Xylem and Phloem Transport of Mineral Nutrients from Solanum tuberosum Roots

The xylem and phloem transport of mineral elements from stemnodal roots to the stem and stolon of growing potato (Solanumtuberosum L. cv. ‘Russet Burbank’) plants was investigated.Adventitious roots, originating from below-ground nodes of thestem of potato seedlings, were exposed to solutions of SrCI₂or MnSO₄. Relative elemental concentrations were measured inthe conductive tissues using energy dispersive X-ray analysis.After a 5 h daylight uptake period, Sr (a Ca-transport analogue)levels were elevated in the stem xylem tissue, but Sr did notincrease in the stem phloem, nor was it present in either ofthe conductive tissues of stolons located 1–2 nodes abovethe treated roots. In contrast, elevated levels of Cl, S, andMn were found in stolon xylem and phloem tissue during the sameperiod. The absence of Sr in the stolon after 5 h suggests thatno xylem flow into the stolon occurred during the uptake periodand, furthermore, phloem flow is responsible for the transportof the Cl, S, and Mn into the stolon. Elevated levels of thesemobile nutrients in the xylem of the stolon were attributedto xylem-to-phloem transfer in the stem or leaves, transportto the stolon in the phloem, and phloem-to-xylem transfer inthe stolon. During a 19 h uptake period, some Sr was observedin the phloem tissue of the stem, demonstrating slow exchangeof Sr with sieve elements or proximal phloem parenchyma andcompanion cells. Key words: Calcium, manganese, X-ray analysis     

Causes and consequences of sectoriality in the clonal herb Glechoma hederacea

The causes of sectoriality and consequences for clone behaviour are examined using data from the stoloniferous herb Glechoma hederacea. The proximal causes of physiological integration patterns are investigated using anatomical studies, acid fuchsin dye to reveal patterns of xylem continuity between ramets, and ¹⁴C as a label to reveal quantitative photoassimilate translocation patterns in the phloem. Dye movement in the xylem was acropetal and sectorial, and the sectoriality was determined by phyllotaxy. Patterns of ¹⁴C-labelled photoassimilate allocation were qualitatively similar to those of xylem based resources, although there was some basipetal movement of photoassimilate. The patterns of physiological integration and independence between ramets are shown to be governed by rules which depend on vascular continuity and discontinuity between ramets. Physiological support to stolon apices results in acquisition of relative branch autonomy (branches become semi-autonomous integrated physiological units, IPUs).This paper evaluates whether observed physiological integration patterns may be modified by altering normal source-sink relationships or by modifying environmental conditions. An experiment using different defoliation intensities, and different defoliation patterns at the same overall intensity, demonstrated that the precise positions of leaves removed from a clone had unique consequences for its subsequent development. Individual ramets of a given clone may be located in microhabitats of differing quality. An experiment in which competition was either present or absent throughout the space occupied by the clone, or patchy in distribution, showed that G. hederacea did not respond to competition at the whole clone level. Instead, connected stolons (IPUs) responded independently to local competition. Sectoriality may promote the restriction of lethal, localised environmental factors within the affected IPU. A study investigating the uptake and translocation of zinc by clones revealed that quantified patterns of zinc distribution resembled patterns of ¹⁴C movement in the phloem, and that there was no significant transport of zinc from one stolon to another.Although sectorial patterns of resource movement in G. hederacea can be modified in the short term, in the long-term, physiological integration may not allow this species to integrate the effects of environmental heterogeneity. A mobile clonal species with a high growth rate and relatively short-lived ramets, such as G. hederacea, is likely to benefit from a semi-autonomous response to patch quality at the level of the stolon, since the alternative of widespread intra-clonal support may increase the residence time of the clone in unfavourable pathches.     

Movement of [14C]sucrose along the stolon of Saxifraga sarmentosa

Summary The characteristics of ¹³⁷Cs transport along the stolon of Saxifraga previously reported have been confirmed for applied sucrose and natural assimilate. Long-distance transport is strictly unidirectional, with a symmetrical short-distance spread from the point of application. Only the latter takes place in a long piece of excised stolon. Transport is readily reversed when the parent plant is darkened and the daugther, plantlet allowed to photosynthesise. These findings strongly support a mass-flow mechanism for the stolon. They also confirm the value of ¹³⁷Cs as a tracer for assimilate movement, though in contrast to assimilate it suffers appreciable lateral leakage. Pulse labelling of the subtending leaf failed to produce a sharp peak of activity in the stolon. A flattening with time of the ¹⁴C profile is considered to be due to differing linear velocities in parallel sieve tubes.This work formed part of that submitted for the degree, of Ph. D. of the University of London.     

The effect of nitrogen on the movement of tracers down the stolon of Saxifraga sarmentosa,with some observations on the influence of light

Summary The movement of applied ¹³⁷Cs and of naturally-assimilated ¹⁴C down the long stolon of Saxifraga is strongly inhibited by confining a length of 10 to 30 cm of the stolon in an atmosphere of nitrogen. The inhibition is reversible, normal transport being restored after less then 4 h when the stolon is returned to air from 5 h in nitrogen. Callose formation does not seem to be involved. There is evidence that local darkness has a similar adverse effect on phloem transport.These findings are considered antagonistic to the pressure-flow hypothesis, but favourable to the active mass-flow theories.This work formed part of that submitted for the degree of Ph.D. of the University of London.     

Movement of Lucifer Yellow CH in potato tuber storage tissues: A comparison of symplastic and apoplastic transport

The fluorescent dye Lucifer Yellow CH (LYCH) was introduced directly into the symplast of potato (Solanum tuberosum L.) tuber storage parenchyma by microinjection and also into the apoplast through cuts made in the stolon cortex. Microinjected LYCH moved away rapidly from a single storage cell and spread radially via the symplast. When the microinjected tissue was subsequently fixed in glutaraldehyde and sectioned the dye was seen clearly to be localised in the cytoplasm but not in the vacuole. In comparison, when LYCH was introduced into cuts made in the stolon cortex the dye entered the tuber by the xylem and subsequently spread apoplastically. No movement of dye was observed in the phloem. In glutaraldehyde-fixed tissues, in which LYCH was introduced to the apoplast, the dye was found within xylem vessels, in the cell walls and in intercellular spaces. Wall regions, possibly associated with plasmodesmata, became stained by the dye as it moved through the apoplast. Three hours after introduction of the dye to the stolon, intense deposits of LYCH were found in the vacuoles of all cells in the tuber, many aligned along the tonoplast. Differentiating vascular parenchyma elements contained large amounts of dye within enlarging vacuoles. However, with the exception of plasmolysed and-or damaged cells, LYCH was absent from the cytoplasm following its introduction to the plasmalemma it is suggested that the most likely pathway from the cell wall to the vacuole was by endocytosis, the dye being transported across the cytoplasm in membrane-bound vesicles. Clathrin-coated vesicles were abundant in the storage cells, providing a possible endocytotic pathway for dye movement. The significance of these observations is discussed in relation to the movement of LYCH in plant tissues and to the movement of solutes within and between storage cells of the tuber.Abbreviation LYCH Lucifer Yellow CH     

<Superscript>11</Superscript>C-imaging: methyl jasmonate moves in both phloem and xylem,promotes transport of jasmonate,and of photoassimilate even after proton transport is decoupled

The long-distance transport and actions of the phytohormone methyl jasmonate (MeJA) were investigated by using the short-lived positron-emitting isotope ¹¹C to label both MeJA and photoassimilate, and compare their transport properties in the same tobacco plants (Nicotiana tabacum L.). There was strong evidence that MeJA moves in both phloem and xylem pathways, because MeJA was exported from the labeled region of a mature leaf in the direction of phloem flow, but it also moved into other parts of the same leaf and other mature leaves against the direction of phloem flow. This suggests that MeJA enters the phloem and moves in sieve tube sap along with photoassimilate, but that vigorous exchange between phloem and xylem allows movement in xylem to regions which are sources of photoassimilate. This exchange may be enhanced by the volatility of MeJA, which moved readily between non-orthostichous vascular pathways, unlike reports for jasmonic acid (which is not volatile). The phloem loading of MeJA was found to be inhibited by parachloromercuribenzenesulfonic acid (PCMBS) (a thiol reagent known to inhibit membrane transporters), and by protonophores carbonyl cyanide 3-chlorophenylhydrazone (CCCP) and 2,4-dinitrophenol (DNP) suggesting proton co-transport. MeJA was found to promote both its own transport and that of recent photoassimilate within 60 min. Furthermore, we found that MeJA can counter the inhibitory effect of the uncoupling agent, CCCP, on sugar transport, suggesting that MeJA affects the plasma membrane proton gradient. We also found that MeJA’s action may extend to the sucrose transporter, since MeJA countered the inhibitory effects of the sulfhydryl reagent, PCMBS, on the transport of photoassimilate.     

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.     

THE PHLOEM OF ETAPTERIS LECLERCQII AND BOTRYOPTERIS TRIDENTATA

The phloem of Etapteris leclercqii and Botryopteris tridentata petioles is described from Lower Pennsylvanian coal balls. Petioles of B. tridentata are characterized in transverse section by an omega-shaped xylem trace, a phloem zone which extends from 2-10 cells in width, and 2-parted cortex. Etapteris leclercqii petioles exhibit a 4–9 cell-wide phloem zone surrounding the central clepsydroid xylem mass, and a 3-parted cortex. In both taxa a 1–2 cell layer parenchyma sheath separates the xylem from the extra-xylary tissues. The phloem of both species consists of sieve elements that average about 20 μm in diam by 200 μm in length in Botryopteris, and 100 μm in length in Etapteris, with horizontal-slightly oblique end walls. In transmitted light, the radial walls of the sieve elements form an irregular reticulate pattern enclosing elliptical lighter areas. With the scanning electron microscope, these areas appear as horizontal-slightly oblique furrows on the cell wall, with many small indentations lining the furrows. These indentations, because of their regular occurrence and size (from a few fractions of a micron up to 1.0 μm in diam), are interpreted as sieve pores, and the elliptical areas that enclose them as sieve areas. The phloem of E. leclercqii and B. tridentata is compared with that described for other fossil genera and with that of extant ferns.     

Investigating the in vivo calcium transport path to developing potato tuber using 45Ca: a new concept in potato tuber calcium nutrition

Calcium is believed to be transported with water in the xylem. Consistent with this proposal, low‐transpiring organs such as potato Solanum tuberosum tubers are known to suffer from calcium deficiency. Although roots on tubers and stolons have been shown to supply water to tubers, there is no direct evidence for the calcium transport pathway to tubers. Both a xylem and a phloem transport pathway have been suggested. We investigated in vivo calcium transport to developing potato, cv. Dark Red Norland and cv. Russet Burbank, tubers using ⁴⁵Ca in a controlled environment facility. Whole plant split pot experiments allowed the placement of ⁴⁵Ca either in the main (basal) root or the tuber and stolon areas of the pot. The results showed that ⁴⁵Ca was transported to the shoot with the transpiration stream from both areas but was not re‐translocated to tubers or the main (basal) root system even 57 days after ⁴⁵Ca application. Radioactivity could only be detected in the tuber when ⁴⁵Ca was fed to the stolon and tuber area. When ⁴⁵Ca was fed to specific tubers, radioactivity was detected in the aerial shoot; however, no activity was detected in other tubers or the main (basal) roots. In another set of experiments, roots on a stolon near a tuber were precisely fed ⁴⁵Ca and Safranin O. The radioactive signal exactly overlapped the water transport pathway in the tuber marked with Safranin O dye, suggesting that water and calcium can be simultaneously transported from stolon roots to the tuber. No transport of ⁴⁵Ca across the tuber periderm was detected 8 days after ⁴⁵Ca was applied to the tuber periderm. This indicated that no significant transport of calcium occurs from the soil across the periderm. Our results provide evidence that: (1) calcium is not re‐translocated via the phloem from the aerial shoot tubers and main (basal) roots; (2) the main root system does not supply calcium to the tuber; (3) calcium is not transported across the periderm to the interior tuber tissue; (4) calcium is transported to the tuber via the xylem along with water, and the roots on the stolon associated with the tuber supply water and calcium to the developing tuber; and (5) transpirational demand is a significant determinant of calcium distribution within the plant.     

Phloem unloading via the apoplastic pathway is essential for shoot distribution of root-synthesized cytokinins

Root-synthesized cytokinins are transported to the shoot and regulate the growth, development, and stress responses of aerial tissues. Previous studies have demonstrated that Arabidopsis (Arabidopsis thaliana) ATP binding cassette (ABC) transporter G family member 14 (AtABCG14) participates in xylem loading of root-synthesized cytokinins. However, the mechanism by which these root-derived cytokinins are distributed in the shoot remains unclear. Here, we revealed that AtABCG14-mediated phloem unloading through the apoplastic pathway is required for the appropriate shoot distribution of root-synthesized cytokinins in Arabidopsis. Wild-type rootstocks grafted to atabcg14 scions successfully restored trans-zeatin xylem loading. However, only low levels of root-synthesized cytokinins and induced shoot signaling were rescued. Reciprocal grafting and tissue-specific genetic complementation demonstrated that AtABCG14 disruption in the shoot considerably increased the retention of root-synthesized cytokinins in the phloem and substantially impaired their distribution in the leaf apoplast. The translocation of root-synthesized cytokinins from the xylem to the phloem and the subsequent unloading from the phloem is required for the shoot distribution and long-distance shootward transport of root-synthesized cytokinins. This study revealed a mechanism by which the phloem regulates systemic signaling of xylem-mediated transport of root-synthesized cytokinins from the root to the shoot.

Phloem unloading via the apoplastic pathway is essential for shoot distribution and long-distance translocation of root-synthesized cytokinins from the root to the shoot through the xylem.     

<Emphasis Type="Italic">LeFRK2</Emphasis> is required for phloem and xylem differentiation and the transport of both sugar and water

It has been suggested that LeFRK2, the major fructose-phosphorylating enzyme in tomato plants, may be required for stem xylem development. Yet, we do not know if this enzyme affects the development of individual vessels, whether it affects water conductance, or whether it affects phloem development and sugar transport. Here, we show that suppression of LeFRK2 results in a significant reduction in the size of vascular cells and slows fiber maturation. The vessels in stems of LeFRK2-antisense plants are narrower than in WT plants and have thinner secondary cell walls. Although the cambium produces rounded secondary vessels, these vessels become deformed during the early stages of xylem maturation. Water conductance is then reduced in stems, roots, and leaves, suggesting that LeFRK2 influences xylem development throughout the entire vascular system. Interestingly, the build-up of positive xylem pressure under static (no-flow) conditions was also decreased. Suppression of LeFRK2 reduced the length and width of the sieve elements, as well as callose deposition. To examine the effect of LeFRK2 suppression on phloem transport, we created triple-grafted plants in which a portion of the wild-type stem was replaced with an antisense interstcok, and compared the contents of the transported sugar, sucrose, in the different portions of these stems. Sucrose contents above and within the LeFRK2-antisense interstock were significantly higher than those below the graft. These results show that the antisense interstock restricted the downward movement of sucrose, suggesting that LeFRK2 is required for both phloem and xylem development. Contribution No. 114/2009 from the Volcani Center ARO.     

Movement of virus and photoassimilate in the phloem: A comparative analysis

Recent progress in the study of short-distance (cell-to-cell) movement of plant virus, facilitated by ‘movement proteins’, has led to a resurgence of interest in long-distance virus transport in the phloem. Relatively little is known about phloem-specific barriers to virus movement or about the form in which virus enters, travels within and exits this tissue. Progress in understanding virus and photoassimilate transport is limited by a paucity of information on the substructure and properties of plasmodesmata at specific interfaces. The direction of virus movement, once it has entered the phloem, can be understood by following photoassimilate translocation, a complex and dynamic process influenced by plant growth, development and vascular topology.     

Identification of high levels of phytochelatins, glutathione and cadmium in the phloem sap of Brassica napus. A role for thiol-peptides in the long-distance transport of cadmium and the effect of cadmium on iron translocation

Phytochelatins (PCs) are glutathione-derived peptides that function in heavy metal detoxification in plants and certain fungi. Recent research in Arabidopsis has shown that PCs undergo long-distance transport between roots and shoots. However, it remains unknown which tissues or vascular systems, xylem or phloem, mediate PC translocation and whether PC transport contributes to physiologically relevant long-distance transport of cadmium (Cd) between shoots and roots. To address these questions, xylem and phloem sap were obtained from Brassica napus to quantitatively analyze which thiol species are present in response to Cd exposure. High levels of PCs were identified in the phloem sap within 24 h of Cd exposure using combined mass spectrometry and fluorescence HPLC analyses. Unexpectedly, the concentration of Cd was more than four-fold higher in phloem sap compared to xylem sap. Cadmium exposure dramatically decreased iron levels in xylem and phloem sap whereas other essential heavy metals such as zinc and manganese remained unchanged. Data suggest that Cd inhibits vascular loading of iron but not nicotianamine. The high ratios [PCs]/[Cd] and [glutathione]/[Cd] in the phloem sap suggest that PCs and glutathione (GSH) can function as long-distance carriers of Cd. In contrast, only traces of PCs were detected in xylem sap. Our results suggest that, in addition to directional xylem Cd transport, the phloem is a major vascular system for long-distance source to sink transport of Cd as PC–Cd and glutathione–Cd complexes.     

白三叶无性系植物种群整合作用格局的研究

利用酸性品红和＾１４Ｃ－标记光合同化物,分析了在维管系统上的资源运输。在木质部中染料的运输是向顶的,在韧皮部中标记的光合同化物的运输方式与木质部中酸性品红的移动是相反的,具有强烈的基性,定量分析表明,尽管有一些向顶的运输,但向基运输是占绝对优势的,但随着无性系生长时间的不同,运输将会发生改变。一个无性系的变化取决于复杂的整合作用,使它们成为许多半自发的整合生理单位（ＩＰＵｓ）,不同匍匐茎的分枝,就     

Characterization of Ricinus communis phloem profilin,RcPRO1

The mature, functional sieve tube, which forms the conduit for assimilate distribution in higher plants, is dependent upon protein import from the companion cells for maintenance of the phloem long-distance translocation system. Using antibodies raised against proteins present in the sieve-tube exudate of Ricinus communis (castor bean) seedlings, a cDNA was cloned which encoded a putative profilin, termed RcPRO1. Expression and localization studies indicated that RcPRO1 mRNA encodes a phloem profilin, with some expression occurring in epidermal, cortex, pith and xylem tissue. Purified, recombinant RcPRO1 was functionally equivalent to recombinant maize profilin ZmPRO4 in a live cell nuclear displacement assay. The apparent equilibrium dissociation constant for RcPRO1 binding to plant monomeric (G-)actin was lower than the previously characterized maize profilins. Moreover, the affinity of RcPRO1 for poly-L-proline (PLP) was significantly higher than that for recombinant maize profilins. Within the sieve-tube exudate, profilin was present in 15-fold molar excess to actin. The data suggest that actin filament formation is prevented within the assimilate stream. These results are discussed in terms of the unique physiology of the phloem.     

Light,temperature and tocopherol status influence foliar vascular anatomy and leaf function in Arabidopsis thaliana

This study addressed whether the winter annual Arabidopsis thaliana can adjust foliar phloem and xylem anatomy both differentially and in parallel. In plants acclimated to hot vs cool temperature, foliar minor vein xylem‐to‐phloem ratio was greater, whereas xylem and phloem responded concomitantly to growth light intensity. Across all growth conditions, xylem anatomy correlated with transpiration rate, while phloem anatomy correlated with photosynthetic capacity for two plant lines (wild‐type Col‐0 and tocopherol‐deficient vte1 mutant) irrespective of tocopherol status. A high foliar vein density (VD) was associated with greater numbers and cross‐sectional areas of both xylem and phloem cells per vein as well as higher rates of both photosynthesis and transpiration under high vs low light intensities. Under hot vs cool temperature, high foliar VD was associated with a higher xylem‐to‐phloem ratio and greater relative rates of transpiration to photosynthesis. Tocopherol status affected development of foliar vasculature as dependent on growth environment. The most notable impact of tocopherol deficiency was seen under hot growth temperature, where the vte1 mutant exhibited greater numbers of tracheary elements (TEs) per vein, a greater ratio of TEs to sieve elements, with smaller individual sizes of TEs, and resulting similar total areas of TEs per vein and transpiration rates compared with Col‐0 wild‐type. These findings illustrate the plasticity of foliar vascular anatomy acclimation to growth environment resulting from independent adjustments of the vasculature's components.     

Cadmium loading into potato tubers: the roles of the periderm, xylem and phloem

The mobility of Cd in potato plants (Solanum tuberosum) was examined using both short‐term radioisotopic labelling with ¹⁰⁹Cd and long‐term growth experiments in soil supplemented with Cd, with an emphasis on the pathways through which Cd is taken up by tubers. Split‐pot experiments showed that tubers and their associated stolons and stolon roots contribute only a minor fraction to the overall Cd absorption by the plant. Most of the Cd was absorbed by the basal roots. ¹⁰⁹Cd absorbed from the soil was rapidly exported to other parts of the plant, especially the stem, with significant amounts appearing in the tubers within 30 h. Application of ¹⁰⁹Cd to leaves showed that Cd can be rapidly distributed via the phloem to all tissues. The results suggest that unlike Ca, Cd has high mobility in plants in both xylem and phloem, and that stems may have an important role in transfer between these two pathways.     

The Transport of Sugar, Water, and Ions into Developing Potato Tubers

Diurnal variations in the pattern of movement of sugars, water,and ions into developing tubers of the potato (Solanum tuberosumL.) were investigated. It was demonstrated using a recordingbalance that large increases in the fresh weight of tubers occurduring a dark period of reduced transpiration. Movement of assimilated¹⁴C did not reflect similar large changes and much of the weightchange observed is considered to be fluctuations in tuber watercontent. This water was shown to be moving predominantly throughthe xylem of the stolon by introducing labelled ions, ³²P and⁸⁹Sr into the plants. ³²P, which moves in both xylem and phloem,was transported to the tuber at a constant rate whereas ⁸⁹Sr,which behaves like calcium and is relatively immobile in thephloem, only moved into the tuber during the dark period. As well as the over-all long-term diurnal fluctuations severalsmaller rapid changes were recorded in the rate of water movement.Switching from darkness to light caused a transient increasefollowed by a rapid decrease in tuber weight. Switching fromlight to darkness caused a rapid increase in tuber weight. Insome experiments small oscillations in tuber weight were recorded.The possibility of these oscillations being directly relatedto cyclic changes in transpiration is considered. The resultsare discussed in relation to solute movement within plants.