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Influence of saline irrigation on growth, ion accumulation and partitioning, and leaf gas exchange of carrot (Daucus carota L.).

Like those of many horticultural crop species, the growth and leaf gas exchange responses of carrot (Daucus carota L.) to salinity are poorly understood. In this study ion accumulation in root tissues (periderm, xylem and phloem tissues) and in leaves of different ages was assessed for carrot plants grown in the field with a low level of salinity (5.8 mM Na(+) and 7.5 mM Cl(-)) and in a glasshouse with salinity ranging from 1-80 mM. At low levels of salinity (1-7.5 mM), in both the field and glasshouse, carrot leaves accumulated high concentrations of Cl(-) (140-200 mM); these appear to be the result of a high affinity for Cl(-) uptake and a low retention of Cl(-) in the root system. However, Cl(-) uptake is under tight control, with an 80-fold increase in external salinity resulting in only a 1.5-fold change in the Cl(-) concentration of the shoot and no increase in the Cl(-) concentration of the root xylem tissue. In contrast to Cl(-), shoot Na(+) concentrations were comparatively low (30-40 mM) but increased by seven-fold when salinity was increased by 80-fold. Growth over the 56-d treatment period in the glasshouse was insensitive to salinity less than 20 mM, but at higher concentrations the yield of carrot tap roots declined by 7 % for each 10 mM increase in salinity. At low levels of salinity the accumulation of high concentrations of Cl(-) (150 mM) in carrot laminae did not appear to limit leaf gas exchange. However, photosynthesis and stomatal conductance were reduced by 38 and 53 %, respectively, for plants grown at a salinity of 80 mM compared with those grown at 1 mM. Salinity-induced reductions in both p(i) and carbon isotope discrimination (delta) were small (2.5 Pa and 1.4 per thousand, respectively, at 80 mM) indicating that the reduction in photosynthesis was only marginally influenced by CO(2) supply. At a salinity of 80 mM the photosynthetic capacity was reduced, with a 30 % reduction in the CO(2)-saturated rate of photosynthesis (A(max)) and a 40 % reduction in both the apparent rate of RuBP-carboxylase-limited CO(2) fixation (V(cmax)) and the electron transport rate limiting RuBP regeneration (J(max)). This study has shown that carrot growth and leaf gas exchange are insensitive to the high leaf Cl(-) concentrations that occur at low levels (1-7 mM) of salinity. However, growth is limited at salinity levels above 20 mM and leaf gas exchange is limited at salinity levels above 8 mM.

Biological Transport↗

Tyloses and ecophysiology of the early carboniferous progymnosperm tree Protopitys buchiana.

Trunk woods of Early Carboniferous Protopitys buchiana show the earliest example of tylose formation and the first record for a progymnosperm. Protopitys tyloses are more densely located in inner trunk woods and near growth layer boundaries. We suggest, therefore, that an altered physiological state of living ray cells, during dormancy and/or following water stress, was necessary to make the woods vulnerable to tylose formation. Coupled with the distribution and proximity of abundant wood ray parenchyma to large xylem conducting cells, the positions of conduits filled with tyloses can be interpreted as ecophysiological responses of the plant to changes in local environment. In addition, some xylem conducting cells might have functioned as vessels. Fungal hyphae are present in some tracheary cells and in some areas with tyloses, but there is no evidence for wood trauma; we conclude, therefore, that these particular cases of tyloses are probably not induced by wound trauma. Protopitys buchiana wood thus shows structure/function similarities to modern woods with vessels, such as those of dicot angiosperms. This implies that ancient and modern plant ecophysiological responses correlate well with the physical parameters of their cellular construction.

Cycadopsida↗

Comparative wood anatomy of epacrids (Styphelioideae, Ericaceae s.L.).

The wood anatomy of 16 of the 37 genera within the epacrids (Styphelioideae, Ericaceae s.l.) is investigated by light and scanning electron microscopy. Several features in the secondary xylem occur consistently at the tribal level: arrangement of vessel-ray pits, distribution of axial parenchyma, ray width, and the presence and location of crystals. The primitive nature of Prionoteae and Archerieae is supported by the presence of scalariform perforation plates with many bars and scalariform to opposite vessel pitting. The wood structure of Oligarrheneae is similar to that of Styphelieae, but the very narrow vessel elements, exclusively uniseriate rays and the lack of prismatic crystals in Oligarrheneae distinguish these two tribes. The secondary xylem of Monotoca tamariscina indicates that it does not fit in Styphelieae; a position within Oligarrheneae is possible. Like most Cosmelieae, all Richeeae are characterized by exclusively scalariform perforation plates with many bars, a very high vessel density and paratracheal parenchyma, although they clearly differ in ray width (exclusively uniseriate rays in Cosmelieae vs. uniseriate and wide multiseriate rays in Richeeae). Several wood anatomical features confirm the inclusion of epacrids in Ericaceae s.l. Furthermore, there are significant ecological implications. The small vessel diameter and high vessel frequency in many epacrids are indicative of a high conductive safety to avoid embolism caused by freeze-thaw cycles, while the replacement of scalariform by simple vessel perforation plates and an increase in vessel diameter would suggest an increased conductive efficiency, which is especially found in mesic temperate or tropical Styphelieae.

Biological Evolution↗

Transpiration, potassium uptake and flow in tobacco as affected by nitrogen forms and nutrient levels.

BACKGROUND AND AIMS: Ammonium can result in toxicity symptoms in many plants when it is supplied as the sole source of N. In this work, influences of different nitrogen forms at two levels (2 and 15 mm N) on growth, water relations and uptake and flow of potassium were studied in plants of Nicotiana tabacum 'K 326'. METHODS: Xylem sap from different leaves was collected from 106-d-old tobacco plants cultured in quartz sand by application of pressure to the root system. Whole-shoot transpiration for each of the treatments was measured on a daily basis by weight determination. KEY RESULTS: Total replacement of NO(3)(-)N by NH(4)(+)-N caused a substantial decrease in dry weight gain, even when plants grew under nutrient deficiency. Increasing nutrient concentration resulted in a greater net dry weight gain when nitrogen was supplied as NO(3)(-) or NH(4)NO(3), but resulted in little change when nitrogen was supplied as NH(4)(+). NH(4)(+)-N as the sole N-source also caused reduction in transpiration rate, changes in plant WUE (which depended on the nutrient levels) and a decrease in potassium uptake. However, the amount of xylem-transported potassium in the plants fed with NH(4)(+) was not reduced: it was 457 % or 596 % of the potassium currently taken up at low or high nutrient level, respectively, indicating a massive export from leaves and cycling of potassium in the phloem. CONCLUSIONS: Ammonium reduces leaf stomatal conductance of tobacco plants. The flow and partitioning of potassium in tobacco plants can be changed, depending on the nitrogen forms and nutrient levels.

Biological Transport↗

Selective transport of zinc, manganese, nickel, cobalt and cadmium in the root system and transfer to the leaves in young wheat plants.

BACKGROUND AND AIMS: The uptake, translocation and redistribution of the heavy metals zinc, manganese, nickel, cobalt and cadmium are relevant for plant nutrition as well as for the quality of harvested plant products. The long-distance transport of these heavy metals within the root system and the release to the shoot in young wheat (Triticum aestivum 'Arina') plants were investigated. METHODS: After the application of 65Zn, 54Mn, 63Ni, 57Co and 109Cd for 24 h to one seminal root (the other seminal roots being excised) of 54-h-old wheat seedlings, the labelled plants were incubated for several days in hydroponic culture on a medium without radionuclides. KEY RESULTS: The content of 65Zn decreased quickly in the labelled part of the root. After the transfer of 65Zn from the roots to the shoot, a further redistribution in the phloem from older to younger leaves was observed. In contrast to 65Zn, 109Cd was released more slowly from the roots to the leaves and was subsequently redistributed in the phloem to the youngest leaves only at trace levels. The content of 63Ni decreased quickly in the labelled part of the root, moving to the newly formed parts of the root system and also accumulating transiently in the expanding leaves. The 54Mn content decreased quickly in the labelled part of the root and increased simultaneously in leaf 1. A strong retention in the labelled part of the root was observed after supplying 57Co. CONCLUSIONS: The dynamics of redistribution of 65Zn, 54Mn, 63Ni, 57Co and 109Cd differed considerably. The rapid redistribution of 63Ni from older to younger leaves throughout the experiment indicated a high mobility in the phloem, while 54Mn was mobile only in the xylem and 57Co was retained in the labelled root without being loaded into the xylem.

Biological Transport, Active↗

Stem photosynthesis not pressurized ventilation is responsible for light-enhanced oxygen supply to submerged roots of alder (Alnus glutinosa).

BACKGROUND AND AIMS: Claims that submerged roots of alder and other wetland trees are aerated by pressurized gas flow generated in the stem by a light-induced thermo-osmosis have seemed inconsistent with root anatomy. Our aim was to seek a verification using physical root-stem models, stem segments with or without artificial roots, and rooted saplings. METHODS: Radial O2 loss (ROL) from roots was monitored polarographically as the gas space system of the models, and stems were pressurized artificially. ROL and internal pressurization were also measured when stems were irradiated and the xylem stream was either CO2 enriched or not. Stem photosynthesis and respiration were measured polarographically. Stem and root anatomy were examined by light and fluorescence microscopy. KEY RESULTS: Pressurizing the models and stems to <or=10 kPa, values much higher than those reportedly generated by thermo-osmosis, created only a negligible density-induced increase in ROL, but ROL increased rapidly when ambient O2 concentrations were raised. Internal pressures rose by several kPa when shoots were exposed to high light flux and ROL increased substantially, but both were due to O2 accumulation from stem photosynthesis using internally sourced CO2. Increased stem pressures had little effect on O2 transport, which remained largely diffusive. Oxygen flux from stems in high light periods indicated a net C gain by stem photosynthesis. Chloroplasts were abundant in the secondary cortex and secondary phloem, and occurred throughout the secondary xylem rays and medulla of 3-year-old stems. Diurnal patterns of ROL, most marked when light reached submerged portions of the stem, were modified by minor variations in light flux and water level. Low root temperatures also helped improve root aeration. CONCLUSIONS: Pressurized gas flow to submerged roots does not occur to any significant degree in alder, but stem photosynthesis, using internally sourced CO2 from respiration and the transpiration stream, may play an important role in root aeration in young trees and measurably affect the overall carbon balance of this and other species.

Alnus↗

Imaging and quantifying carbohydrate transport to the developing ovaries of maize.

BACKGROUND AND AIMS: Shade or inadequate water can inhibit photosynthesis and limit the development of maize (Zea mays) ovaries around the time of pollination, potentially reducing the number of kernels at harvest. This study investigated whether the decreased photosynthesis diminished only the sugar supply or also altered the transport path to the ovaries. METHODS: Photosynthesis and water potentials (Psiw) were measured in the leaves while dry matter delivery was monitored in the ovaries. Ovary glucose, starch and acid invertase activities were measured in situ. Stems were fed xylem-mobile safranin or phloem-mobile carboxyfluorescein (CF), and the dye transport to the ovaries was determined. KEY RESULTS: Under normal conditions, the ovaries gained in dry mass, and starch accumulated in the pedicel and ovary wall. Glucose accumulated in the pedicel, apparently in the apoplast where insoluble (cell-wall-bound) acid invertase acted on the arriving sucrose. A glucose gradient developed from pedicel to nucellus. Safranin moved in the xylem and did not reach the ovary, but CF moved in the phloem and arrived at the ovary. CF also spread into the pedicel but unlike glucose it did not enter the nucellus. Low Psiw or shade decreased leaf photosynthesis, ovary dry mass accumulation, invertase activities, pedicel glucose, starch accumulation and CF delivery. Removal of these treatments reversed the effects. CONCLUSIONS: The success of CF in tracing the general path and rate of carbohydrate transport gave visual evidence that phloem transport to the ovary decreased at low Psiw or in the shade but otherwise remained functional. The decreases indicated that losses in carbohydrate delivery are central features of failed ovary development under these conditions. The selectivity of transport into the nucellus resembled the situation later when embryo and endosperm are present and selective uptake occurs from the apoplast.

Acids↗

Anatomical significance of the hygrochastic movement in Anastatica hierochuntica.

BACKGROUND AND AIMS: Unlike the dispersal mechanisms of many desert plants, the whole dead skeleton of Anastatica hierochuntica is involved in seed dispersal and preservation. This process depends on the hygrochastic nature of the lignified conducting tissue that bends when dry and straightens under wet conditions. An anatomical interpretation of this mechanical movement was investigated. METHODS: An anatomical study of the stem was conducted on the juvenile plants raised under different water treatments and on the branch-orders of adult A. hierochuntica size-classes. KEY RESULTS: In the juvenile stem of A. hierochuntica, the area of cortex, conducting tissue and pith increased with water availability. However, the hydraulic conductance decreased, resulting in a better withdrawal of water in water-stressed plants. The anatomical investigation of the hygrochastic mechanism revealed an asymmetric distribution of the cortical tissues, with the conducting tissues of the stem of juvenile and adult plants being larger in the lower side. The hydraulic conductance was better in the basal and middle branch-orders than the terminal ones, permitting better conductance of water to the subsequent branch-orders. CONCLUSIONS: The lignified conducting tissue of the whole stem, having a hygrochastic nature, controls the movement of the branches. The greater amount of conducting tissue associated with a higher density of wide xylem vessels was observed in the lower side of the stem as compared with the upper side. Consequently, the conducting tissue in the lower side of the stem was suggested to be more effective in the opening process of the curled dry branches through better and more rapid conductance of water. Alternatively, due to the few narrow xylem vessels in the upper side of the stem, it was likely that the conducting tissue in the upper side is more effective in the closing process by providing more rapid drying. The mechanical rise of water and the related hygrochastic efficiency were maximized in the basal and middle branch-orders that are mostly involved in the mechanical movement.

Acclimatization↗

Effect of local heating and cooling on cambial activity and cell differentiation in the stem of Norway spruce (Picea abies).

UNLABELLED: BACKGROUND AND AIMS The effect of heating and cooling on cambial activity and cell differentiation in part of the stem of Norway spruce (Picea abies) was investigated. METHODS: A heating experiment (23-25 degrees C) was carried out in spring, before normal reactivation of the cambium, and cooling (9-11 degrees C) at the height of cambial activity in summer. The cambium, xylem and phloem were investigated by means of light- and transmission electron microscopy and UV-microspectrophotometry in tissues sampled from living trees. KEY RESULTS: Localized heating for 10 d initiated cambial divisions on the phloem side and after 20 d also on the xylem side. In a control tree, regular cambial activity started after 30 d. In the heat-treated sample, up to 15 earlywood cells undergoing differentiation were found to be present. The response of the cambium to stem cooling was less pronounced, and no anatomical differences were detected between the control and cool-treated samples after 10 or 20 d. After 30 d, latewood started to form in the sample exposed to cooling. In addition, almost no radially expanding tracheids were observed and the cambium consisted of only five layers of cells. Low temperatures reduced cambial activity, as indicated by the decreased proportion of latewood. On the phloem side, no alterations were observed among cool-treated and non-treated samples. CONCLUSIONS: Heating and cooling can influence cambial activity and cell differentiation in Norway spruce. However, at the ultrastructural and topochemical levels, no changes were observed in the pattern of secondary cell-wall formation and lignification or in lignin structure, respectively.

Cell Differentiation↗

Transmission of Xylella fastidiosa to grapevines by Homalodisca coagulata (Hemiptera: Cicadellidae).

Pierce's disease (PD) of grapevines is caused by a xylem-limited bacterium Xylella fastidiosa (Wells, Raju, Hung, Weisburg, Mandelco-Paul, and Brenner) that is transmitted to plants by xylem sap-feeding insects. The introduction of the sharpshooter leafhopper Homalodisca coagulata (Say) into California has initiated new PD epidemics in southern California. In laboratory experiments, the major characteristics of H. coagulata's transmission of X. fastidiosa to grapevines were the same as reported for other vectors: short or absent latent period; nymphs transmitted but lost infectivity after molting and regained infectivity after feeding on infected plants; and infectivity persisted in adults. Adult H. coagulata acquired and inoculated X. fastidiosa in <1 h of access time on a plant. Inoculation rates increased with access time, but acquisition efficiency (20% per individual) did not increase significantly beyond 6-h access. Estimated inoculation efficiency per individual per day was 19.6, 17.9, and 10.3% for experiments where plant access was 1, 2, and 4 d, respectively. Freshly molted adults and nymphs acquired and transmitted X. fastidiosa more efficiently than did older, field-collected insects. H. coagulata transmitted X. fastidiosa to 2-yr-old woody tissues of grapevines as efficiently as to green shoots. H. coagulata transmitted X. fastidiosa 3.5 mo after acquisition, demonstrating persistence of infectivity in adults. About half (14/29) of the H. coagulata from which we failed to culture X. fostidiosa from homogenized heads (with a detection threshold of 265 CFU/head) transmitted the pathogen to grape, and 17 of 24 from which we cultured X. fastidiosa transmitted.

Animals↗

pH, abscisic acid and the integration of metabolism in plants under stressed and non-stressed conditions: cellular responses to stress and their implication for plant water relations.

A paradigm for the response of plants to stress is presented which suggests that plants move towards a state of minimal metabolic activity as a stress intensifies and remain in that state until that stress is relieved. The paradigm is based on the proposition that cells that interface with the transpiration stream employ variations on the following theme to move towards that state. Tension on the apoplastic water opens a mechanosensitive Ca2+ channel, a response that is augmented by apoplastic ABA. The resulting elevated cytoplasmic Ca2+ deactivates a plasmalemma H+/ATPase and also activates a K(+)-H+ symport. The inflow of K+ and H+ depolarizes the membrane and renders the apoplast less acidic, the protons being removed to the vacuole and the K+ ions being re-exported via the K+ outward rectifying channel. The onset of darkness in guard and mesophyll cells deactivates the plasmalemma H+/ATPase and then the events outlined above ensue except that these cells do not appear to utilize either Ca2+ or ABA during these changes. In stressed cells it is proposed that elevated cytoplasmic Ca2+ activates the release of an ABA precursor from a stored form. ABA is then released in the apoplast after export of the precursor if the activity of the K(+)-H+ symport has brought the apoplastic pH close to 7.0. It is proposed that aquaporins in the xylem parenchyma and mesophyll cells are opened by elevated cytoplasmic Ca2+ when the water potential of the transpiration stream is high so that water can be stored in the 'xylem parenchyma reservoir'. The water in this reservoir is then used to increase the water potential in the transpiration stream when the water column is under tension and to help repair embolisms by a mechanism that resembles stomatal closure.

Abscisic Acid↗

Water uptake by roots: effects of water deficit.

The variable hydraulic conductivity of roots (Lp(r)) is explained in terms of a composite transport model. It is shown how the complex, composite anatomical structure of roots results in a composite transport of both water and solutes. In the model, the parallel apoplastic and cell-to-cell (symplastic and transcellular) pathways play an important role as well as the different tissues and structures arranged in series within the root cylinder (epidermis, exodermis, cortex, endodermis, stelar parenchyma). The roles of Casparian bands and suberin lamellae in the root's endo- and exodermis are discussed. Depending on the developmental state of these apoplastic barriers, the overall hydraulic resistance of roots is either more evenly distributed across the root cylinder (young unstressed roots) or is concentrated in certain layers (exo- and endodermis in older stressed roots). The reason for the variability of root Lp(r), is that hydraulic forces cause a dominating apoplastic flow of water around protoplasts, even in the endodermis and exodermis. In the absence of transpiration, water flow is osmotic in nature which causes a high resistance as water passes across many membranes on its passage across the root cylinder. The model allows for a high capability of roots to take up water in the presence of high rates of transpiration (high demands for water from the shoot). By contrast, the hydraulic conductance is low, when transpiration is switched off. Overall, this results in a non-linear relationship between water flow and forces (gradients of hydrostatic and osmotic pressure) which is otherwise hard to explain. The model allows for special root characteristics such as a high hydraulic conductivity (water permeability) in the presence of a low permeability of nutrient ions once taken up into the stele by active processes. Low root reflection coefficients are in line with the idea of some apoplastic bypasses for water within the root cylinder. According to the composite transport model, the switch from the hydraulic to the osmotic mode is purely physical. In the presence of heavily suberized roots, the apoplastic component of water flow may be too small. Under these conditions, a regulation of radial water flow by water channels dominates. Since water channels are under metabolic control, this component represents an 'active' element of regulation. Composite transport allows for an optimization of the water balance of the shoot in addition to the well-known phenomena involved in the regulation of water flow (gas exchange) across stomata. The model is employed to explain the responses of plants to water deficit and other stresses. During water deficit, the cohesion-tension mechanism of the ascent of sap in the xylem plays an important role. Results are summarized which prove the validity of the coehesion/tension theory. Effects of the stress hormone abscisic acid (ABA) are presented. They show that there is an apoplastic component of the flow of ABA in the root which contributes to the ABA signal in the xylem. On the other hand, (+)-cis-trans-ABA specifically affects both the cell level (water channel activity) and water flow driven by gradients in osmotic pressure at the root level which is in agreement with the composite transport model. Hydraulic water flow in the presence of gradients in hydrostatic pressure remains unchanged. The results agree with the composite transport model and resemble earlier findings of high salinity obtained for the cell (Lp) and root (Lp(r)) level. They are in line with known effects of nutrient deprivation on root Lp(r )and the diurnal rhythm of root Lp(r )recently found in roots of LOTUS.

Abscisic Acid↗

Sensitivity of growth of roots versus leaves to water stress: biophysical analysis and relation to water transport.

Water transport is an integral part of the process of growth by cell expansion and accounts for most of the increase in cell volume characterizing growth. Under water deficiency, growth is readily inhibited and growth of roots is favoured over that of leaves. The mechanisms underlying this differential response are examined in terms of Lockhart's equations and water transport. For roots, when water potential (psi) is suddenly reduced, osmotic adjustment occurs rapidly to allow partial turgor recovery and re-establishment of psi gradient for water uptake, and the loosening ability of the cell wall increases as indicated by a rapid decline in yield-threshold turgor. These adjustments permit roots to resume growth under low psi. In contrast, in leaves under reductions in psi of similar magnitude, osmotic adjustment occurs slowly and wall loosening ability either does not increase substantially or actually decreases, leading to marked growth inhibition. The growth region of both roots and leaves are hydraulically isolated from the vascular system. This isolation protects the root from low psi in the mature xylem and facilitates the continued growth into new moist soil volume. Simulations with a leaky cable model that includes a sink term for growth water uptake show that growth zone psi is barely affected by soil water removal through transpiration. On the other hand, hydraulic isolation dictates that psi of the leaf growth region would be low and subjected to further reduction by high evaporative demand. Thus, a combination of transport and changes in growth parameters is proposed as the mechanism co-ordinating the growth of the two organs under conditions of soil moisture depletion. The model simulation also showed that roots behave as reversibly leaky cable in water uptake. Some field data on root water extraction and vertical profiles of psi in shoots are viewed as manifestations of these basic phenomena. Also discussed is the trade-off between high xylem conductance and strong osmotic adjustment.

Biological Transport↗

Hormonal changes induced by partial rootzone drying of irrigated grapevine.

Partial rootzone drying (PRD) is a new irrigation technique which improves the water use efficiency (by up to 50%) of wine grape production without significant crop reduction. The technique was developed on the basis of knowledge of the mechanisms controlling transpiration and requires that approximately half of the root system is always maintained in a dry or drying state while the remainder of the root system is irrigated. The wetted and dried sides of the root system are alternated on a 10-14 d cycle. Abscisic acid (ABA) concentration in the drying roots increases 10-fold, but ABA concentration in leaves of grapevines under PRD only increased by 60% compared with a fully irrigated control. Stomatal conductance of vines under PRD irrigation was significantly reduced when compared with vines receiving water to the entire root system. Grapevines from which water was withheld from the entire root system, on the other hand, show a similar reduction in stomatal conductance, but leaf ABA increased 5-fold compared with the fully irrigated control. PRD results in increased xylem sap ABA concentration and increased xylem sap pH, both of which are likely to result in a reduction in stomatal conductance. In addition, there was a reduction in zeatin and zeatin-riboside concentrations in roots, shoot tips and buds of 60, 50 and 70%, respectively, and this may contribute to the reduction in shoot growth and intensified apical dominance of vines under PRD irrigation. There is a nocturnal net flux of water from wetter roots to the roots in dry soil and this may assist in the distribution of chemical signals necessary to sustain the PRD effect. It was concluded that a major effect of PRD is the production of chemical signals in drying roots that are transported to the leaves where they bring about a reduction in stomatal conductance.

Plant Growth Regulators↗

Solute balance of a maize (Zea mays L.) source leaf as affected by salt treatment with special emphasis on phloem retranslocation and ion leaching.

Strategies for avoiding ion accumulation in leaves of plants grown at high concentration of NaCl (100 mol m(-3)) in the rooting media, i.e. retranslocation via the phloem and leaching from the leaf surface, were quantified for fully developed leaves of maize plants cultivated hydroponically with or without salt, and with or without sprinkling (to induce leaching). Phloem sap, apoplastic fluid, xylem sap, solutes from leaf and root tissues, and the leachate were analysed for carbohydrates, amino acids, malate, and inorganic ions. In spite of a reduced growth rate Na(+) and Cl(-) concentrations in the leaf apoplast remained relatively low (about 4-5 mol m(-3)) under salt treatment. Concentrations of Na(+) and Cl(-) in the phloem sap of salt-treated maize did not exceed 12 and 32 mol m(-3), respectively, and thus remained lower than described for other species. However, phloem transport rates of these ions were higher than reported for other species. The relatively high translocation rate of ions found in maize may be due to the higher carbon translocation rate observed for C(4) plants as opposed to C(3) plants. Approximately 13-36% of the Na(+) and Cl(-) imported into the leaves through the xylem were exported by the phloem. It is concluded that phloem transport plays an important role in controlling the NaCl content of the leaf in maize. Surprisingly, leaching by artificial rain did not affect plant growth. Ion concentrations in the leachate were lower than reported for other plants but increased with NaCl treatment.

Biological Transport↗

Hydraulic vulnerability, vessel refilling, and seasonal courses of stem water potential of Sorbus aucuparia L. and Sambucus nigra L.

Differences in the seasonal variation in stem water potential between the two shrub species Sorbus aucuparia and Sambucus nigra were related with their vulnerability to xylem cavitation. It was also demonstrated indirectly that the two species differ in the extent to which they reverse cavitation. Seasonal variation in stem water potential was investigated during three growing seasons with in situ stem psychrometers. Sorbus experienced wide water potential variations and reached a minimum of -4.2 MPa during drought. Under the same microclimatic conditions, Sambucus experienced consistent stem water potentials with a minimum of -1.7 MPa. The relationship between percentage loss in hydraulic conductivity (PLC) and water potential (hydraulic vulnerability curve) of the two species differed in shape: a flat curve with nearly total loss of conductivity at -6 MPa was found for SORBUS: Sambucus showed a steep vulnerability curve with 90% loss conductivity at -2.2 MPa. Thus, Sambucus is extremely vulnerable to cavitation, but Sorbus is an almost invulnerable species. This different cavitation resistance adjusted the ranges of field stem water potential that the species experienced. Finally, seasonal courses of naturally occurring (native) embolism were compared with calculated PLC courses. This comparison indicates that Sorbus did not refill embolized xylem vessels whereas Sambucus reversed embolism. It was concluded that species which are highly vulnerable to cavitation and drought-induced embolism need refilling of embolized vessels as well as isohydric water potential patterns as two strategies of survival.

Biological Transport↗

Solute flux into parasitic plants.

Parasitic plants form intimate contacts with host tissue in order to gain access to host solutes. There are a variety of cell types within the host which parasitic plants could access to extract solutes. Depending on the degree to which the parasite has embraced the parasitic lifestyle, the extent of solute flux and the pathways used to transfer solutes from host to parasite will vary. To date, a variety of experimental approaches argue for diversity in the mechanisms and the routes by which parasites accumulate host solutes. Contact between host and parasite ranges from direct lumen-to-lumen links between host and parasite xylem and continuity between the sieve elements of host and parasite, to the involvement of transfer cells between host and parasite. Progress has been slow since Solms-Laubach distinguished types of parasitic plants that fed from host phloem or xylem in 1867, but advances in clearly delineating the pathways that link host and parasite should now be possible using fluorescent proteins expressed and restricted to particular cell types of the host. This will initially necessitate using Arabidopsis, but should allow the types of connection, i.e. symplasmic or apoplasmic, to be determined and then the identification of parasite transporters responsible for solute flux.

Adaptation, Physiological↗

Partitioning of nutrient transport processes in roots.

Roots have a range of cell types that each contribute to the acquisition of nutrients and their subsequent transfer to the xylem. The activities of these cells must be co-ordinated to ensure that delivery of nutrients to the shoot occurs at a rate that matches the demands of growth. The partitioning of transport processes between different cell types is thus essential for roots to function effectively. This partitioning is considered at the level of proteins, organelles and cells in relation to the accepted concepts of how nutrients are taken up by roots and delivered to the xylem. Using K+ as an example, the evidence underpinning current concepts is examined, gaps in understanding identified and the contribution of some new approaches assessed.

Biological Transport, Active↗