红松雌球果发育对枝叶营养生长和生物量积累的影响

通过对红松生殖枝和营养枝当年的枝叶生长及生物量积累差异的分析,研究2年生雌球果发育对枝叶营养生长和生物量积累的影响。结果表明:2年生雌球果抑制了1年生雌球果的发育,生殖枝当年的营养生长比营养枝当年的营养生长旺盛;生殖枝当年的生长量(顶枝芽除外)与母枝的枝长和基径呈显著正相关(P0.05),生殖枝当年的顶枝和侧枝生物量与母枝的枝生物量呈显著正相关(P0.05); 2年生雌球果数量与生殖枝当年的顶枝基径、侧枝长、侧枝基径、侧枝芽数量和侧枝数量呈显著正相关(P0.05),2年生雌球果生物量与当年的顶枝和侧枝生物量及总生物量呈显著正相关(P0.05),说明2年生雌球果的发育促进了生殖枝当年的营养生长和生物量积累;生殖枝当年的总生物量中分配给顶枝、侧枝、顶枝针叶、侧枝针叶的生物量比营养枝多,生殖枝中超过87%的当年生物量分配给2年生雌球果,仅有0.22%的比例分配给1年生雌球果;通径分析表明,2年生雌球果的生物量积累直接影响生殖枝母枝的生物量积累,对生殖枝和营养枝当年生物量积累没有直接影响,而生殖枝母枝的生物量直接影响生殖枝和营养枝当年的生物量积累。     

环剥对红松(Pinus koraiensis)韧皮部和木质部碳水化合物的影响

树干环剥可以阻碍韧皮部光合产物的运输并进一步影响光合产物的分配。长时期内,环剥能够导致环痕上部可溶性糖和淀粉的积累,但对于短期内如何影响碳水化合物在木质部和韧皮部内的运输模式所知甚少。以38年生红松(Pinus koraiensis Sieb.etZucc.)为研究材料,分别对环剥上部、下部每隔1~2d采样,区分木质部和韧皮部(树皮)进行可溶性糖和淀粉含量及树干糖呼吸消耗速率测定,确定环剥后的日变化和周变化,并对木质部可溶性糖、淀粉含量与韧皮部中相应指标进行相关关系的回归分析。结果发现:(1)环剥后4周内,在环剥痕上、下部间木质部可溶性糖和淀粉含量,韧皮部中淀粉含量均不存在显著差异(p>0.05),而韧皮部内可溶性糖含量,环剥后第2周出现显著差异,从第4周出现环剥上部显著高于下部的碳水化合物积累现象(p<0.05);(2)环剥阻隔了韧皮部可溶性糖的纵向运输,但是并不影响木质部的纵向运输,而且环剥并没有影响木质部和韧皮部之间的糖和淀粉的相关关系;(3)环剥第1周内环剥上部和下部呼吸消耗速率差异不显著,第2周环剥上部显著高于环剥下部,从第3周开始环剥下部呼吸消耗速率显著下降。推断认为,在环剥处理的4周内,环剥上部冠层新形成的碳水化合物很大一部分均被呼吸消耗掉,导致环剥上部较环剥下部可溶性糖稍有增加;红松胸高直径以下部分所储藏的碳水化合物足以保障2周内红松树干呼吸。     

基于空间变异性的树冠非结构性碳含量估算：以胡桃楸和春榆为例

以胡桃楸和春榆为例,采用枝解析法测定不同基径枝条的非结构性碳水化合物（NSC）浓度的变异,进而估算并评价不同取样方法对树冠NSC含量估算的误差.结果表明： 器官对两种树种树冠NSC浓度的影响显著,叶、新枝、老枝和死枝的可溶性糖与淀粉的总和（TNC）的平均浓度分别为17.6%、12.6%、5.7%和2.9%.叶和新枝的NSC浓度随枝基径、枝龄、枝长和枝相对高度的变异多不显著,但老枝的NSC浓度随枝基径、枝龄和枝长增大而降低,而随枝相对高度增加而增加,其中枝基径是老枝NSC浓度的最佳预测变量（R²在0.87~0.95）.两树种叶、新枝和老枝平均TNC含量分别占其树冠TNC含量的28%、2%和70%.分析树冠枝NSC的空间变异性对树冠NSC含量估算误差的影响发现,采用新枝和直径约3 cm老枝枝段的NSC浓度估算冠层NSC含量是较简易而精确的方法.     

去叶对青杨雌雄植株生长和非结构性碳水化合物的影响

近年来,森林食叶害虫在全世界呈爆发趋势。树木的非结构性碳水化合物(NSC)如何响应叶片损失对其生长和生存至关重要。雌雄异株植物在维持森林生态系统稳定性方面扮演着重要角色。然而,目前对该类植物性别之间如何响应去叶的研究还比较少。本文以我国重要的经济和生态恢复树种青杨(Populus cathayana)为研究材料,比较了雌雄青杨幼苗的生长、NSC含量和储量对去叶(0,50%和100%叶片去除)的响应差异。结果表明:随去叶强度的增加,植物的生物量和植株NSC呈降低趋势,且根系(尤其是粗根)的生物量和NSC比地上部分受去叶的影响更大;雌株叶、粗根、细根和植株NSC储量总是高于雄株;随去叶强度增加,雄株的生物量积累和NSC含量和储量降低得比雌株更多。这些结果表明,青杨雌雄植株生长和NSC对不同去叶强度的响应存在性别差异,且去叶对青杨雄株的影响更大。这暗示了雌性青杨对去叶的耐性比雄性强。这些结果有助于理解雌雄异株植株性别水平上的碳平衡机理,也可为杨树人工林的选育提供支撑。     

氮添加对稀土尾砂地猴樟幼苗根系生长、生物量分配及非结构性碳水化合物的影响

为探究氮添加对稀土矿尾砂地猴樟(Cinnamomun bodinieri)幼苗生长及非结构性碳水化合物(NSC)含量的影响，以1年生猴樟扦插苗为研究对象，选用硝酸铵钙作为氮肥(含N 15%)，设置3种氮肥水平(CK(0)、N1(1.8 g·株^-1)、N2(3.6 g·株^-1))，分析不同氮添加水平下猴樟幼苗根系生长、生物量分配和NSC等指标的差异，探讨稀土尾砂地猴樟对氮添加的响应。结果发现：稀土尾砂地氮添加均增加了猴樟幼苗不同组织的生物量积累，其中N1处理下叶生物量、冠层生物量分别较N2处理下显著提升了44.75%、57.43%(P<0.05);N2处理下叶比重分别较CK、N1处理显著提升了123.53%、15.85%(P<0.05)。不论是粗根(直径>2 mm)，还是细根(直径≤2 mm)，氮添加均显著增加了植物的根长和根表面积(P<0.05)，其中N1处理的促进效果最显著(P<0.05);N2处理下的比根长、比表面积均高于CK与N1，且与CK显著差异(P<0.05)。对NSC来说，N1处理提升了叶、茎中...     

林地覆盖经营对雷竹叶片非结构性碳水化合物与氮、磷关系的影响

为揭示雷竹生理生态对林地覆盖经营的响应与适应机制,测定了不同覆盖经营年限(CK、1、3和6年)雷竹林1~3年生立竹叶片可溶性糖、淀粉、氮(N)和磷(P)含量,分析了覆盖经营对雷竹林叶片非结构性碳水化合物(NSC)与N、P含量及其化学计量关系的影响.结果表明： 覆盖经营1年雷竹林叶片NSC及可溶性糖含量较CK显著升高,N/P显著降低,N限制作用增强;覆盖经营3年雷竹林叶片可溶性糖含量较CK显著降低,淀粉含量显著升高,而NSC含量变化不明显,单位质量N、P的NSC含量最高;覆盖经营6年雷竹林叶片NSC、可溶性糖含量较CK显著降低,淀粉含量和N/P显著升高,P限制作用增强.短期覆盖(≤3年)经营雷竹林叶片NSC含量与N、P含量呈显著正相关,与N/P呈显著负相关;长期覆盖(6年)经营雷竹林叶片淀粉含量与N、P含量呈显著负相关,与N/P呈显著正相关.研究表明,短期覆盖(≤3年)经营雷竹林叶片N/P的降低可促使叶片淀粉分解和可溶性糖积累,明显增强雷竹生长活性,而长期覆盖(6年)经营雷竹林叶片N/P升高则促进叶片淀粉积累,雷竹生长由N限制转变为P限制,立竹生长活性明显降低,引起雷竹林退化.因此建议雷竹林连续覆盖经营不宜超过3年.      

不同修剪措施对薄壳山核桃枝条生长及枝条和叶片碳氮代谢物积累的影响

以薄壳山核桃品种‘马罕’（Caryaillinoensis‘Mahan’）的5年生嫁接苗为实验材料,研究枝条短截（1/4、1/3和1/2短截）以及枝条和主干的环剥和环割对其枝条生长及枝条和叶片中碳氮代谢物积累的影响。结果显示：经不同程度短截处理后,枝条萌芽率均显著高于对照（未经任何修剪）,新枝的数量、长度和直径也均不同程度高于对照,而比叶质量及叶绿素含量总体上与对照无显著差异;经1/2和1/3短截处理后,长度0-10cm和30cm以上的新枝比例明显提高;枝条和叶片中可溶性糖含量和C/N比均高于对照、全N含量均低于对照,枝条中淀粉含量低于对照而叶片中淀粉含量高于对照。经枝条环剥和环割处理后,枝条萌芽率和比叶质量均高于对照但无显著差异,枝条平均长度增长量和叶绿素含量均显著低于对照、枝条平均直径增长量均显著高于对照;枝条和叶片中可溶性糖和淀粉含量以及C/N比均高于对照,全N含量均低于对照。经主干环剥和环割处理后,枝条的萌芽率和平均直径增长量以及比叶质量均显著高于对照,枝条平均长度增长量和叶绿素含量均显著低于对照;枝条和叶片中可溶性糖和淀粉含量以及C/N比均高于对照,枝条中全N含量高于对照而叶片中全N含量则低于对照。此外,品种‘马罕’的结果枝长度为0-30cm,其中长度0-10cm的结果枝数量最多。研究结果表明：不同短截措施均能提高薄壳山核桃的萌芽率、促进新枝伸长和增粗;而枝条和主干的环剥和环割处理对枝条萌芽率无明显促进作用,但有利于枝条增粗;不同修剪措施总体上有利于其叶片及枝条中碳水化合物的合成和积累。总体上,1/3短截及枝条和主干的适度环剥可促进品种‘马罕’结果枝的形成。     

干旱胁迫下接种丛枝菌根真菌对七子花非结构性碳水化合物积累及C、N、P化学计量特征的影响

以濒危植物七子花二年生幼苗为研究材料,采用盆栽试验方法,研究干旱胁迫和接种丛枝菌根真菌(AMF)处理对幼苗不同器官C、N、P化学计量关系和非结构性碳水化合物(NSC)含量的影响。试验共设计4个处理:对照(CK)、干旱胁迫(D)、接种AMF(AMF)、干旱胁迫和接种AMF(D+AMF)。结果表明: 在干旱胁迫下七子花根系AMF的侵染率显著下降,但接种AMF处理植株的株高、叶片数显著高于未接种处理。接种AMF显著提高了干旱胁迫下植株根、叶可溶性糖和NSC含量及茎、叶淀粉含量,且茎和叶可溶性糖与淀粉比显著下降。干旱胁迫导致植株C含量在根和叶中显著增加,P含量在茎中显著减少;与干旱胁迫相比,胁迫下接种AMF植株根、茎、叶P含量及叶C含量显著提高,而根C、N含量及茎C含量显著降低。胁迫下接种AMF植株根、茎C∶N、C∶P、N∶P和叶N∶P均显著低于单一胁迫处理。NSC与C∶N∶P计量比的相关性分析表明,根、叶P含量与可溶性糖和NSC含量呈显著正相关,茎P含量与淀粉和NSC含量呈显著正相关,各器官N∶P与NSC含量呈显著负相关。综上,干旱胁迫显著抑制了七子花幼苗的生长,接种AMF通过提高植株根和叶的可溶性糖含量、根的可溶性糖/淀粉,增加地上部分淀粉含量,促进P元素吸收和降低各器官N∶P来增强植株耐旱性,从而提高七子花幼苗在干旱环境中的存活率。     

杉木幼苗生物量分配格局对氮添加的响应

大气氮(N)沉降的急剧增加可能会对植物碳(C)固定和分配产生深远影响。然而, N添加如何影响碳水化合物在植物不同器官之间的分配动态并不十分清楚。该研究利用杉木(Cunninghamia lanceolata)幼苗盆栽试验, 设置N添加处理, 测定分析幼苗非结构性碳水化合物(NSC)与结构性碳水化合物(SC)含量和库的变化, 以探讨N添加后杉木幼苗不同器官中NSC与SC的分配模式及调控机制。结果发现: (1) N添加虽然显著增加叶片净光合速率(143.96%), 但却降低了叶片中的NSC含量和库; N添加导致一年生茎的淀粉含量显著下降, 而可溶性糖含量的变化不显著, 当年生茎的NSC组分含量和库没有显著变化; 幼苗根系的NSC及其组分含量和库也有降低的趋势。(2) N添加后地下与地上生物量的比值降低22.09%, 其中SC库比值降低31.07%, 而NSC库比值无显著变化。(3) N添加使地上部分的磷(P)库显著增加, 使地下与地上P库的比值降低了57.02%, 而N库的比值无显著变化。(4) N添加后土壤pH由(4.94 ± 0.09)显著降低到(4.02 ± 0.04), 铵态N和硝态N含量分别增加7.17倍和11.55倍, 土壤有效P含量也增加了42.86%, 而土壤中脲酶(62.75%)和酸性磷酸酶(56.52%)的活性显著降低。研究表明, 低养分条件下杉木幼苗主要通过构建根系结构增加养分吸收, 而非通过向根系分配更多的NSC, 而N添加驱动的养分缓解使更多的碳水化合物分配到地上器官, 导致地上部分SC积累。     

养分异质条件下结缕草克隆分株生长、碳水化合物及可溶性蛋白的生理整合

通过对连接和断开的结缕草分株进行不同的养分处理,研究了养分异质条件下结缕草克隆分株生长、碳水化合物及可溶性蛋白含量的变化.结果表明: 在养分异质条件下,处于中、高养分水平的母株可以显著提高与其相连子株的地上、地下及总生物量,高养分下分别提高32.5%、22.1%和24.8%,降低根冠比、可溶性糖及非结构性碳水化合物(NSC)含量,高养分下分别降低7.7%、15.2%和13.1%,但是对淀粉、纤维素、可溶性蛋白含量无显著影响.处于中、高养分水平的子株对与其相连的母株生长、碳水化合物及可溶性蛋白含量没有显著影响.养分异质条件下结缕草母株对子株生物量、根冠比、可溶性糖及NSC含量有明显的生理整合,其整合强度与养分梯度呈正比,而对淀粉、纤维素及可溶性蛋白含量没有显现显著的生理整合效应.子株对母株各项指标都没有显著的生理整合,结缕草母株和子株间是一种单向的生理整合.     

摘叶造成的碳限制对刺槐碳素分配和水力学特性的影响

以3年生刺槐（Robinia pseudoacacia Linn.）为研究对象,通过对其进行连续3次摘叶造成严重碳限制,检测摘叶后刺槐的生物量分配、叶片形态和不同部位的非结构性碳（NSC）浓度,同时检测其根压和根系导水率、枝条水势和导水率损失值（PLC）及茎的抗栓塞能力,研究摘叶造成的碳限制对刺槐碳素分配和水力学特性的影响。结果显示,摘叶显著降低了刺槐不同部位的生物量,其中细根生物量降低程度最大;摘叶还造成了刺槐不同部位NSC浓度显著降低,茎韧皮部、茎木质部、根韧皮部和根木质部的NSC浓度分别为对照的29.6%、20.2%、10.2%和8.7%,且根部NSC的降低程度显著高于茎;碳限制显著降低了刺槐苗木的根压和根系导水率,增加了枝条凌晨和正午的PLC,降低了其抗栓塞能力。研究结果表明摘叶造成的碳限制改变了刺槐的碳素分配模式,限制了碳素向根的分配,抑制细根的发生,进而限制根的水分吸收能力,加重枝条栓塞程度,同时还会导致枝条抗栓塞能力下降,从而降低植物水分输导的安全性。     

THE ROLE OF RESERVES IN LEAVES,BRANCHES, STEMS,AND ROOTS ON SHOOT GROWTH OF RED PINE

By defoliation, girdling, and various combinations of these before the growing season began, the role of foods in old needles, branches, main stems, and roots on shoot growth of 8-yr-old red pine (Pinus resinosa) trees was studied in northern Wisconsin. Defoliation and girdling, alone or in combination, reduced shoot growth but defoliation reduced shoot dry weight more than it reduced shoot elongation. Reductions in shoot growth due to treatment were in the following decreasing order: branch girdling + needle removal > girdling at stem base + needle removal > needle removal > branch girdling > base girdling. The old needles were the major source of food for shoot growth and accounted for four-fifths or more of all shoot growth. The combined reserves in the branches, main stem, and roots accounted for less than 15% of shoot growth. The contribution of reserves from tissues other than old leaves was in the following order: branches > main stem > roots. Defoliation weakened apical dominance relations. In defoliated trees, many secondary axes elongated more than the terminal leader, and secondary axes in lower whorls often grew more than those in upper ones. The data suggest an important role of nutritional factors in correlative growth inhibition.     

Interguild interactions between a non-native phloem feeder and a native outbreaking defoliator

1. Competitive and synergistic interactions directly or indirectly drive community dynamics of herbivorous insects. Novel interactions between non-native and native insects are unpredictable and not fully understood. 2. We used manipulative experiments on mature red spruce trees to test interactions between a non-native phloem feeding insect, the brown spruce longhorn beetle (BSLB), and an outbreaking native defoliator, the spruce budworm. We subjected treatment trees to defoliation by three densities of spruce budworm larvae. Treatment trees were: stressed by (i) girdling (to mimic beetle feeding) or (ii) girdling + BSLB before spruce budworm larvae were introduced on branches in sleeve cages. Budworm larvae then fed on foliage and developed to pupation. We assessed all branches for budworm performance, defoliation, shoot production and shoot growth. 3. Shoot length did not differ in response to stress from girdling or BSLB infestation. Neither stress from girdling, nor interactions with BSLB feeding affected spruce budworm performance or defoliation. Intraspecific impacts on performance and defoliation in relation to budworm density were stronger than the effects of tree stress. 4. Prior infestation of red spruce by BSLB in our experimental set-up did not influence spruce budworm performance. BSLB is a successful invader that has blended into its novel ecological niche because of ecological and phylogenetic similarities with a native congener, Tetropium cinnamopterum. 5. Outbreaks by BSLB will not likely impede or facilitate spruce budworm outbreaks if they co-occur. It would be useful to evaluate the reverse scenario of BSLB success after defoliation stress by spruce budworm.     

Impact of phloem girdling on leaf gas exchange and hydraulic conductance in hybrid aspen

We investigated phloem-xylem interactions in relation to leaf hydraulic capacity in hybrid aspen (Populus tremula L. × P. tremuloides Michx.) by using phloem girdling method. Removal of bark tissues (phloem girdling) at the branch base resulted in a substantial decline in stomatal conductance (g_S), net photosynthetic rate (P_N), and leaf hydraulic efficiency, and in increase of leaf water potential (Ψ_L). Although gS declined more than P_N (83 versus 78 %), the ratio of intercellular to ambient CO₂ concentrations (c_i/c_a) increased from 0.67 to 0.87 in three days after girdling. Girdling induced a decrease in leaf hydraulic conductance (K_L) on average by 43 % (P = 0.006). The changes in gS and leaf conductance to water vapour were co-ordinated with K_L only in girdled branches whereas intrinsic water-use efficiency was invariant to K_L. The declines in K_L with girdling were not accompanied by changes in potassium ion concentration ([K⁺]), electrical conductivity, or pH of xylem sap. The results suggest that phloem girdling at the branch base does not influence the recirculation of ions between the phloem and xylem in hybrid aspen and the decrease of K_L in response to the manipulation is not related to changes in [K⁺] and total ionic content of xylem sap.     

Water stress, shoot growth and storage of non-structural carbohydrates along a tree height gradient in a tall conifer

We analysed concentrations of starch, sucrose, glucose and fructose in upper branch wood, foliage and trunk sapwood of Douglas-fir trees in height classes ranging from ~2 to ~57 m. Mean concentrations of non-structural carbohydrates (NSC) for all tissues were highest in the tallest height class and lowest in the lowest height class, and height-related trends in NSC were most pronounced in branches. Throughout a 17-month sampling period, mean values of branch NSC from the 57 m trees ranged between 30 and 377% greater than the 2 m trees. Branch NSC was inversely correlated with midday shoot water potential (Ψ(l)), shoot osmotic potential at full turgor (Ψ) and shoot extension. Temporal fluctuation in branch NSC was inversely correlated with height, and positively correlated with midday Ψ(l) , Ψ and shoot extension. The positive correlation between height and storage of NSC, and the negative correlation between NSC storage and shoot extension provide evidence that size-related growth decline in trees is not strongly associated with constraints on photosynthesis. The negative correlation between height and fluctuation in NSC suggests that mobilization of photosynthate in taller trees is constrained by some factor such as reductions in turgor-driven cell expansion or constraints on phloem transport.     

长白山阔叶红松林3个主要树种的非结构性碳储存特征

非结构性碳水化合物(NSC,包括可溶性糖和淀粉)作为树木生命代谢的关键物质之一,在树木器官中的储存特征受到了广泛关注,但NSC在器官内部具有不同功能的组织间(韧皮部与木质部)的分配和权衡还不清楚.本研究以长白山阔叶红松林的3个优势树种——红松、水曲柳和紫椴为研究对象,对比分析NSC在根、树干韧皮与木质部中的浓度和分配特征.结果表明: 树木韧皮部和木质部间的NSC浓度差异显著,总体分配趋势为韧皮部以可溶性糖为主,而木质部以淀粉为主.树干外侧(以年轮划分,0～20年)、中段(20～40年)和内侧(>40年)的NSC浓度在不同树种间差异显著,而根中的差异不显著.红松和水曲柳树干韧皮部可溶性总糖浓度显著高于紫椴,在木质部中差异不显著.本研究结果表明,树体内部NSC在韧皮部和木质部上的分配存在明显分异,这与树种的演替阶段及组织的功能进化有关.研究结果对于深入理解温带树木的碳储存特征和分配机理具有参考意义.     

Transport Exchange of Carbon,Nitrogen and Water in the Context of Whole Plant Growth and Functioning — Case History of a Nodulated Annual Legume

The economy of carbon, nitrogen and water during growth of nodulated, nitrogen-fixing plants of white lupin (Lupinus albus L.) was studied by measuring C, N and H₂O content of plant parts, concentrations of C and N in bleeding sap of xylem and phloem, transpirational losses of whole shoots and shoot parts, and daily exchanges of CO₂ between shoot and root parts and the surrounding atmosphere. Relationships were studied between water use and dry matter accumulation of shoot and fruits, and between net photosynthesis rate and leaf area, transpiration rate and nitrogen fixation. Conversion efficiencies were computed for utilization of net photosynthate for nitrogen fixation and for production of dry matter and protein in seeds. Partitioning of the plant's intake of C, N and H₂O was described in terms of growth, transpiration, and respiration of plant parts. An empirically-based model was developed to describe transport exchanges in xylem and phloem for a 10-day interval of growth. The model depicted quantitatively the mixtures of xylem and phloem streams which matched precisely the recorded amounts of C, N and H₂O assimilated, absorbed or consumed by the various parts of the plant. The model provided information on phloem translocation of carbon and nitrogen to roots from shoots, the cycling of carbon and nitrogen through leaves, the relationship between transpiration and nitrogen partitioning to shoot organs through the xylem, the relative amount of the plant's water budget committed to phloem translocation, and the significance of xylem to phloem transfer of nitrogen in stems as a means of supplying nitrogen to apical regions of the shoot.     

Modelling of the Partitioning, Assimilation and Storage of Nitrate within Root and Shoot Organs of Castor Bean (Ricinus communis L.)

An experimentally-based modelling technique was developed todescribe quantitatively the uptake, flow, storage and utilizationof NO₃-N over a 9 d period in mid-vegetative growth of sandcultured castor bean (Ricinus communis L.) fed 12 mol m^–3nitrate and exposed to a mean salinity stress of 128 mol m^–3NaCl. Model construction used information on increments or lossesof NO₃-N or total reduced N in plant parts over the study periodand concentration data for NO₃-N and reduced (amino acid) Nin phloem sap and pressure-induced xylem exudates obtained fromstem, petiole and leaf lamina tissue at various levels up ashoot. The resulting models indicated that the bulk (87%) of incomingnitrate was reduced, 51% of this in the root, the remainderprincipally in the laminae of leaves. The shoot was 60% autotrophicfor N through its own nitrate assimilation, but was oversuppliedwith surplus reduced N generated by the root and fed to theshoot through the xylem. The equivalent of over half (53%) ofthis N returned to the root as phloem translocate and, mostly,then cycled back to the shoot via xylem. Nitrate comprised almosthalf of the N of most xylem samples, but less than 1% of phloemsap N. Laminae of leaves of different age varied greatly inN balance. The fully grown lower three leaves generated a surplusof reduced N by nitrate assimilation and this, accompanied byreduced N cycling by xylem to phloem exchange, was exportedfrom the leaf. Leaf 4 was gauged to be just self-sufficientin terms of nitrate reduction, while also cycling reduced N.The three upper leaves (5–7) met their N balance to varyingextents by xylem import, phloem import (leaves 6 and 7 only)and assimilation of nitrate. Petioles and stem tissue generallyshowed low reductase activities, but obtained most of theirN by abstraction from xylem and phloem streams. The models predictedthat nodal tissue of lower parts of the stem abstracted reducedN from the departing leaf traces and transferred this, but notnitrate, to xylem streams passing further up the shoot. As aresult, xylem sap was predicted to become more concentratedin N as it passed up the shoot, and to decrease the ratio ofNO₃-N to reduced N from 0·45 to 0·21 from thebase to the top of the shoot. These changes were reflected inthe measured N values for pressure-induced xylem exudates fromdifferent sites on the shoot. Transfer cells, observed in thexylem of leaf traces exiting from nodal tissue, were suggestedto be involved in the abstraction process. Key words: Ricinus communis, nitrogen, nitrate, nitrate reduction, partitioning, phloem, xylem, flow models     

Modeling the transport and utilization of carbon and nitrogen in a nodulated legume

An empirical modeling technique was developed for depicting quantitatively the transport and partitioning of photosynthetically fixed C and symbiotically fixed N during 10-day intervals of a 40-day period in the growth of nodulated plants of white lupin (Lupinus albus L. cv. Ultra). Model construction utilized data for C and N consumption of plant parts and C:N weight ratios of the xylem and phloem fluids serving specific plant organs. Formulas were derived from calculating the net transport of C and N between plant parts in xylem and phloem. The models provided quantitative information on the dependence of growing organs on xylem and phloem for their supply of C and N, the cycling of N through leaflets and of C through nodules, the extent of direct incorporation of fixed N into growing nodules, and the involvement of N from shoot translocate in the nutrition of the nodulated root. Stem plus petioles abstracted considerably more N from xylem than expected from their transpirational activity. Xylem to phloem transfer of recently fixed N in mature stem and petioles was substantiated by the models, being depicted as a device for dispensing N to growing parts of the shoot extra to that attracted transpirationally in xylem or received as translocate from leaflets.