AUX1 promotes lateral root formation by facilitating indole-3-acetic acid distribution between sink and source tissues in the Arabidopsis seedling

Arabidopsis root architecture is regulated by shoot-derived signals such as nitrate and auxin. We report that mutations in the putative auxin influx carrier AUX1 modify root architecture as a result of the disruption in hormone transport between indole-3-acetic acid (IAA) source and sink tissues. Gas chromatography-selected reaction monitoring-mass spectrometry measurements revealed that the aux1 mutant exhibited altered IAA distribution in young leaf and root tissues, the major IAA source and sink organs, respectively, in the developing seedling. Expression studies using the auxin-inducible reporter IAA2::uidA revealed that AUX1 facilitates IAA loading into the leaf vascular transport system. AUX1 also facilitates IAA unloading in the primary root apex and developing lateral root primordium. Exogenous application of the synthetic auxin 1-naphthylacetic acid is able to rescue the aux1 lateral root phenotype, implying that root auxin levels are suboptimal for lateral root primordium initiation in the mutant.     

Relationships of endogenous plant hormones to accumulation of grain protein and starch in winter wheat under different post-anthesis soil water statusses

Accumulation of protein and starch in grain is a key process determining grain yield and quality in wheat. Under drought or waterlogging, endogenous plant hormone levels will change and may have an impact on the yield and quality of wheat. In a greenhouse experiment, four winter wheat (Triticum aestivum L.) varieties differing in grain protein content, Heimai 76, Wanmai 38, Yangmai 10 and Yangmai 9, were subjected to drought (SRWC = 4550%, DR), waterlogging (WL) and moderate water supply (SRWC = 7580%, CK), beginning from 4 days post-anthesis (DPA) to maturity. On the 10 (grain enlargement stage) and 20 (grain filling stage) DPA, endogenous abscisic acid (ABA), gibberellins (GA₁₊₃), indole-3-acetic acid (IAA) and zeatin riboside (ZR) were determined in sink and source organs of wheat plants by enzyme linked immunosorbent assay (ELISA). The patterns of hormonal changes were similar in four varieties. The ABA levels were much higher under DR and WL than under CK. Compared with CK, GA₁₊₃ levels in whole-plant under DR and WL changed a little at 10 DPA, but markedly decreased under DR and WL at 20 DPA. Changes of endogenous IAA level under DR and WL exhibited a complicated pattern, depending on organs and growth stages. Particularly at the 20 DPA, the mean levels of IAA in roots, leaves and grains decreased significantly under DR and WL. In comparison with CK, ZR levels in all organs significantly decreased under DR and WL at both stages. The correlation analyses between yields and contents of starch and protein in grains and levels and ratios of four hormones in source and sink organs indicated that the changes in yield and content of grain starch and protein under DR and WL were associated with the reduced IAA, ZR and GA₁₊₃ levels and elevated ABA level in plants, especially in grains. It was proposed that the changed levels of endogenous hormones under post-anthesis DR and WL might indirectly affect protein and starch accumulation in grains by influencing the regulatory enzymes and processes.     

Gibberellic-acid-promoted transport of assimilates in stems of Phaseolus vulgaris L.

Gibberellic acid (GA₃), applied as a dispersion in aqueous lanolin to the stumps of decapitated stems of P. vulgaris plants, was found to promote the transfer of ¹⁴C-and ³²P-labelled assimilates to the site of hormone application. Measurements of the component transfer processes, operating between source and sink (site of hormone application), showed that GA₃ was not acting to promote assimilate transfer by increasing the photosynthetic rate of, or the assimilate export rate from the source, nor by altering the mobilizing ability of the competing root sink. Here, it also was found that the time between GA₃ application and detection of an enhanced transport flux was independent of the length of the transport pathway. Overall, the evidence obtained indicated that GA₃ was not acting on any transfer process remote from its point of hormone application but was acting locally at this latter point.Abbreviations GA₃ gibberellic acid - IAA indol-3yl-acetic acid     

Improvement of pea biomass and seed productivity by simultaneous increase of phloem and embryo loading with amino acids

The development of sink organs such as fruits and seeds strongly depends on the amount of nitrogen that is moved within the phloem from photosynthetic‐active source leaves to the reproductive sinks. In many plant species nitrogen is transported as amino acids. In pea (Pisum sativum L.), source to sink partitioning of amino acids requires at least two active transport events mediated by plasma membrane‐localized proteins, and these are: (i) amino acid phloem loading; and (ii) import of amino acids into the seed cotyledons via epidermal transfer cells. As each of these transport steps might potentially be limiting to efficient nitrogen delivery to the pea embryo, we manipulated both simultaneously. Additional copies of the pea amino acid permease PsAAP1 were introduced into the pea genome and expression of the transporter was targeted to the sieve element‐companion cell complexes of the leaf phloem and to the epidermis of the seed cotyledons. The transgenic pea plants showed increased phloem loading and embryo loading of amino acids resulting in improved long distance transport of nitrogen, sink development and seed protein accumulation. Analyses of root and leaf tissues further revealed that genetic manipulation positively affected root nitrogen uptake, as well as primary source and sink metabolism. Overall, the results suggest that amino acid phloem loading exerts regulatory control over pea biomass production and seed yield, and that import of amino acids into the cotyledons limits seed protein levels.     

Separation and Determination of Heavy Metals Associated with Low Molecular Weight Chelators in Xylem Saps of Indian Mustard (<Emphasis Type="Italic">Brassica juncea</Emphasis>) by Size Exclusion Chromatography and Atomic Absorption Spectrometry

To elucidate the role of low molecular weight chelators in long-distance root-to-shoot transport of heavy metals in Indian mustard, an “off-line” size exclusion high-performance liquid chromatography–graphite furnace atomic absorption spectrometry was developed to investigate heavy metals associated with low molecular weight chelators in xylem saps of Indian mustard (Brassica juncea). The size exclusion chromatogram presented only the peaks with molecular weight for all xylem saps and directly indicated the long-distance transport of phytochelatins (PCs) of Indian mustard under Cd stress. In the absence of Cd stress, only organic acids and inorganic anions participated in the long-distance transport of Cd, but organic acids, inorganic anions, glutathione (GSH), and cysteine might relate to the long-distance transport of Cu or Zn. In the presence of Cd stress, PCs were induced, and Cd ions in xylem saps were associated with the induced PCs. As the Cd levels in nutrient solution increased, more Cd in xylem saps adopted the form of PC–Cd. Although PCs might participate in the long-distance transport of Cd under Cd stress, the majority of Cd was still transported by organic acids and inorganic anions in xylem vessels. Moreover, results indicated the existence of complexation competition for GSH and cysteine between Cd and Cu (or Zn) and complexation competition for Cd between PCs and GSH (or cysteine) in xylem vessels. Our work might be very useful for understanding the mechanism of long-distance transport of heavy metals in hyperaccumulator.     

Effective carbon partitioning driven by exotic phloem-specific regulatory elements fused to the Arabidopsis thaliana AtSUC2 sucrose-proton symporter gene

Background  

AtSUC2 (At1g22710) from Arabidopsis thaliana encodes a phloem-localized sucrose/proton symporter required for efficient photoassimilate transport from source tissues to sink tissues. AtSUC2 plays a key role in coordinating the demands of sink tissues with the output capacity of source leaves, and in maintaining phloem hydrostatic pressure during changes in plant-water balance. Expression and activity are regulated, both positively and negatively, by developmental (sink to source transition) and environmental cues, including light, diurnal changes, photoassimilate levels, turgor pressure, drought and osmotic stress, and hormones.     

Physiological implications of SWEETs in plants and their potential applications in improving source–sink relationships for enhanced yield

The sugars will eventually be exported transporters (SWEET) family of transporters in plants is identified as a novel class of sugar carriers capable of transporting sugars, sugar alcohols and hormones. Functioning in intercellular sugar transport, SWEETs influence a wide range of physiologically important processes. SWEETs regulate the development of sink organs by providing nutritional support from source leaves, responses to abiotic stresses by maintaining intracellular sugar concentrations, and host–pathogen interactions through the modulation of apoplastic sugar levels. Many bacterial and fungal pathogens activate the expression of SWEET genes in species such as rice and Arabidopsis to gain access to the nutrients that support virulence. The genetic manipulation of SWEETs has led to the generation of bacterial blight (BB)-resistant rice varieties. Similarly, while the overexpression of the SWEETs involved in sucrose export from leaves and pathogenesis led to growth retardation and yield penalties, plants overexpressing SWEETs show improved disease resistance. Such findings demonstrate the complex functions of SWEETs in growth and stress tolerance. Here, we review the importance of SWEETs in plant–pathogen and source–sink interactions and abiotic stress resistance. We highlight the possible applications of SWEETs in crop improvement programmes aimed at improving sink and source strengths important for enhancing the sustainability of yield. We discuss how the adverse effects of the overexpression of SWEETs on plant growth may be overcome.     

Carbon allocation in fruit trees: from theory to modelling

Carbon allocation within a plant depends on complex rules linking source organs (mainly shoots) and sink organs (mainly roots and fruits). The complexity of these rules comes from both regulations and interactions between various plant processes involving carbon. This paper presents these regulations and interactions, and analyses how agricultural management can influence them. Ecophysiological models of carbon production and allocation are good tools for such analyses. The fundamental bases of these models are first presented, focusing on their underlying processes and concepts. Different approaches are used for modelling carbon economy. They are classified as empirical, teleonomic, driven by source–sink relationships, or based on transport and chemical/biochemical conversion concepts. These four approaches are presented with a particular emphasis on the regulations and interactions between organs and between processes. The role of plant architecture in carbon partitioning is also discussed and the interest of coupling plant architecture models with carbon allocation models is highlighted. As an illustration of carbon allocation models, a model developed for peach trees, describing carbon transfer within the plant, and based on source–sink and Münch transport theory is presented and used for analyzing the link between roots, shoots and reproductive compartments. On this basis, the consequences of fruit load or plant pruning on fruit and vegetative growth can be evaluated.     

Regulation of nitrogen partitioning in field-grown almond trees: Effects of fruit load and foliar nitrogen applications

Two treatments were employed to influence the amount of amino nitrogen (N) transport in phloem. In walnut trees (Juglans regia L.), developing fruit significantly reduced the efflux of foliar-applied ¹⁵N-enriched urea from treated spurs over a 33-day period in comparison with similarly-treated defruited spurs. Those data suggest that local aboveground demand for N influences vascular transport of amino N. In another experiment, a 1% urea solution was applied foliarly to 5-year old `Mission' almond trees [Prunus dulcis (Mill.) D. A. Webb] to increase the concentration of amino N in the phloem. The effect of foliar N treatments on a) the transport and distribution of labelled urea N within the trees over the experimental period and b) the uptake of soil-applied labelled N were determined by replicated whole tree excavation, fractionation into various tree components and mass spectrometric analyses of the ¹⁴N/¹⁵N ratios. Concentrations and composition of amino acids in the phloem and xylem saps of control trees and trees receiving foliar-applied urea were also determined. In foliar urea-treated trees, the amino acid concentrations increased significantly in leaf and bark phloem exudate, within 24 and 96 h, respectively. Foliar-applied urea N was translocated to the roots of almond trees over the experimental period and decreased soil N uptake. The results of these experiments are consistent with the hypothesis that aboveground N demand affects the amount of amino N cycling between shoots and roots, and may be involved in the regulation of soil N uptake. This revised version was published online in June 2006 with corrections to the Cover Date.     

高等植物光合同化物的运输与分配

高等植物光合同化物的运输受维管束发育状况影响较大,有时会限制产量。而同化物在各库器官间的分配主要决定于库本身的特性,它常用库强度和优先权来描述。库强度是库容量和库活力的乘积,库容量用细胞数目来度量,而库活力常用相对生长速度来度量。近年来人们也用酶少戌一来度量库活力或库强度。而库的优先权描述的是各库器官需求同化物的优先次序,种子被认为是优等权最高的库。同化物的运输分配不仅决定于植物本身源、流、库的特     

Phloem transport and differential unloading in pea seedlings after source and sink manipulations

Phloem transport was investigated in pea seedlings after application of [¹⁴C]sucrose to the cotyledons. The accumulation of the label in segments of young seedlings shows a differential unloading along the plant axis. Shoot and root exhibit tip-to-base gradients of sink strength. In the primary root, the sink-strength profiles reflect not only the importance of the apical meristem, but show also the starting points of cambial activity and production of secondary vascular elements. Experiments including partial removal of the source and manipulations of the sink strength indicate that translocation of pea seedlings is sink-regulated and responds rapidly to changed apoplastic conditions in the apical root region. Here, a lowered water potential leads to an increase of phloem unloading that is suggested to supply the assimilate demand for the short-term osmoregulation of affected cells via the symplasmic pathway.Abbreviation PCMBS parachloromercuribenzenesulfonic acid Discussions with Prof. R. Kollmann, Botanisches Institut, Universität Kiel, and financial support from the Deutsche Forschungsgemeinschaft are gratefully acknowledged.     

Proteomics and metabolomics analyses reveal the cucurbit sieve tube system as a complex metabolic space

The plant vascular system, and specifically the phloem, plays a pivotal role in allocation of fixed carbon to developing sink organs. Although the processes involved in loading and unloading of sugars and amino acids are well characterized, little information is available regarding the nature of other metabolites in the sieve tube system (STS) at specific sites along the pathway. Here, we elucidate spatial features of metabolite composition mapped with phloem enzymes along the cucurbit STS. Phloem sap (PS) was collected from the loading (source), unloading (apical sink region) and shoot–root junction regions of cucumber, watermelon and pumpkin. Our PS analyses revealed significant differences in the metabolic and proteomic profiles both along the source–sink pathway and between the STSs of these three cucurbits. In addition, metabolite profiles established for PS and vascular tissue indicated the presence of distinct compositions, consistent with the operation of the STS as a unique symplasmic domain. In this regard, at various locations along the STS we could map metabolites and their related enzymes to specific metabolic pathways. These findings are discussed with regard to the function of the STS as a unique and highly complex metabolic space within the plant vascular system.     

器官间关系对叶片衰老的影响

叶片衰老是整个植株生理特性的敏感表现,受根系、茎、生殖器官和其他叶片等器官的影响。器官间关系影响叶片衰老可能是通过竞争体内营养、水分等物质、竞争环境因子、源库关系、激素等信息系统调节等机制实现的。从整株水平上加强叶片衰老的生理机制和控制技术研究,将为生产上控制衰老、减少叶片异常衰老造成的产量和品质损失提供有效的技术途径。     

器官间关系对叶片衰老的影响

叶片衰老是整个植株生理特性的敏感表现,受根系、茎、生殖器官和其他叶片等器官的影响。器官间关系影响叶片衰老可能是通过竞争体内营养、水分等物质、竞争环境因子、源库关系、激素等信息系统调节等机制实现的。从整株水平上加强叶片衰老的生理机制和控制技术研究,将为生产上控制衰老、减少叶片异常衰老造成的产量和品质损失提供有效的技术途径。     

Overexpression of GmAAP6a enhances tolerance to low nitrogen and improves seed nitrogen status by optimizing amino acid partitioning in soybean

Amino acid transport via phloem is one of the major source‐to‐sink nitrogen translocation pathways in most plant species. Amino acid permeases (AAPs) play essential roles in amino acid transport between plant cells and subsequent phloem or seed loading. In this study, a soybean AAP gene, annotated as GmAAP6a, was cloned and demonstrated to be significantly induced by nitrogen starvation. Histochemical staining of GmAAP6a:GmAAP6a‐GUS transgenic soybean revealed that GmAAP6a is predominantly expressed in phloem and xylem parenchyma cells. Growth and transport studies using toxic amino acid analogs or single amino acids as a sole nitrogen source suggest that GmAAP6a can selectively absorb and transport neutral and acidic amino acids. Overexpression of GmAAP6a in Arabidopsis and soybean resulted in elevated tolerance to nitrogen limitation. Furthermore, the source‐to‐sink transfer of amino acids in the transgenic soybean was markedly improved under low nitrogen conditions. At the vegetative stage, GmAAP6a‐overexpressing soybean showed significantly increased nitrogen export from source cotyledons and simultaneously enhanced nitrogen import into sink primary leaves. At the reproductive stage, nitrogen import into seeds was greatly enhanced under both sufficient and limited nitrogen conditions. Collectively, our results imply that overexpression of GmAAP6a enhances nitrogen stress tolerance and source‐to‐sink transport and improves seed quality in soybean. Co‐expression of GmAAP6a with genes specialized in source nitrogen recycling and seed loading may represent an interesting application potential in breeding.     

Regulation by kinetin and abcisic acid of correlative senescence in relation to grain maturation,source-sink relationship and yield of rice (Oryza sativa L.)

When kinetin was applied to the source organ (flag leaf) of rice (Oryza sativa L. cv. Ratna), foliar senescence was delayed and grain yield per plant (as evidenced by grain weight, grain/straw weight ratio and 1,000 grain growth) was increased through the increase of sink activity (increase in dry weight of the grains/plant), duration of sink capacity as well as photosynthetic ability of the glumes (as determined by the chlorophyll content of the glumes of the developing grains). However, application of kinetin to the sink organs (fruits), promoted senescence of the source but increased the yield by increasing the sink capacity and 1,000 grain growth mostly at the earlier stage of reproductive development. Lower sterility percentage was associated with higher grain yield of the plant by kinetin treatments. ABA applied either to the source or the sink promoted leaf senescence and reduced the grain yield by reducing the sink activity, harvest index, sink capacity duration and increasing the sterility percentage. Thousand grain dry weight at harvest did not vary significantly amongst the treatments. It was concluded that nutrient drainage was associated with the correlative influence of fruit on the monocarpic senescence of rice plant and that a competetion for differential allocation of cytokinin and ABA in the source and sink organs initiates this senescence syndrome.     

Is polyphenol induction simply a result of altered carbon and nitrogen accumulation?

Carbon translocation in plants is shaped by phyllotaxis and regulated by source/sink interactions that respond to the demands of growth and defense. We have studied this extensively in poplar saplings, and recently showed that unlike carbon import, nitrogen is not translocated to sink leaves in response to application of jasmonic acid. Here we report that this is also true for young trees in the field. We discuss the importance of transport processes in establishing local C:N ratios, and suggest that the JA-induced flow of C but not N to sink tissues, and their corresponding increases in C-based defenses, may simply reflect a plant adaptation to handle excess reduced carbon and energy.     

源库关系改变对棉叶内源激素的影响

以中棉18号为材料,在大田条件下研究了摘蕾引起源库关系改变对主要源叶（对应的主茎叶和对位果枝叶）中内源激素的影响。结果表明;去蕾处理推迟了相应主茎叶和果枝叶中吲哚乙酸（IAA）峰出现,降低了玉米素及其核苷（Z ZR）含量,推迟了果枝叶中异戊烯基腺嘌呤及其核苷（iP iPA)的积累,提前了二氢玉米素及其核苷（DHZ DHZR）的积累,降低了脱落酸（ABA）含量。去除一个蕾对相应叶片内源激素的影响是短期的,随整株源库关系的协调,与对照趋同。激素的这种变化与叶片功能变化是一致的,表明棉花源库关系的协调不仅是物质上的关系,植物激素可能是其中重要的信息调控系统。     

An assessment of auxin-promoted transport in decapitated stems and whole shoots of Phaseolus vulgaris L.

¹⁴C-photosynthate transfer in decapitated stems of P. vulgaris plants, treated with IAA (indol-3yl-acetic acid), appeared, as ascertained by microautoradiography, to be restricted to cells of sieve-element appearance. The IAA-induced promotion of photosynthate transport was found not to depend on any artifacts caused by the decapitation procedure. Rather, decapitation primarily served the purpose of removing photosynthate sources above the point of hormone application which otherwise suppressed the expression of the IAA effect on acropetal photosynthate transport. Furthermore, by manipulating stem levels of endogenous auxins with the inhibitor of polar auxin transport, 1-(2¹-carboxyphenyl)-3-phenylpropane-1,3-dione (ACP1.55), evidence was obtained indicating that photosynthate transfer to the shoot apex depended, at least in part, on endogenous levels of auxins at site(s) remote from the apical sink (i.e. shoot apex).Abbreviations ACP1.55 1-(2¹-carboxyphenyl)-3-phenylpropane-1,3-dione - DCMU 3-(3,4-dichlorophenyl)-1,1-dimethylurea - IAA indol-3yl-acetic acid