资源交互斑块性生境中两种不同分枝型匍匐茎植物的克隆内分工

以青藏高原和四川盆地过渡带两种不同分枝型匍匐茎植物野草莓 (Fragaria vesca)和过路黄 (Lysimachia christinae)为对象 ,研究它们在高光照低养分斑块和低光照高养分斑块组成的资源交互斑块性生境中的克隆内分工。结果显示 ,与资源的空间同质性处理 (I)和 (II)相比 ,资源的空间异质性处理 (III)和 (IV)中野草莓和过落黄的近端、远端和整个克隆片段的生物量和分株数均获得显著增加。生长在低光高养条件下的远端分株 ,若与高光低养的近端分株相连 ,相比连接到低光高养的近端分株 ,它们分配更多的生物量到地下部分 ;生长在高光低养条件下的远端分株 ,若与低光高养的近端分株相连 ,相比连接到高光低养的近端分株 ,它们分配更多的生物量到地上部分 ;生长在高光低养条件下的近端分株 ,若与低光高养的远端分株相连 ,相比连接到高光低养的远端分株 ,它们分配更多的生物量到地上部分。实验结果表明 ,资源交互斑块性生境中野草莓和过路黄均发生了克隆内分工。通过克隆内分工 ,克隆植物能有效的利用异质性分布的资源 ,缓解资源交互斑块性分布对克隆植物生长的不利影响     

不同程度南方菟丝子寄生对入侵植物三叶鬼针草生长的影响

生物入侵严重影响了生态系统的结构和功能。菟丝子属植物可以有效地抑制入侵植物的生长,恢复本地群落,是一种有效的生物防治剂。本文以本地寄生植物南方菟丝子（Cuscuta australis）和三叶鬼针草（Bidens pilosa L.）为研究对象,采用不同数量的南方菟丝子茎段（长度15cm）寄生入侵植物三叶鬼针草来形成不同的寄生程度（寄生I、II、III分别由1、2、3段南方菟丝子茎段寄生）,分析不同程度的寄生对寄主三叶鬼针草补生长的影响,并探讨寄生植物对寄主补偿生长的诱导。南方菟丝子寄生34d之后,三种程度的寄生均可显著抑制三叶鬼针草生长。寄生III处理组的南方菟丝子生物量显著大于寄生I和寄生Ⅱ处理组。不同寄生强度对三叶鬼针草生长的影响具有一定的差异。寄生II和寄生III的三叶鬼针草叶生物量、茎生物量、地上生物量、地下生物量、总生物量、株高与冠幅均显著低于寄生I,但相互之间不存在显著性差异;寄生II的三叶鬼针草的叶面积、根长和根体积均显著低于寄生I,寄生III的叶面积和根长高于寄生II,但与寄生I及寄生II之间不存显著性差异,寄生III的根体积显著高于寄生II,但与寄生I之间不存在显著性关异。与对照相比,不同程度寄生下三叶鬼针草的NAR和RGR极显著降低。寄生II的三叶鬼针草的RGR显著低于寄生I,寄生III的三叶鬼针草的RGR高于寄生II,但与寄生I及寄生II之间均不存在显著性差异。寄生II与寄生III的三叶鬼针草的NAR显著低于寄生I组,但相互之间不存在显著性差异。除寄生I处理组外,寄生II与寄生III处理导致三叶鬼针草与南方菟丝子的累积生物量均显著低于对照。寄生II处理组的累积生物量要低于寄生I处理组,寄生III处理组的累积生物量要高于寄生II处理组,但显著低于对照组,表明寄生III处理组的三叶鬼针草产生了低补偿生长。由于南方菟丝子寄生显著抑制入侵植物的生物量,因此南方菟丝子仍具有防治入侵植物的潜力。     

放牧对小嵩草草甸生物量及不同植物类群生长率和补偿效应的影响

基于小嵩草(Kobresia parva)草甸连续2 a的牦牛放牧试验,研究了暖季和冷季放牧草场地上地下生物量及其分配规律、不同植物类群的绝对生长率生长率,探讨了放牧制度和放牧强度对不同植物类群补偿效应的影响。结果表明,随着放牧强度的增加地上总生物量呈减小趋势,放牧强度对暖季草场地上总生物量的影响极显著（P?0.01）,对冷季草场地上总生物量的影响不显著（P?0.05）;两季放牧草场各土壤层地下生物量随放牧强度的增加呈明显下降趋势,放牧强度对暖季放牧各土壤层地下生物量的影响显著（P?0.05）,对冷季放牧各土壤层地下生物量的影响不显著（P?0.05）;冷季放牧草场牧草生长季地下生物量与地上生物量的比值随放牧强度的增大而减小,暖季放牧草场对照区地下生物量与地上生物量的比值低于轻度放牧和中度放牧、高于重度放牧;暖季放牧草场各放牧处理不同植物类群均存在超补偿生长,但莎草科和禾本科植物的超补偿生长在8月份,阔叶植物的超补偿生长发生在6月和7月份,禾本科植物的超补偿生长效应强于莎草科植物和阔叶植物,轻度和中度放牧的补偿效应更明显;冷季放牧下不同植物类群也存在超补偿生长,但补偿效应不明现。因此,暖季适度（轻、中度）放牧利用更有利于产生超补偿生长,而重度利用对植被的稳定产生潜在的不利影响。     

放牧格局和生境资源对矮嵩草（Kobresia humilis）分株生物量分配和补偿性生长的影响

为研究密集型克隆植物对放牧扰动和生境资源变化的生物量分配和补偿生长响应特性,验证克隆植物的觅食模型和3个有关植物个体补偿反应的假说(①放牧优化假说,grazing optimization hypothesis,GOH;②反应连续谱假说,continuum of responses hypothesis,CRH;③增长率模型,growth rate model,GRM),在具有不同放牧利用格局和土壤养分水平的高寒矮嵩草草甸(Ⅰ.畜圈草地:重度放牧、资源丰富;Ⅱ.牧道草地:中度放牧、资源贫乏;Ⅲ.封育草地:不放牧、资源贫乏)中通过设置扣笼/无扣笼样方对其建群种矮嵩草(Kobresia humilis)进行了研究.结果表明不论当年解除家畜放牧与否,春季采摘率越高(畜圈草地),分株生物量向生长的投入越少,向贮藏器官的投入越多.繁殖分配在中度采摘下最高(牧道草地),扣笼内外分株各部分的生物量分配无差异.矮嵩草分株在中度采摘×资源贫乏条件下产生了超补偿响应,在重度采摘×资源丰富条件下为等量补偿,重度采摘导致分株密度显著减少.生物量分配格局与觅食模型的预测不符.补偿生长响应特性证实了GOH和GRM的预测,但与CRH的预测不符.这说明在研究地区放牧扰动格局对克隆植物矮嵩草分株的生物量分配和补偿生长具有重要影响,适度放牧利用更利于引起超补偿,而重度利用可能会对该种群的长期保持产生不利影响.     

根部水淹和土壤养分提升对三峡库区消落带水蓼生长和繁殖特性的影响

水淹和土壤养分是影响三峡库区消落带植物生长的主要环境因子。消落带不同高程的植物长期经历不同的淹水强度和土壤养分条件。该研究假设同一物种来自于消落带不同高程的植株可能产生性状分化, 从而对根部淹水和土壤养分变化具有不同的生长和繁殖响应策略。为了验证以上假设, 选取在三峡库区消落带高低高程均广泛分布的物种水蓼(Polygonum hydropiper)为研究对象, 采集自然种群的种子。在温室同质园条件下, 研究了根部水淹和土壤养分提升对高低高程水蓼植株生长和繁殖特性的影响。研究结果表明根部水淹显著或趋于显著降低了水蓼植株功能叶的叶长、叶宽、总分枝数、叶生物量、花生物量和总生物量; 低养分处理显著或趋于显著降低了水蓼植株的总节数、总分枝数、根生物量、花生物量和总生物量, 表明根部水淹和低土壤养分对水蓼的生长和繁殖能力具有抑制作用。同时, 根部水淹和土壤养分的交互作用显著影响植株的根生物量, 表明根部水淹条件下高土壤养分更有利于植株根生物量的积累。高高程植株的根生物量和叶生物量显著或趋于显著高于低高程植株, 而低高程植株的始花时间早于高高程植株, 且繁殖分配也显著高于高高程植株, 表明高低高程水蓼植株对资源的分配策略不同。该研究结果表明水蓼的生长和繁殖特性受根部水淹和土壤养分共同限制, 但对根部水淹条件下高土壤养分生境具有较好的适应性; 同时, 低高程植株可以通过调整其生长和繁殖特性以提高对所处生境胁迫的适应性。     

镉胁迫对桂花生长和养分积累、分配与利用的影响

桂花(Osmanthus fragrans var. thunbergii)是长江流域镉污染地区普遍栽植兼具绿化、观赏和净化环境等重要价值的园林树种之一。为了解镉胁迫条件下桂花生长适应特性, 采用盆栽试验研究了不同镉浓度处理下(CK: 0 mg·kg^-1; I: 25 mg·kg^-1; II: 50 mg·kg^-1; III: 100 mg·kg^-1; IV: 200 mg·kg^-1)一个生长季节内一年生桂花生物量生产、生物量分配格局以及C、N、P积累、分配与利用特征。植物各器官生物量生产及C、N和P积累量均表现出随镉处理浓度的增加而降低的趋势, 较高浓度镉处理(II、III、IV)明显抑制了桂花的生物量生产、C、N和P的积累, 显著改变了生物量及其C、N和P积累量的分配格局, 但相对较低浓度镉处理(I)对桂花生物量生产以及C、N和P的积累与分配特征影响并不显著。一定浓度的镉胁迫处理(I、II、III)表现出提高桂花N的利用效率而降低P的利用效率的趋势, 但重度镉胁迫(IV)均降低了桂花N和P的利用效率。结果表明桂花具有一定的抗镉胁迫能力, 但较高程度的镉胁迫显著影响了桂花生长及养分格局。     

基于根系形态可塑性的空心莲子草克隆分工特征

资源在空间和时间上不均匀分布现象往往形成资源异质性斑块,克隆植物凭借强大的侧向生长能力占据广阔空间,分株间的生理连接促进了其对异质性生境的适应。克隆分株首先通过资源获取结构的功能特化来提高从各种资源富养斑块中的养分获取,然后通过克隆整合作用实现分株间的养分传输,这种功能特化和资源共享模式被称为‘分工’。该文以入侵克隆植物空心莲子草(Alternanthera philoxeroides)为研究对象,研究其根系对资源异质性分布的形态可塑性响应;通过调节光照强度和土壤养分来实现资源的异质性分布,共设置4个处理:1近端分株高光低养—远端分株高光低养(HL-HL),2近端分株低光高养—远端分株低光高养(LH-LH),3近端分株高光低养—远端分株低光高养(HL-LH),4近端分株低光高养—远端分株高光低养(LH-HL);使用WinRHIZO Pro软件分析相关根系指标,SPSS 18.0单因素方差(one-way ANOVA)分析方法分析异质性条件对近、远端分株以及整个克隆片段的影响。结果表明:异质性斑块中经历高光低养的分株分配更多的生物量到地上部分,经历低光高养的分株分配更多的生物量到地下部分,空心莲子草通过调整对地上和地下部分的生物量分配比例实现了克隆分工;异质性斑块中,生长在富养斑块中的空心莲子草分株根系有更高的根生物量、根长、根表面积、根体积以及分枝系数等,表明空心莲子草分株根系通过对异质性斑块的形态可塑性变化提高了土壤养分的吸收能力。由此可见,空心莲子草通过对资源获取结构的功能特化提高了其资源吸收能力,这可能是其具强入侵能力的重要原因。     

不同生境下空心莲子草响应模拟昆虫采食的生长和化学防御策略

外来植物往往可以入侵多种生境并受到多种昆虫的采食,而不同生境条件将可能会影响这些入侵植物对昆虫采食的防御策略。以入侵我国的克隆植物——空心莲子草为研究对象,分别选择生长在水生生境、水陆两栖生境和陆生生境中的无性个体(分株),通过50%去叶处理模拟昆虫采食,分析不同生境下空心莲子草对模拟昆虫采食处理的生长及化学防御响应的差异。模拟昆虫采食处理显著抑制了陆生生境、水陆两栖生境以及水生生境下空心莲子草的根、茎、叶和总生物量,但对3种生境下空心莲子草的生物量分配(根冠比、根生物量分配、茎生物量分配和叶生物量分配)均无显著影响。陆生生境下空心莲子草根、茎和总生物量显著高于水陆两栖生境和水生生境,根冠比显著低于水陆两栖生境和水生生境。模拟昆虫采食处理显著降低了空心莲子草的木质素含量,而对单宁和总酚含量影响不显著。生境对木质素含量无显著影响,但陆生生境下空心莲子草单宁含量显著高于水陆两栖生境和水生生境,且总酚含量显著高于水陆两栖生境,表明陆生生境中空心莲子草具有更强的防御能力。空心莲子草木质素含量与总生物量无显著相关性,但在模拟采食情况下,其总酚含量与总生物量呈显著负相关,而无论模拟昆虫采食处理存在与否,空心莲子草单宁含量与总生物量均呈显著正相关。因此,空心莲子草存在昆虫介导的生长和化学防御之间的权衡,在昆虫采食的情况下可通过减少生长来增加对化学防御物质的投入,但生境对空心莲子草这种生长-防御权衡的影响十分有限。     

异质性生境中匍匐茎草本野草莓(Fragaria vesca)的克隆内资源共享

研究了不同海拔高度(1800和3900m)的匍匐茎克隆植物野草莓(Fragaria vesca)种群对光照和养分资源斑块性分布生境的响应。结果表明:与资源的空间同质性处理(Ⅰ)和(Ⅱ)相比,资源的空间异质性处理(Ⅲ)和(Ⅳ)中2个种群的野草莓的近端、远端分株部分和整个克隆片段的生物量和分株数均明显增加。当近端分株部分经历低光高养,而与其相连的远端分株部分经历高光低养时,相比于整个克隆片段都处于低光高养的同质性生境,来自2个海拔的种群的近端分株部分都会增加对根的生物量分配;当近端分株部分经历高光低养,而与其相连的远端分株部分经历低光高养时,相比于整个克隆片段都处于低光高养的同质性生境,来自2个海拔的种群的近端分株部分都会减少对根的生物量分配,远端分株部分也被观察到类似的生物量分配格局。相比于高光低养的同质性生境,当与低光高养的远端分株部分相连时,经历高光低养的近端分株部分有更大的叶面积;相比于低光高养的同质性生境,当与低光高养近端分株部分相连时,经历高光低养的远端分株部分有更大的叶面积。结果表明,野草莓在资源交互斑块性生境中发生了克隆内分工,克隆内分工有利于克隆植物对异质性资源的利用,对克隆植物在资源斑块性分布生境中的生存和生长具有重要的意义。     

不同生境中鬼针草(Bidens pilosa)碳氮磷化学计量特征及其营养利用策略

该研究通过野外采样和实验室测定的方法,研究了三种生境中鬼针草叶和根碳(C)、氮(N)、磷(P)化学计量特征及其与土壤养分的相关性。结果表明:(1)鬼针草的生境具有不同的资源水平。三种生境的土壤全氮(TN)和速效氮贫乏,有机碳(C)和全磷(TP)较充裕;生境Ⅰ土壤TN含量显著低于其它两个生境,生境II土壤TP含量显著低于其它两个生境,生境III土壤TN、TP含量均大于其它两个生境。(2)不同生境的鬼针草对磷(P)分配策略不同。低N生境的鬼针草叶片P含量根P含量,P较多地分配到植物体地上部分;N、P含量较高的生境中鬼针草根P含量叶P含量,P更多地分配到地下部分。(3)不同生境的鬼针草其地上部分和根的生长速率不同。低N生境的鬼针草叶片N/P和C/P值小,植物体具有较高的相对生长速率,具有地上生长竞争优势;低P生境的植物叶片N/P和C/P值大,植物体具有较慢的相对生长速率;高N、高P生境中根N/P和C/P值小,根具有较高的生长速率,保证了鬼针草的地下生长竞争优势。(4)鬼针草叶片N/P和根N/P之间呈现不显著的负相关关系,植物地上部分和地下部分为异速生长。不同生境的鬼针草具有不同的营养利用和分配策略,保证了植物强大的竞争力和入侵性。     

Plant compensatory growth: a conquering strategy in plant–herbivore interactions?

We present a theoretical analysis that considers the phenotypic trait of compensatory growth ability in a context of population dynamics. Our model depicts a system of three interactors: herbivores and two different plant types referred to as ordinary and compensating. The compensating plant type has the ability to increase its intrinsic rate of biomass increase as a response to damage. This compensatory growth ability is maintained at the expense of a reduced growth rate in the absence of damage, where the ordinary plant type has the higher growth rate. Analysis of this system suggests that, even though a compensatory capacity of this kind will not imply an increase in equilibrium plant density, it will give a competitive advantage in relation to other plants, in the presence of a sufficiently efficient herbivore. Invasion of compensating plants into a population of non-compensating plants is facilitated by a high compensatory growth ability and a high intrinsic rate of plant biomass increase. Conversely, an ordinary plant can invade and outcompete a compensating plant when the herbivore is characterised by a relatively low attack rate, and/or when plant intrinsic growth rate is decreased.     

Compensatory growth and oxidative stress in a damselfly

Physiological costs of compensatory growth are poorly understood, yet may be the key components in explaining why growth rates are typically submaximal. Here we tested the hypothesized direct costs of compensatory growth in terms of oxidative stress. We assessed oxidative stress in a study where we generated compensatory growth in body mass by exposing larvae of the damselfly Lestes viridis to a transient starvation period followed by ad libitum food. Compensatory growth in the larval stage was associated with higher oxidative stress (as measured by induction of superoxide dismutase and catalase) in the adult stage. Our results challenge two traditional views of life-history theory. First, they indicate that age and mass at metamorphosis not necessarily completely translate larval stress into adult fitness and that the observed physiological cost may explain hidden carry-over effects. Second, they support the notion that costs of compensatory growth may be associated with free-radical-mediated trade-offs and not necessarily with resource-mediated trade-offs.     

Compensatory growth of a submerged macrophyte (<Emphasis Type="Italic">Vallisneria spiralis</Emphasis>) in response to partial leaf removal: effects of sediment nutrient levels

Many plants mitigate damage due to loss of tissues through compensatory growth, yet their compensatory abilities vary depending on physical and environmental conditions. We conducted an outdoor experiment using a 2 × 2 factorial experimental design (leaf damage and nutrient level), in order to evaluate the compensatory growth response of Vallisneria spiralis (a submerged macrophyte widely distributed in China) to partial leaf removal in two nutrient regimes. Our results reveal that under both high- and low-nutrient conditions, V. spiralis exhibited substantial compensatory growth response to partial leaf removal via accelerated growth rates, with significantly greater compensatory abilities observed in the high-nutrient sediments. These observations suggest that V. spiralis has a strong compensatory ability to partial leaf removal, providing arguably one of the major mechanisms for the coexistence of this plant with herbivores, in particular, in eutrophic freshwater ecosystems (e.g. Lake Taihu).     

刈割、施肥和浇水对矮嵩草补偿生长的影响

通过对青海海北高寒矮嵩草(Kobresia humilis)草甸进行为期3年的野外控制试验, 研究了刈割(留茬1 cm、3 cm及不刈割)、施肥(2.5 g·m^-2尿素+ 0.6 g·m^-2磷酸二胺、不施肥)和浇水(20.1 kg·m^-2、不浇水)处理对矮嵩草补偿生长(包括分株密度、株高和分株地上生物量)的影响, 及其比叶面积、叶片净光合速率和相对增长率的变化, 探讨矮嵩草补偿生长的机制。研究结果表明: 刈割后, 矮嵩草的补偿生长高度和比叶面积显著降低; 分株密度有增加的趋势, 但会随刈割强度的增加而下降; 株高和生物量的相对增长率随刈割强度的增加而呈上升趋势; 补偿地上生物量在重度刈割处理下最高。施肥能显著增加矮嵩草的补偿高度、分株密度、补偿地上生物量、株高相对增长率、生物量相对增长率、比叶面积和净光合速率; 与不浇水处理相比, 浇水处理对重度刈割处理下的分株地上生物量、密度相对增长率、比叶面积和净光合速率无影响, 而显著降低了中度刈割处理下的补偿高度和株高相对增长率, 提高了不刈割处理下的分株密度和重度刈割处理下的生物量相对增长率。刈割、施肥和浇水处理的交互作用也显示出刈割与施肥对矮嵩草补偿生长具有拮抗效应, 而刈割与浇水具有协同效应。上述结果说明, 矮嵩草在刈割后可通过增加分株密度和相对增长率等途径来提高补偿能力, 弥补在生长高度上出现的低补偿, 而施肥可显著抵消刈割的不利影响, 提高矮嵩草的补偿能力。     

Compensatory growth offsets poor condition in native trout populations

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Overall grazing tolerance index (overall GTI) is not an ideal predictor for describing a single‐species tolerance to grazing

Plants' pattern of compensatory growth is often used to intuitively estimate their grazing tolerance. However, this tolerance is sometimes measured by the overall grazing tolerance index (overall GTI), which assumes that tolerance is a multivariate linear function of various underlying mechanisms. Because the interaction among mechanisms is not independent, the grazing tolerance expression based on overall GTI may be inconsistent with that based on compensatory growth. Through a manipulative field experiment from 2007 to 2012, we measured the responses of 12 traits of Elymus nutans to clipping under different resource availabilities in an alpine meadow and explored the compensatory aboveground biomass and the overall GTI to assess the possible differences between the two expressions of tolerance. Our results showed that these two expressions of tolerance were completely opposite. The expression based on overall GTI was over‐compensatory and did not vary with clipping and resource availability, while the expression based on compensatory aboveground biomass was under‐compensatory and altered to over‐compensation after fertilization. The over‐expression of highly variable traits with extremely high negative mean GTI to defoliation damage, the influence of random errors contained in traits considered, and the doubling weight of functional redundant traits greatly inflated the overall GTI, which leads to the inconsistency of the two tolerance expressions. This inconsistency is also associated with the different determining mechanisms of the two tolerance expressions. Our data suggest that plants' grazing tolerance is not a multivariate linear function of traits or mechanisms that determine grazing tolerance; the overall GTI is only a measure of traits' variability to defoliation damage. Our findings highlight that the tolerance of E. nutans mainly depends on the response of traits with lower variability to defoliation, and the overall GTI is not an ideal predictor for describing a single‐species tolerance to grazing.     

Autocrine activation of the MET receptor tyrosine kinase in acute myeloid leukemia

Although the treatment of acute myeloid leukemia (AML) has improved substantially in the past three decades, more than half of all patients develop disease that is refractory to intensive chemotherapy. Functional genomics approaches offer a means to discover specific molecules mediating the aberrant growth and survival of cancer cells. Thus, using a loss-of-function RNA interference genomic screen, we identified the aberrant expression of hepatocyte growth factor (HGF) as a crucial element in AML pathogenesis. We found HGF expression leading to autocrine activation of its receptor tyrosine kinase, MET, in nearly half of the AML cell lines and clinical samples we studied. Genetic depletion of HGF or MET potently inhibited the growth and survival of HGF-expressing AML cells. However, leukemic cells treated with the specific MET kinase inhibitor crizotinib developed resistance resulting from compensatory upregulation of HGF expression, leading to the restoration of MET signaling. In cases of AML where MET is coactivated with other tyrosine kinases, such as fibroblast growth factor receptor 1 (FGFR1), concomitant inhibition of FGFR1 and MET blocked this compensatory HGF upregulation, resulting in sustained logarithmic cell killing both in vitro and in xenograft models in vivo. Our results show a widespread dependence of AML cells on autocrine activation of MET, as well as the key role of compensatory upregulation of HGF expression in maintaining leukemogenic signaling by this receptor. We anticipate that these findings will lead to the design of additional strategies to block adaptive cellular responses that drive compensatory ligand expression as an essential component of the targeted inhibition of oncogenic receptors in human cancers.     

Growth, Nitrogen Uptake and Flow in Maize Plants Affected by Root Growth Restriction

The objective of the present study was to investigate the influence of a reduced maize root-system size on root growth and nitrogen (N) uptake and flow within plants. Restriction of shoot-borne root growth caused a strong decrease in the absorption of root: shoot dry weight ratio and a reduction in shoot growth. On the other hand, compensatory growth and an increased N uptake rate in the remaining roots were observed. Despite the limited long-distance transport pathway in the mesocotyl with restriction of shoot-borne root growth, N cycling within these plants was higher than those in control plants, implying that xylem and phloem flow velocities via the mesocotyl were considerably higher than in plants with an intact root system. The removal of the seminal roots in addition to restricting shoot-borne root development did not affect whole plant growth and N uptake, except for the stronger compensatory growth of the primary roots. Our results suggest that an adequate N supply to maize plant is maintained by compensatory growth of the remaining roots, increased N uptake rate and flow velocities within the xylem and phloem via the mesocotyl, and reduction in the shoot growth rate.