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.     

Heterogeneous nutrient supply promotes maize growth and phosphorus acquisition: additive and compensatory effects of lateral roots and root hairs

Background and AimsRoot proliferation is a response to a heterogeneous nutrient distribution. However, the growth of root hairs in response to heterogeneous nutrients and the relationship between root hairs and lateral roots remain unclear. This study aims to understand the effects of heterogeneous nutrients on root hair growth and the trade-off between root hairs and lateral roots in phosphorus (P) acquisition.MethodsNear-isogenic maize lines, the B73 wild type (WT) and the rth3 root hairless mutant, were grown in rhizoboxes with uniform or localized supply of 40 (low) or 140 (high) mg P kg⁻¹ soil.ResultsBoth WT and rth3 had nearly two-fold greater shoot biomass and P content under local than uniform treatment at low P. Significant root proliferation was observed in both WT and rth3 in the nutrient patch, with the WT accompanied by an obvious increase (from 0.7 to 1.2 mm) in root hair length. The root response ratio of rth3 was greater than that of WT at low P, but could not completely compensate for the loss of root hairs. This suggests that plants enhanced P acquisition through complementarity between lateral roots and root hairs, and thus regulated nutrient foraging and shoot growth. The disappearance of WT and rth3 root response differences at high P indicated that the P application reduced the dependence of the plants on specific root traits to obtain nutrients.ConclusionsIn addition to root proliferation, the root response to a nutrient-rich patch was also accompanied by root hair elongation. The genotypes without root hairs increased their investment in lateral roots in a nutrient-rich patch to compensate for the absence of root hairs, suggesting that plants enhanced nutrient acquisition by regulating the trade-off of complementary root traits.     

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.     

Root Growth of the Annual Tillering Grass Panicum miliaceum in Heterogeneous Nutrient Environments (in English)

To study growth responses of the roots of Panicum miliaceum L. to heterogeneous supply of nutrients. The authors analyzed the effects of the nutrient levels in both original patches (O) and destination patches (D) on the root growth of P. miliaceum when its roots were allowed to extend from original patch into destination patch. When the nutrient levels in the original patches were low, coarse root biomass ratio (coarse root biomass in the D/total coarse root biomass), coarse root length ratio (coarse root length in the D/total coarse root length), coarse root surface area ratio (coarse root surface area in the D/total coarse root surface area) and fine root length ratio (fine root length in the D/total fine root length) were greater in the destination patches with lower nutrient levels than in the destination patches with higher nutrient levels, while fine root length, fine root length density, fine root surface index, and fine root surface area density were smaller in the former than in the latter. When the nutrient levels in the original patches were high, fine root length, fine root length density, fine root surface area index and fine root surface density were greater in the destination patches with lower nutrient levels than in the destination patches with higher nutrient levels, coarse roots did not respond to the nutrient levels in the destination patches significantly. When the roots extended from the original patches with the same nutrient level into the destination patches with contrasting nutrient levels, fine root biomass and its percentage allocation did not respond to the nutrient levels in the destination patches significantly, whereas both root length and root surface area did. This indicates that the fine roots of P. miliaceum responded to difference in nutrient supply by plasticity in their length and surface area, rather than in their root biomass.     

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。     

马尾松种源在异质养分环境中的觅养行为差异

选择广东信宜、福建武平、广西岑溪3个不同磷效率特性的马尾松种源,构建同质和异质两种养分环境开展盆栽实验,研究马尾松搜寻利用异质分布养分的获取机制及不同种源觅养行为差异。结果表明,与同质营养环境相比,异质营养环境中马尾松种源具有较高的苗高、地径生长量、较强光合速率和干物质生产能力。研究证实了根系形态可塑性和生理可塑性在马尾松获取异质分布养分中的重要性。马尾松可通过在富养斑块中须侧根的大量增生、对N、P、K等元素的有效吸收提高其觅养能力。马尾松在拓殖富养斑块的初期主要依靠新生侧根的增加和侧根的延长,在拓殖一段时期之后则主要靠新生侧根的生成和须根数量、须根密度的增加来搜寻异质分布养分。异质养分环境中的根系具有较高的养分吸收效率主要缘由在富养斑块中对N、P、K大量的吸收。综合比较分析认为,在3个参试种源中广西岑溪和福建武平种源在异质营养环境中拓殖富养斑块和觅养能力较强,广东信宜种源拓殖和觅养能力相对较弱。     

Soil nutrient heterogeneity and competitive ability of three grass species (<Emphasis Type="Italic">Festuca ovina,Arrhenatherum elatius</Emphasis> and <Emphasis Type="Italic">Calamagrostis epigejos</Emphasis>) in experimental conditions

We studied the effects of differences in root growth and nutrient pool on the competitive ability of Festuca ovina (short grass), Arrhenatherum elatius and Calamagrostis epigejos (tall grasses) grown in monocultures and in mixtures of homogeneous and heterogeneous environments during two growing seasons. Analysis of variance revealed a significant effect of plant species on nutrient concentrations in above-ground biomass and of substrate type on contents of N, K, Ca, Mg in biomass. The ANOVA also confirmed the significant effect of competitive environment on the concentration of N, K in above-ground biomass. In heterogeneous environments, both tall grasses (in competition with F. ovina) were able to produce more roots in the nutrient-rich patches and to accumulate more nitrogen in plant tissues, which was associated with higher yield of their above-ground biomass. Thus, the relative competitive ability for nutrients of both tall grasses was higher than that of F. ovina. This competitive ability of A. elatius to C. epigejos increased in heterogeneous treatments.     

Root morphological plasticity and nutrient acquisition of perennial grass species from habitats of different nutrient availability

We studied the root foraging ability and its consequences for the nutrient acquisition of five grass species that differ in relative growth rate and that occur in habitats that differ widely in nutrient availability. Foraging responses were quantified, based on the performance of the plants in homogeneous and heterogeneous soil environments of the same overall nutrient availability. Although all species tended to produce a significantly higher root length density in a nutrient-rich patch, this response was significant only for the faster-growing species. The increased root length density resulted from small, though not significant, changes in root biomass and specific root length. The effectiveness of root proliferation was determined by quantifying the total amount of nutrients (N and P) accumulated by the plants over the course of the experiment. Plants acquired more N in a heterogeneous environment than in a homogeneous environment, although the total nutrient availability was the same. The ability to acquire nutrients (N or P) in the heterogeneous environment was not related to the ability of species to increase root length density in response to local nutrient enrichment. In contrast to other studies, our results suggest that the role of morphological plasticity of roots in acquiring patchily distributed resources is limited. Possible reasons for this discrepancy are discussed. Received: 11 September 1997 / Accepted: 28 February 1998     

Growth Consequences of Soil Nutrient Heterogeneity for two Old-field Herbs,Ambrosia artemisiifoliaandPhytolacca americana, Grown Individually and in Combination

Plant species can respond to small scale soil nutrient heterogeneityby proliferating roots or increasing nutrient uptake kineticsin nutrient-rich patches. Because root response to heterogeneitydiffers among species, it has been suggested that the distributionof soil resources could influence the outcome of interspecificcompetition. However, studies testing how plants respond toheterogeneity in the presence of neighbours are lacking. Inthis study, individuals of two species,Phytolacca americanaL.andAmbrosia artemisiifoliaL. were grown individually and incombination in soils with either a homogeneous or heterogeneousnutrient distribution. Above-ground biomass of individuallygrown plants of both species was greater when fertilizer waslocated in a single patch than when the same amount of fertilizerwas distributed evenly throughout the soil. Additionally, bothspecies proliferated roots in high-nutrient patches.A. artemisiifoliaexhibitedlarger root:shoot ratios, increased nitrogen depletion fromnutrient patches, and a higher growth rate thanP. americana,suggestingA. artemisiifoliais better suited to find and rapidlyexploit nutrient patches. In contrast to individually grownplants, soil nutrient distribution had no effect on final above-groundplant biomass for either species when grown with neighbours,even though roots were still concentrated in high nutrient patches.This study demonstrates that increased growth of isolated plantsas a consequence of localized soil nutrients is not necessarilyan indication that heterogeneity will affect interspecific encounters.In fact, despite a significant below-ground response, soil nutrientheterogeneity was inconsequential to above-ground performancewhen plants were grown with neighbours.Copyright 1999 Annalsof Botany Company Phytolacca americana, pokeweed,Ambrosia artemisiifolia, ragweed, nutrient heterogeneity, root proliferation, plasticity, foraging, nutrient patches.     

Effect of heterogeneous distribution of nutrients on root growth, ABA content and drought resistance of wheat plants

Heterogeneous distribution of nutrients influenced morphology of wheat roots; they were shorter and had more laterals at the site of high nutrient content. The roots outside the nutrient-rich patch were longer and penetrated deeper in the soil than those of the plants supplied with the same quantity of nutrients but having them homogeneously distributed. The ABA content of roots sampled from the site of high nutrient content was greater than in those elsewhere. It is suggested that this might be important for inhibition of their extension growth and stimulation of laterals in the nutrient rich patch. Heterogeneously distributed nutrients beneficially influenced accumulation of ¹⁵N and assimilation of CO₂ by the plants, enabling an increased supply of nutrients to the parts of the root system beyond the nutrient rich patch and sustained their rapid extension. Enhanced penetration of roots into the deeper soil layers could contribute to the observed increase in drought resistance of plants grown with localised placement of fertilisers under field conditions.     

Nitrate and phosphate availability and distribution have different effects on root system architecture of Arabidopsis

Plant root systems can respond to nutrient availability and distribution by changing the three-dimensional deployment of their roots: their root system architecture (RSA). We have compared RSA in homogeneous and heterogeneous nitrate and phosphate supply in Arabidopsis. Changes in nitrate and phosphate availability were found to have contrasting effects on primary root length and lateral root density, but similar effects on lateral root length. Relative to shoot dry weight (DW), primary root length decreased with increasing nitrate availability, while it increased with increasing phosphate supply. Lateral root density remained constant across a range of nitrate supplies, but decreased with increasing phosphate supply. In contrast, lateral root elongation was suppressed both by high nitrate and high phosphate supplies. Local supplies of high nitrate or phosphate in a patch also had different effects. Primary root growth was not affected by a high nitrate patch, but growth through a high phosphate patch reduced primary root growth after the root left the patch. A high nitrate patch induced an increase in lateral root density in the patch, whereas lateral root density was unaffected by a high phosphate patch. However, both phosphate- and nitrate-rich patches induced lateral root elongation in the patch and suppressed it outside the patch. This co-ordinated response of lateral roots also occurs in soil-grown plants exposed to a nutrient-rich patch. The auxin-resistant mutants axrl, axr4 and aux1 all showed the wild-type lateral root elongation responses to a nitrate-rich patch, suggesting that auxin is not required for this response.     

供磷水平及方式对杉木幼苗根系生长和磷利用效率的影响

为研究杉木幼苗根系生长、形态学指标及养分利用效率对土壤磷素异质分布的响应规律,选择杉木种子园单株采种培育的半同胞家系实生幼苗为研究对象,采用室内沙培控磷盆栽试验,设计低浓度供磷(8 mg/kg KH_2PO_4)、正常供磷(16 mg/kg KH_2PO_4)和高浓度供磷(32 mg/kg KH_2PO_4)3个供磷水平,每个供磷水平分别采用2种供磷方式(局部供磷和均匀供磷)进行根部施磷。结果表明:(1)从供磷水平来看,低浓度供磷下的杉木根长、根系生物量、根冠比、根系及全株的磷素利用效率均显著大于正常供磷和高浓度供磷,而根平均直径相反;随着供磷水平的提高,杉木苗高和地上部生物量无显著差异,而比根长表现出逐渐降低的趋势。(2)从供磷方式来看,局部供磷处理的杉木苗高、根长、根系表面积、比根长、地上部生物量、根系及全株的磷素利用效率均显著大于均匀供磷处理,而根平均直径和根冠比则相反。总体上,低浓度局部供磷处理下杉木可明显增强其根系的形态可塑性,从而优化根系在养分异质土壤里的空间分布,并通过提高根系磷素利用效率以维持地上部的正常生长。     

Heterogeneity in spatial P-distribution and foraging capability by Zea mays: effects of patch size and barriers to restrict root proliferation within a patch

• Background and Aims Localized proliferation of rootsin nutrient-enriched patches seems to be an adaptive responsein many plants, but its function is still debatable. To understandthe efficiency and limitation of foraging behaviour, the impactof patch size and the presence or absence of a barrier to rootproliferation within phosphorus (P)-enriched patches was examined. • Methods In pots filled with P-poor soil, six treatmentsof heterogeneous P supply were prepared: three patch sizes withor without a root barrier between patches. In addition, a homogeneousP supply treatment was also prepared. Irrespective of thesetreatments, each pot received the same total amount of P. Maize(Zea mays) was grown in each pot for 45 d in a greenhouse. • Key Results P content and biomass were greatest in plantsgrown in the largest patch due to successful root proliferation,and were higher in the presence of a root barrier. Interestingly,plants preferentially developed adventitious nodal roots projectingfrom the stem into the P-enriched soil, particularly in thelargest patch with a root barrier. Removal of the barrier reducedthe P-uptake capacity per unit root surface area or volume inP-enriched patches, revealing that the P-uptake capacity perroot can be suppressed even in P-rich soil if other portionson the root axis encounter P-poor conditions. • Conclusions The results suggest that the efficiency ofroot morphological plasticity is largely determined by the sizeof the P-enriched patch. Furthermore, the results imply a novelaspect of P-uptake physiology that roots in heterogeneous Pcannot demonstrate their potential capacity, as would be observedin roots encountering P continuously; this effect is probablymediated by an internal root factor.     

Interactive effects of soil nutrient heterogeneity and belowground herbivory on the growth of plants with different root foraging traits

Aims

Plants with precise root foraging patterns can proliferate roots preferentially in nutrient-rich soil patches. When nutrients are distributed heterogeneously, this trait is often competitively advantageous in pot experiments but not field experiments. We hypothesized that this difference is due to belowground herbivory under field conditions.

Methods

We performed pot experiments using seedlings of Lolium perenne (a more precise root foraging species) and Plantago lanceolata (a less precise root foraging species). The experiment had a two-way factorial randomized block design, with nutrient distribution pattern (homogeneous or heterogeneous) and belowground herbivore (present or absent) as the two factors. Each pot contained one seedling of each species.

Results

With no herbivore present, plant biomass was smaller in the heterogeneous nutrient treatment than in the homogeneous treatment in P. lanceolata, but not in L. perenne. Under homogeneous nutrient distribution, plant biomass was lower in both species with a herbivore present than with no herbivore. Under heterogeneous nutrient distribution, biomass reduction due to herbivory occurred only in L. perenne.

Conclusions

Roots of the precise root foraging species were grazed more under the heterogeneous nutrient distribution, suggesting that the herbivore more efficiently foraged for roots in nutrient-rich soil patches.     

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.     

Root proliferation,nitrate inflow and their carbon costs during nitrogen capture by competing plants in patchy soil

The responses of roots to nitrogen- and phosphorus-rich patches of soil include proliferation of laterals and stimulation of nutrient inflow (uptake rate per unit root length) within the patch. Nitrate uptake from an N-rich patch is thereby maximised and, perhaps, compensates for an uneven supply of nitrate to the whole root system. Paradoxically, the often weak correlation between root length density and N uptake found in experiments on single plants and crop monocultures suggests that root proliferation in patches has only a minor compensatory influence on N capture. This paradox was resolved when it was realised that localised root proliferation during inter-specific competition for nitrate can lead to a strong association between root length density and nitrate capture. Here, a simple model of inter-specific competition is used to estimate the stimulation in inflow required in one plant to match the N capture of a competitor that responds only by root proliferation, and to estimate associated carbon costs. The model predicts that nitrate inflow must increase proportionally more than root length density to achieve the same N capture. For example, the N capture possible with a 10% increase in root length density can be matched by increasing N inflow by anything from 20% to 20-fold, depending on the initial conditions: the faster the rate of change in root length density, the greater the required relative increase in inflow. In those terms, proliferation would seem the better option, but one that may be more costly in terms of its carbon requirement.     

Variation in resource acquisition and utilization traits between native and invasive perennial forbs

Understanding the functional traits that allow invasives to outperform natives is a necessary first step in improving our ability to predict and manage the spread of invaders. In nutrient-limited systems, plant competitive ability is expected to be closely tied to the ability of a plant to exploit nutrient-rich microsites and use these captured nutrients efficiently. The broad objective of this work was to compare the ability of native and invasive perennial forbs to acquire and use nutrients from nutrient-rich microsites. We evaluated morphological and physiological responses among four native and four invasive species exposed to heterogeneous (patch) or homogeneous (control) nutrient distribution. Invasives, on average, allocated more biomass to roots and allocated proportionately more root length to nutrient-rich microsites than did natives. Invasives also had higher leaf N, photosynthetic rates, and photosynthetic nitrogen use efficiency than natives, regardless of treatment. While these results suggest multiple traits may contribute to the success of invasive forbs in low-nutrient environments, we also observed large variation in these traits among native forbs. These observations support the idea that functional trait variation in the plant community may be a better predictor of invasion resistance than the functional group composition of the plant community.     

Mycorrhizal respiration: implications for global scaling relationships

Most plant species form mycorrhizas, yet these are neglected by plant physiologists. One consequence of this neglect is reduced ability to predict plant respiration, because respiration rate (R) in mycorrhizal roots might be higher than in non-mycorrhizal roots owing to increased substrate availability associated with enhanced nutrient uptake, coupled with increased respiratory product demand. Other predictions include that mycorrhizal colonization will affect scaling of R with tissue nitrogen concentrations; that mycorrhizal and non-mycorrhizal root R differ in their response to nutrient supply; and that the impact of colonization on R is related to fungal biomass. Failure to examine properly the role of colonization in determining root R means that current interpretations of root and soil respiration data might be flawed.     

Carbon allocation to shoots and roots in relation to nitrogen supply is mediated by cytokinins and sucrose: Opinion

In this paper we firstly show some general responses of biomass partitioning upon nitrogen deprivation. Secondly, these responses are explained in terms of allocation of carbon and nitrogen, photosynthesis and respiration, using a simulation model. Thirdly, we present a hypothesis for the regulation of biomass partitioning to shoots and roots.Shortly after nitrogen deprivation, the relative growth rate (RGR) of the roots generally increases and thereafter decreases, whereas that of the shoot decreases immediately. The increased RGR of the root and decreased RGR of the shoot shortly after a reduction in the nitrogen supply, cause the root weight ratio (root weight per unit plant weight) to increase rapidly.We showed previously that allocation of carbon and nitrogen to shoots and roots can satisfactorily be described as a function of the internal organic plant nitrogen concentration. Using these functions in a simulation model, we analyzed why the relative growth rate of the roots increases shortly after a reduction in nitrogen supply. The model predicts that upon nitrogen deprivation, the plant nitrogen concentration and the rate of photosynthesis per unit plant weight rapidly decrease, and the allocation of recently assimilated carbon and nitrogen to roots rapidly increases. Simulations show that the increased relative growth rate of the root upon nitrogen deprivation is explained by decreased use of carbon for root respiration, due to decreased carbon costs for nitrogen uptake. The stimulation of the relative growth rate of the root is further amplified by the increased allocation of carbon and nitrogen to roots. Using the simple relation between the plant nitrogen concentration and allocation, the model describes plant responses quite realistically.Based on information in the literature and on our own experiments we hypothesize that allocation of carbon is mediated by sucrose and cytokinins. We propose that nitrogen deprivation leads to a reduced cytokinin production, a decreased rate of cytokinin export from the roots to the shoot, and decreased cytokinin concentrations. A reduced cytokinin concentration in the shoot represses cell division in leaves, whereas a low cytokinin concentration in roots neutralizes the inhibitory effect of cytokinins on cell division. A reduced rate of cell division in the leaves leads to a reduced unloading of sucrose from the phloem into the expanding cells. Consequently, the sucrose concentration in the phloem nearby the expanding cells increases, leading to an increase in turgor pressure in the phloem nearby the leaf's division zone. In the roots, cell division continues and no accumulation of sugars occurs in dividing cells, leading to only marginal changes in osmotic potential and turgor pressure in the phloem nearby the root's cell division zone. These changes in turgor pressure in the phloem of roots and sink leaves affect the turgor pressure gradients between source leaf-sink leaf and source leaf-root in such a way that relatively more carbohydrates are exported to the roots. As a consequence RWR increases after nitrogen deprivation. This hypothesis also explains the strong relationship between allocation and the plant nitrogen status.     

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.