细根形态和生物量进行了研究。结果表明,桢楠种源间各级细根的平均直径、总根长和表面积差异显著,在种源内细根的平均直径随根序的增加而增加,但根序间总根长和表面积差异规律不明显。根序生物量分配随根序增加而增加,1~4级根生物量分配分别为6.33%、14.47%、25.03%和54.17%。通过综合评价,以HT、LF、ES和WC种源的根系最优,具有较高的生长潜力。     

西北干旱区灌草型白刺-芨芨草群落根系分布与互作

本文以宁夏沙湖一个典型的干旱区灌草型白刺(Nitraria tangutorum)—芨芨草(Achnatherum splendens)群落为研究对象,将该群落自植物聚生丛冠层覆盖区至丛间裸地依次划分为冠盖区、过渡区和空旷区3个微生境分区,通过比较两优势种的根系生物量密度以及根系消弱系数(反映根系总体埋深)在这3个分区中的分布特征,研究了干旱胁迫环境下两物种的根系分布与共存机制。结果表明:两物种的总根与细根均集中分布于冠盖区,但芨芨草根系生物量密度自冠盖区至空旷区的下降幅度更大;白刺总根及细根生物量密度重心自冠盖区至空旷区逐渐下移,而芨芨草的总根和细根生物量密度重心在空旷区较过渡区有所抬升;另外,白刺细根生物量密度在冠盖区10~30 cm土层中最高,而芨芨草在0~10cm层最高;各微生境分区内两物种的根系消弱系数均具有显著性(或接近于显著性)差异。这暗示了白刺与芨芨草的根系分布存在明显的生态位分离现象。"地上聚生,地下分离"的共生方式可能是白刺与芨芨草共同适应干旱环境的生长策略。     

Positive tree diversity effect on fine root biomass: via density dependence rather than spatial root partitioning

The importance of species richness to ecosystem functioning and services is a central tenet of biological conservation. However, most of our theory and mechanistic understanding is based on diversity found aboveground. Our study sought to better understand the relationship between diversity and belowground function by studying root biomass across a plant diversity gradient. We collected soil cores from 91 plots with between 1 and 12 aboveground tree species in three natural secondary forests to measure fine root (≤ 2 mm in diameter) biomass. Molecular methods were used to identify the tree species of fine roots and to estimate fine root biomass for each species. This study tested whether the spatial root partitioning (species differ by belowground territory) and symmetric growth (the capacity to colonize nutrient-rich hotspots) underpin the relationship between aboveground species richness and fine root biomass. All species preferred to grow in nutrient-rich areas and symmetric growth could explain the positive relationship between aboveground species richness and fine root biomass. However, symmetric growth only appeared in the nutrient-rich upper soil layer (0–10 cm). Structural equation modelling indicated that aboveground species richness and stand density significantly affected fine root biomass. Specifically, fine root biomass depended on the interaction between aboveground species richness and stand density, with fine root biomass increasing with species richness at lower stand density, but not at higher stand density. Overall, evidence for spatial (i.e. vertical) root partitioning was inconsistent; assumingly any roots growing into deeper unexplored soil layers were not sufficient contributors to the positive diversity–function relationship. Alternatively, density-dependent biotic interactions affecting tree recruitment are an important driver affecting productivity in diverse subtropical forests but the usual root distribution patterns in line with the spatial root partitioning hypothesis are unrealistic in contexts where soil nutrients are heterogeneously distributed.     

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.     

Nutrient foraging traits in 10 co-occurring plant species of contrasting life forms

1 Responses to spatial heterogeneity of soil nutrients were tested in 10 plant species that differ in life form and successional status, but which co-occur in the South Carolina coastal plain. The morphological responses of the root system were tested by assessing scale (represented by root mass and root length densities), precision (preferential proliferation of roots in nutrient-rich patches compared with less fertile patches) and discrimination (ability to detect and proliferate within the richest patches when patches vary in nutrient concentration). We also investigated sensitivity (growth benefits gained as spatial heterogeneity of nutrients increases, measured as total biomass).
2 Ten individuals of each species were grown in pots under four treatments that had differing nutrient distribution but the same overall nutrient addition. Plants were harvested when roots reached pot edge.
3 We observed high variation between species in scale, precision and sensitivity. No significant discrimination responses were observed, although greatest root mass density occurred at intermediate fertility levels for all species.
4 We rejected the hypothesis that scale and precision are negatively correlated. Indeed, in herbaceous species alone, scale and precision were positively correlated.
5 Sensitivity was not closely related to precision, indicating that proliferation of roots in fertile patches does not always yield growth benefits in heterogeneous soils. Further, some sensitive species had very low precision, suggesting that a positive growth response in heterogeneous environments may be related to plasticity in physiology or root life span, rather than morphology.
6 Plant life form was not correlated with precision or sensitivity. However, scale of response was greater in herbs than in woody plants, possibly because the two life forms develop root systems at different rates.     

Soil nutrient status determines how elephant utilize trees and shape environments

1. Elucidation of the mechanism determining the spatial scale of patch selection by herbivores has been complicated by the way in which resource availability at a specific scale is measured and by vigilance behaviour of the herbivores themselves. To reduce these complications, we studied patch selection by an animal with negligible predation risk, the African elephant. 2. We introduce the concept of nutrient load as the product of patch size, number of patches and local patch nutrient concentration. Nutrient load provides a novel spatially explicit expression of the total available nutrients a herbivore can select from. 3. We hypothesized that elephant would select nutrient-rich patches, based on the nutrient load per 2500 m(2) down to the individual plant scale, and that this selection will depend on the nitrogen and phosphorous contents of plants. 4. We predicted that elephant would cause more adverse impact to trees of lower value to them in order to reach plant parts with higher nutrient concentrations such as bark and root. However, elephant should maintain nutrient-rich trees by inducing coppicing of trees through re-utilization of leaves. 5. Elephant patch selection was measured in a homogenous tree species stand by manipulating the spatial distribution of soil nutrients in a large field experiment using NPK fertilizer. 6. Elephant were able to select nutrient-rich patches and utilized Colophospermum mopane trees inside these patches more than outside, at scales ranging from 2500 down to 100 m(2) . 7. Although both nitrogen and phosphorus contents of leaves from C. mopane trees were higher in fertilized and selected patches, patch choice correlated most strongly with nitrogen content. As predicted, stripping of leaves occurred more in nutrient-rich patches, while adverse impact such as uprooting of trees occurred more in nutrient-poor areas. 8. Our results emphasize the necessity of including scale-dependent selectivity in foraging studies and how elephant foraging behaviour can be used as indicators of change in the availability of nutrients.     

土壤养分空间异质性与植物根系的觅食反应

植物在长期进化过程中,为了最大限度地获取土壤资源,对养分的空间异质性产生各种可塑性反应．包括形态可塑性、生理可塑性、菌根可塑性等．许多植物种的根系在养分丰富的斑块中大量增生,增生程度种间差异较大,并受斑块属性(斑块大小、养分浓度)、营养元素种类和养分总体供应状况的影响．植物还通过调整富养斑块中细根的直径、分枝角、节问距以及空间构型来实现斑块养分的高效利用．根系的生理可塑性及菌根可塑性可能在一定程度上影响其形态可塑性．生理可塑性表现为处于不同养分斑块上的根系迅速调整其养分吸收速率,从而增加单位根系的养分吸收,对在时间上和空间上变化频繁的空间异质性土壤养分的利用具有重要意义,可在一定程度上弥补根系增生反应的不足．菌根可塑性目前研究较少,一些植物种的菌根代替细根实现在富养斑块中的增生．菌根增生的碳投入养分吸收效率较高、根系增生对增加养分吸收的作用较复杂,取决于养分离子在土壤中的移动性能以及是否存在竞争植物;对植物生长(竞争能力)的作用因种而异,一些敏感种由此获得生长效益,而其它一些植物种受影响较小．植物个体对土壤养分空间异质性反应能力和生长差异,影响其在群落中的地位和命运,最终影响群落组成及其结构．     

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.     

Root morphological and physiological plasticity of perennial grass species and the exploitation of spatial and temporal heterogeneous nutrient patches

Root morphological and physiological characteristics of four perennial grass species were investigated in response to spatial and temporal heterogeneous nutrient patches. Two species from nutrient-rich habitats (i.e. Holcus lanatus and Lolium perenne) and two species from nutrient-poor habitats (i.e. Festuca rubra and Anthoxanthum odoratum) were included in the study. Patches were created by injecting equal amounts of nutrient solution into the soil either on one location (i.e. spatial heterogeneity) or on several, alternating locations (i.e. temporal heterogeneity) within the pot. The consequences of changes in root morphology and the implications for the exploitation of the nutrient patches by individual plants were quantified by the amount of ¹⁵N captured from the enriched patches. The effects of nutrient heterogeneity on the acquisition of nutrients by species were determined by comparing the total nitrogen and phosphorus acquisition of the species in the two heterogeneous habitats with the total nitrogen and phosphorus acquisition in a homogeneous treatment. In this homogeneous treatment the same amount of nutrient solution was supplied homogeneously over the soil surface. The experiment lasted for 27 days and comprised one harvest. In response to the spatial enrichment treatment, all species produced significantly more root biomass within the enriched patch. The magnitude of the response was similar for species from nutrient-rich and nutrient-poor habitats. In contrast to this response of root biomass, root morphology, including specific root length, branching frequency and mean lateral root length was not affected by the treatments. In response to the temporal enrichment treatment, all species were able to increase the nitrogen uptake rate per unit of root biomass. The species from nutrient-poor habitats had, on average, higher uptake rates per unit root biomass than the species from nutrient-rich habitats, but the magnitude of the response did not differ between the species. These results question the general validity of the assumptions that root foraging characteristics differ among species from nutrient-rich and nutrient-poor habitats. As a result of these root responses, all species captured an equal amount of ¹⁵N from the spatial and temporal enriched nutrient patches and all species acquired significantly more nitrogen in the heterogeneous treatments than in homogeneous treatment. Hence, the ability to exploit local and temporal nutrient heterogeneity does not appear to differ between species from nutrient-rich and nutrient-poor habitats, but is achieved by these species in different ways. The ecological implications of these differences are discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

The effects of shading and N status on root proliferation in nutrient patches by the perennial grass Agropyron desertorum in the field

Competition for light can affect exploitation of spatially heterogeneous soil resources. To evaluate the influence of shoot status on root growth responses in nutrient-rich soil patches, we studied the effects of shading and whole-plant nitrogen status on root growth in N-enriched and nonenriched patches by mature Agropyron desertorum plants growing in the field with below-ground competition. Roots in enriched patches had greater length to weight ratios (specific root length, SRL), indicating increased absorptive surface areas, compared with roots in control patches. Increased SRL was due to increased production and length of higher order laterals rather than morphological changes in roots of the same branching order. Although the pattern of root growth rates in patches was the same for shaded and unshaded plants, the magnitude of this response to enriched patches was damped by shading. Root relative growth rates (RGR) in N-enriched patches were reduced by more than 50% by short-term shading treatments (60% reduction in photosynthetic flux density), while root RGR in unenriched patches was unaffected by shading. Unexpectedly, plants with higher nitrogen status had greater root RGR in enriched patches than plants that had not received nitrogen supplement, again with no detectable effect on root RGR in the unenriched patches. Therefore, while both shading and plant N status affected the ability of roots to exploit enriched patches by proliferation, there was no stimulation or suppression of root growth in the unenriched, control patches. Thus, plants already under competitive pressure above ground for light and below ground for nutrients should be less able to rapidly respond to opportunities presented in nutrient patches and pulses.     

Habitat selection in spatially heterogeneous environments: a test of foraging behaviour in the clonal submerged macrophyte Vallisneria spiralis

1. To test whether clonal macrophytes can select favourable habitats in heterogeneous environments, clonal fragments of the stoloniferous submerged macrophyte Vallisneria spiralis were subjected to conditions in which light intensity and substratum nutrients were patchily distributed. The allocation of biomass accumulation and ramet production of clones to the different patches was examined. 2. The proportion of both biomass and ramet number of clones allocated to rich patches was significantly higher than in poor patches. The greatest values of both clone and leaf biomass were produced in the heterogeneous light treatment, in which clones originally grew from light‐rich to light‐poor patches, while clones produced the most offspring ramets in the treatments with heterogeneous substratum nutrients. Similarly, root biomass had the highest values in nutrient‐rich patches when clones grew from nutrient‐rich to nutrient‐poor patches. 3. The quality of patches in which parent ramets established significantly influenced the foraging pattern. When previously established in rich patches, a higher proportion of biomass was allocated to rich patches, whereas a higher proportion of ramet number was allocated to rich patches when previously established in poor patches. 4. Results demonstrate that the clonal macrophyte V. spiralis can exhibit foraging in submerged heterogeneous environments: when established under resource‐rich conditions V. spiralis remained in favourable patches, whereas if established in adverse conditions it could escape by allocating more ramets to favourable patches.