Arabidopsis Lateral Root Development 3 is essential for early phloem development and function, and hence for normal root system development

We have identified a gene, Lateral Root Development 3 (LRD3), that is important for maintaining a balance between primary and lateral root growth. The lrd3 mutant has decreased primary root growth and increased lateral root growth. We determined that the LRD3 gene encodes a LIM-domain protein of unknown function. LRD3 is expressed only in the phloem companion cells, which suggested a role in phloem function. Indeed, while phloem loading and export from the shoot appear to be normal, delivery of phloem to the primary root tip is limited severely in young seedlings. Abnormalities in phloem morphology in these seedlings indicate that LRD3 is essential for correct early phloem development. There is a subsequent spontaneous recovery of normal phloem morphology, which is correlated tightly with increased phloem delivery and growth of the primary root. The LRD3 gene is one of very few genes described to affect phloem development, and the only one that is specific to early phloem development. Continuous growth on auxin also leads to recovery of phloem development and function in lrd3, which demonstrates that auxin plays a key role in early phloem development. The root system architecture and the pattern of phloem allocation in the lrd3 root system suggested that there may be regulated mechanisms for selectively supporting certain lateral roots when the primary root is compromised. Therefore, this study provides new insights into phloem-mediated resource allocation and its effects on plant root system architecture.     

盐胁迫对苹果器官中钙镁铁锌含量的影响

以盆栽2年生富士苹果树(砧木为平邑甜茶M.hupehensisReld)为试材,研究了盐胁迫对苹果矿质营养平衡的影响.结果表明,在盐胁迫下,苹果各器官不同时期的单位干样中Ca、Mg、Fe和Zn含量的平均值没有明显变化,但各元素与Na的比值明显下降,特别是在高盐(3‰NaCl)胁迫下下降更为明显,从而破坏了树体内元素平衡.在无盐和盐胁迫下,苹果各器官中Ca含量的顺序为主干韧皮部＞叶片、新梢＞根＞主干木质部;Mg含量为新梢、根＞主干木质部、主干韧皮部、叶片;Fe含量为根＞叶片＞主干韧皮部、新梢＞主干木质部;Zn含量为新梢＞叶片＞根、主干韧皮部＞主干木质部.与对照相比,器官中各元素含量在胁迫期间表现出不同程度的波动性.     

Thermodynamic analysis of the interaction of the xylem water and phloem sugar solution and its significance for the cohesion theory

The cohesion theory explains water transport in trees by the evaporation of water in the leaves (transpiration), which in turn generates the tension required for sap ascent, i.e. the flow of pure water from the soil through the root system and the non-living cells of the tree (xylem tracheids) up to the leaves. Only a small part of this water flow entering the leaves is used in photosynthesis to produce sugar solution, which is transported from the leaves through the living cells (phloem) to everywhere in the tree where it is needed and used. The phloem sieves are connected to the xylem tracheids by water transparent membranes, which means that the upflow of pure water and downflow of sugar solution interact with each other, causing the osmotic pressure in the sugar solution (Münch model). In this paper we analyse this interaction with a thermodynamic approach and we show that some open questions in the cohesion theory can then perhaps be better understood. For example, why under a quite high tension the water can flow in the xylem mostly without any notable cavitation, and how the suction force itself depends on the cavitation. Minimizing Gibbs energy of the system of xylem and phloem, we derive extended vapor pressure and osmotic pressure equations, which include gas bubbles in the xylem conduits as well as the cellulose-air-water interface term. With the aid of the vapor pressure equation derived here, we estimate the suction force that the cavitation controlled by the phloem sugar solution can generate at high moisture contents. We also estimate the suction force that the transpiration can generate by moisture gradient at low moisture contents. From the general osmotic pressure equation we derive an equation for calculating the degree of cavitation with different sugar solution concentrations and we show the conditions under which the cavitation in the xylem is totally avoided. Using recent field measurement results for a Scotch pine, the theory is demonstrated by showing its predictions for possible amounts of cavitation or embolism from morning hours to late afternoon.     

Sugars en route to the roots. Transport,metabolism and storage within plant roots and towards microorganisms of the rhizosphere

In plants, the root is a typical sink organ that relies exclusively on the import of sugar from the aerial parts. Sucrose is delivered by the phloem to the most distant root tips and, en route to the tip, is used by the different root tissues for metabolism and storage. Besides, a certain portion of this carbon is exuded in the rhizosphere, supplied to beneficial microorganisms and diverted by parasitic microbes. The transport of sugars toward these numerous sinks either occurs symplastically through cell connections (plasmodesmata) or is apoplastically mediated through membrane transporters (MST, mononsaccharide tranporters, SUT/SUC, H+/sucrose transporters and SWEET, Sugar will eventually be exported transporters) that control monosaccharide and sucrose fluxes. Here, we review recent progresses on carbon partitioning within and outside roots, discussing membrane transporters involved in plant responses to biotic and abiotic factors.     

Distribution and pathway for phloem-dependent movement of Melon necrotic spot virus in melon plants

The translocation of Melon necrotic spot virus (MNSV) within tissues of inoculated and systemically infected Cucumis melo L. 'Galia' was studied by tissue-printing and in situ hybridization techniques. The results were compatible with the phloem vascular components being used to spread MNSV systemically by the same assimilate transport route that runs from source to sink organs. Virus RNAs were shown to move from the inoculated cotyledon toward the hypocotyl and root system via the external phloem, whereas the upward spread through the stem to the young tissues took place via the internal phloem. Virus infection was absent from non-inoculated source tissues as well as from both shoot and root apical meristems, but active sink tissues such as the young leaves and root system were highly infected. Finally, our results suggest that the MNSV invasion of roots is due to virus replication although a destination-selective process is probably necessary to explain the high levels of virus accumulation in roots. This efficient invasion of the root system is discussed in terms of natural transmission of MNSV by the soil-borne fungal vector.     

Cytokinins in the vascular saps of Ricinus communis

Roots are recognised as the major sites of cytokinin synthesis and shoots receive a continuous supply of cytokinins from the roots. Although reports are available on the xylem mobility of putative free bases and their ribosides, relatively few studies on the phloem mobility of cytokinins have been reported. The origin of phloem-mobile cytokinins is uncertain but there is evidence which implicates a recirculation from the root source. This study is the first report in which zeatin and zeatin riboside from the root pressure exudate and phloem sap of Ricinus have been identified by full-scan GC-MS and quantified by GC-MS selective-ion-monitoring. In this study, the concentration of cytokinins in root pressure exudate was similar, but lower, and in the phloem sap higher than that reported previously. The concentration of cytokinins quantified in the phloem sap confirms their transport in the sieve tubes. The relatively high concentration of zeatin riboside detected in the root pressure exudate and of zeatin detected in the phloem sap indicate a possible vascular recirculation of these hormones.     

DEVELOPMENT AND STRUCTURE OF THE VASCULAR CONNECTION BETWEEN THE PRIMARY AND LATERAL ROOT OF LYCOPERSICON ESCULENTUM

The primary xylem connection between the diarch parent root and the diarch lateral root was derived from the pericycle and stelar parenchyma. Early in lateral root development stelar parenchyma that was positioned between the parent xylem and the primordium divided transversely. These transverse divisions produced a plate of cells, most of which subsequently differentiated into vessel element connectors. After emergence of the lateral root, xylem maturation began in the stelar vessel element connectors and maturation proceeded acropetally into the lateral root. Protoxylem of the lateral root was connected to the metaxylem of the parent root via stelar vessel element connectors. The circular phloem connection was pericyclic in origin. Axial phloem connections which vascularized the lateral root were established with sieve tube elements of both parent phloem poles. Maturation of the phloem connection occurred prior to lateral root emergence. Transaxial phloem, positioned in arches above and below the lateral root vascular cylinder, was derived from the pericycle; and each arch consisted of three to four sieve tube elements. No transfer cells were found in the transaxial phloem.     

Effects of Wounding on Transport in the Phloem

The problem dealt with was the nature of the mechanisms of therecovery of the phloem of Phaseolus vulgaris from the effectsof wounds. The influence of root removal on the transport of[¹⁴C]sucrose and of fluorescein was followed both in the directionof the root and in the opposite direction, towards the intactshoot. The removal of the root was found to cause a sharp buttemporary decrease in the transport of sucrose towards the shoot;this was interpreted as the direct influence of the wound onthe transport system. Similar results were observed concerningfluorescein transport, which also showed that the resumptionof transport was in the entire phloem and not only in newlydifferentiated elements.     

Are phloem amino acids involved in the shoot to root control of NO3- uptake in Ricinus communis plants?

The putative role of phloem amino acids as negative feedback signals for root NO3^- uptake was investigated in Ricinus communis L. The NO3^--grown plants were subjected to N-deficiency due either to complete N-deprivation, or to localized N-deprivation on one side of a split-root system. In comparison with controls, complete N-deprivation resulted in a transient increase in ¹⁵NO3^- influx, and in profound changes in downward phloem transport of amino acids. Total amino acid concentration in the phloem sap decreased by 40%, but responses markedly differed between the individual amino acids. Concentrations of Gln and Ser were rapidly lowered by 50%, while those of Val, Phe, Leu, and Ile displayed a marked increase. Localized N-deprivation on one side of the split root system also resulted in the up-regulation of ¹⁵NO3^- influx in the roots still supplied with NO3^-. However, the amino acid composition of the phloem sap directed to these roots was not modified by the treatment, and remained similar to that in N-sufficient control plants. Only amino acid transport to the N-deprived roots was affected as observed in response to complete N-deprivation. The results from split-root plants indicate that the response of root NO3^- influx to N-deficiency is controlled by shoot-borne regulatory signals, and provide a case study where these signals are not related to a qualitative change or a significant decrease in downward phloem transport of amino acids.     

Bark-stripping by Grey squirrels (Sciurus carolinensis)

To investigate why Grey squirrels strip bark from young trees, squirrel populations and tree quality were studied at 30 English Midland woods. In young beech and sycamore woodlands, the area of bark which Grey squirrels stripped from each tree was strongly related to the phloem width (volume per unit area) of the tree, and not to the phloem sugar content. Variation in bark-stripping between woods was strongly related to average phloem widths in each plantation. The extent of bark-stripping also correlated with juvenile squirrel density, but only in years with relatively little squirrel breeding. There was a tendency for damage to recur in previously damaged areas, in a pattern suggesting that squirrels had learned the habit. The results indicate that bark-stripping can be initiated by young squirrels, perhaps through agonistic gnawing or exploratory feeding, or by older squirrels which have learned the habit, but that severe damage occurs only where the tree phloem is suitable.     

Effects of nutrient solution zinc activity on net uptake,translocation, and root export of cadmium and zinc by separated sections of intact durum wheat (Triticum turgidum L. var durum) seedling roots

Cd accumulation in durum wheat presents a potential health risk to consumers. In an effort to understand the physiological mechanisms involved with Cd accumulation, this study examined the effects of Zn on Cd root uptake and phloem translocation in a split– root system. Durum wheat seedlings were grown in chelate-buffered nutrient solution with intact root systems divided into two sections. Each root section grew in a separate 1 l pot, one of which contained 0.2 μM CdSO₄. In addition, each two-pot system contained ZnSO₄ in the following combinations (in μm) (for -cd root system: +cd root system): 1:1, 1:10, 10:1,10:10, 1:19, and 19:1. Harvested plant material was analyzed for Cd and Zn. In addition, rates of Cd and Zn net uptake, translocation to the shoot, and root export (translocation from one root segment to the other) between days 8 and 22 were calculated. Results show that Zn was not translocated from one root section to its connected root section. Uptake rates of Cd increased as solution Zn concentrations increased. Cd translocation from one root section to the other decreased significantly when the Zn concentration in either pot was greater than 1 μM. These results show the potential of Zn to inhibit movement of Cd via the phloem, and suggests that providing adequate Zn levels may limit phloem loading of Cd into wheat grain. Increasing the rhizosphere activity of Zn²⁺ in Cd-containing soils may therefore result in reduced Cd accumulation in grain even while net Cd uptake is slightly enhanced. This revised version was published online in June 2006 with corrections to the Cover Date.     

Transport of resistance-inducing sterols in phloem sap of barley

After root application of [7alpha-3H]-7beta-hydroxysitosterol and [3alpha,6beta-3H2]-6alpha-hydroxylathosterol these sterols could be detected in the leaves and phloem sap feeding aphids. These results imply that the phloem sap is a sterol transport system in barley plants.     

Photoassimilate partitioning in nodulated soybean II. The effect of changes in photoassimilate availability shows that nodule permeability to gases is not linked to the supply of solutes or water

It is concluded that the permeability of the soybean nodule to gases is not linked to the supply of solutes or water via the phloem to the nodule. Nodule respiration and nitrogenase activity were less affected by diel variation and shading treatments than partitioning to the nodule, as assessed using a non-invasive ¹¹C-based technique. Thus C import to the nodule was not matched to C requirement by the nodule. Transit times of tracer to, and within, the nodulated root increased under conditions of reduced photosynthetic rate. The increase in transit time was interpreted as a reduction in the flux of phloem sap. Thus the fluxes of both water and C to the nodule decreased following a reduction in photosynthetic rate. The change in partitioning of recent photosynthate to soybean roots and nodules in response to changes in photoassimilate availability was also used to assess the 'priority' of these sinks. Partitioning from the leaf to the root system was greatly decreased when photoassimilate availability was limited, indicating that root system priority is lower than that of the shoot, as reported for other systems. However, partitioning of tracer arriving in the root system between the nodulated and non-nodulated zones of the root was not affected by changes in photoassimilate availability, as caused by diel change, shading, or steaming of branch roots. Thus although nodules are sinks of high sink 'activity', they have 'priority' equal to that of other root sinks. It is suggested that there are similar phloem unloading kinetics, despite the very different metabolic destiny of the carbohydrate within the two organs.     

Determination of phytohormones in phloem exudate from tree species by radioimmunoassay

A sensitive and specific radioimmunoassay was used to determine quantitatively four of the most important phytohormones in the phloem exudate from 14 different tree species of 8 genera. For cytokinins and indole-3-acetic acid (IAA) we found higher concentrations than those reported previously for other species. The gibberellin values were of the same order of magnitude as in earlier analyses (with different methods) of tree phloem exudates, but lower than the ones reported for Ricinus. Free abscisic acid (ABA) was found in tree phloem exudates in similar concentrations as before in Yucca or palm phloem exudate, but at considerably lower ones than reported for Ricinus and in Lupinus phloem exudate.Abbreviations IAA indole-3-acetic acid - ABA abscisic acid - GA gibberellic acid     

Carbon import into barley roots: effects of sugars and relation to cell expansion

The import of photosynthate labelled with ¹¹C from a sourceleaf into the two halves of a split root system of an intactbarley plant was studied. When applied to one half of a splitroot system sugars that are absorbed and metabolized reducesubsequent import of ¹¹C into that root half. The non-metabolizedsugar analogue 3-O-methyl glucose has no effect on import, whilstmannose and 2–deoxyglucose inhibit both root elongationand import of ¹¹C. EDTA, PCMBS, and apoplastic pH in the range4–7, have little effect on partitioning. These resultsare interpreted in terms of a suggestion that phloem unloadsdirectly into expanding cells in the elongation zone of roottips. Key words: Carbon partitioning, roots, ¹¹C, sucrose, phloem, sugars, cell expansion     

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.     

Comparative axial widening of phloem and xylem conduits in small woody plants

Key message

Along the stem axis phloem’s sieve elements increase in diameter basally at rates comparable to those of xylem conduits and in agreement with principles of hydraulic optimization.

Abstract

Plant physiology relies on the efficiency of the two long-distance transport systems of xylem and phloem. Xylem architecture comprises conduits of small dimensions towards the stem apex, where transpiration-induced tensions are the highest along the root-to-leaves hydraulic pathway, and widen basally to minimize the path length resistance to water flow. Instead, information on phloem anatomy and allometry is extremely scarce, although potentially relevant for the efficiency of sugar transportation. We measured the hydraulic diameter (Dh) of both xylem conduits and phloem sieve elements in parallel at different heights along the stem of a small tree of Picea abies, Fraxinus excelsior and Salix eleagnos. Dh increased from the stem apex to base in both xylem and phloem, with a higher scaling exponent (b) of sieve elements than that of tracheids in the conifer (0.19 vs. 0.14) and lower than that of vessels in the angiosperms (0.14–0.22 vs. 0.19–0.40). In addition, sieve elements were larger than tracheids in P. abies and narrower than angiosperms vessels at any height along the stem. In conclusion, axial conduit widening would seem to be a key feature of both xylem and phloem long-distance transport architectures.     

Real-time imaging of phloem unloading in the root tip of Arabidopsis

Confocal laser scanning microscopy (CLSM) has been used to image phloem transport and unloading in the root tip of Arabidopsis. The fluorescent probe 5(6) carboxyfluorescein (CF) was ester loaded into a single cotyledon and the entire seedling placed within an observation chamber under the microscope. Translocation of CF to the root tip was rapid, followed by unloading into discrete concentric files of cells. The position of the prominent unloading ‘zone’ corresponded precisely with that of the two protophloem files of sieve elements, demonstrating a functional role of these cells in symplastic sieve-element unloading. Symplastic transport following unloading was confined to the elongating zone of the root with little basipetal transport to more mature cells. Following photobleaching of the unloading zone, phloem transport was restored immediately into the protophloem sieve elements, followed rapidly by lateral, symplastic sieve-element unloading. The results demonstrate that phloem transport processes can now be imaged in real time, and non-invasively, within an intact plant system.