Changes in root hydraulic conductivity in wheat (Triticum aestivum L.) in response to salt stress and day/night can best be explained through altered activity of aquaporins.

Salt stress reduces plant water flow during day and night. It is not known to which extent root hydraulic properties change in parallel. To test this idea, hydroponically grown wheat plants were grown at four levels of salt stress (50, 100, 150 and 200 mM NaCl) for 5-8d before harvest (d14-18) and subjected to a range of analyses to determine diurnal changes in hydraulic conductivity (Lp) at cell, root and plant level. Cell pressure probe analyses showed that the Lp of cortex cells was differentially affected by salt stress during day and night, and that the response to salt stress differed between the main axis of roots and lateral roots. The Aquaporin (AQP) inhibitor H2 O2 reduced Lp to a common, across treatments, level as observed in salt-stressed plants during the night. Analyses of transpiring plants and exuding root systems provided values of root Lp which were in the same range as values modeled based on cell-Lp. The results can best be explained through a change in root Lp in response to salt stress and day/night, which results from an altered activity of AQPs. qPCR gene expression analyses point to possible candidate AQP isoforms.

Diurnal changes in apoplast bypass flow of water and ions in salt-stressed wheat (Triticum aestivum L.) and barley (Hordeum vulgare L.).

The aim of the present study was to quantify the contribution of apoplastic bypass flow to the uptake of water and salt across the root cylinder of wheat and barley during day and night. Plants were grown on hydroponics until they were 14-17 days old and then analysed over a single day (16 h) or night (8 h) period while being exposed to different concentrations of NaCl (50, 100, 150 and 200 mM NaCl). Exposure to salt started just before the experiment (short-term stress) or had started 6d before (longer-term stress). Bypass flow was quantified using the apoplastic tracer dye 8-hydroxy-1,3,6-pyrenesulphonic acid (PTS). The percent contribution of bypass flow to root water uptake increased in response to salt stress and during the night and amounted to up to 4.4%. Bypass flow across the root cylinder of Na+ and Cl- made up 2%-12% of the net delivery of these ions to the shoot; this percentage changed little (wheat) or decreased (barley) during the night. Changes in the contribution of bypass flow to the net uptake of water, Na+ and Cl- in response to salt stress and day/night are the combined result of changes in xylem tension, the contribution of alternative cell-to-cell flow path and the requirement to generate xylem osmotic pressure.

Does night-time transpiration provide any benefit to wheat (Triticum aestivum L.) plants which are exposed to salt stress?

The study aimed to test whether night-time transpiration provides any potential benefit to wheat plants which are subjected to salt stress. Hydroponically grown wheat plants were grown at four levels of salt stress (50, 100, 150, and 200 mM NaCl) for 5-8 days prior to harvest (day 14-18). Salt stress caused large decreases in transpiration and leaf elongation rates during day and night. The quantitative relation between the diurnal use of water for transpiration and leaf growth was comparatively little affected by salt. Night-time transpirational water loss occurred predominantly through stomata in support of respiration. Diurnal gas exchange and leaf growth were functionally linked to each other through the provision of resources (carbon, energy) and an increase in leaf surface area. Diurnal rates of water use associated with leaf cell expansive growth were highly correlated with the water potential of the xylem, which was dominated by the tension component. The tissue-specific expression level of nine candidate aquaporin genes in elongating and mature leaf tissue was little affected by salt stress or day/night changes. Growing plants under conditions of reduced night-time transpirational water loss by increasing the relative humidity (RH) during the night to 95% had little effect on the growth response to salt stress, nor was the accumulation of Na+ and Cl- in shoot tissue altered. We conclude that night-time gas exchange supports the growth in leaf area over a 24 h day/night period. Night-time transpirational water loss neither decreases nor increases the tolerance to salt stress in wheat.

Salt Stress-Regulation of Root Water Uptake in a Whole-Plant and Diurnal Context.

This review focuses on the regulation of root water uptake in plants which are exposed to salt stress. Root water uptake is not considered in isolation but is viewed in the context of other potential tolerance mechanisms of plants-tolerance mechanisms which relate to water relations and gas exchange. Plants spend between one third and half of their lives in the dark, and salt stress does not stop with sunset, nor does it start with sunrise. Surprisingly, how plants deal with salt stress during the dark has received hardly any attention, yet any growth response to salt stress over days, weeks, months and years is the integrative result of how plants perform during numerous, consecutive day/night cycles. As we will show, dealing with salt stress during the night is a prerequisite to coping with salt stress during the day. We hope to highlight with this review not so much what we know, but what we do not know; and this relates often to some rather basic questions.

Effect of Salinity on Stomatal Conductance, Leaf Hydraulic Conductance, HvPIP2 Aquaporin, and Abscisic Acid Abundance in Barley Leaf Cells.

The stomatal closure of salt-stressed plants reduces transpiration bringing about the maintenance of plant tissue hydration. The aim of this work was to test for any involvement of aquaporins (AQPs) in stomatal closure under salinity. The changes in the level of aquaporins in the cells were detected with the help of an immunohistochemical technique using antibodies against HvPIP2;2. In parallel, leaf sections were stained for abscisic acid (ABA). The effects of salinity were compared to those of exogenously applied ABA on leaf HvPIP2;2 levels and the stomatal and leaf hydraulic conductance of barley plants. Salinity reduced the abundance of HvPIP2;2 in the cells of the mestome sheath due to it being the more likely hydraulic barrier due to the deposition of lignin, accompanied by a decline in the hydraulic conductivity, transpiration, and ABA accumulation. The effects of exogenous ABA differed from those of salinity. This hormone decreased transpiration but increased the shoot hydraulic conductivity and PIP2;2 abundance. The difference in the action of the exogenous hormone and salinity may be related to the difference in the ABA distribution between leaf cells, with the hormone accumulating mainly in the mesophyll of salt-stressed plants and in the cells of the bundle sheaths of ABA-treated plants. The obtained results suggest the following succession of events: salinity decreases water flow into the shoots due to the decreased abundance of PIP2;2 and hydraulic conductance, while the decline in leaf hydration leads to the production of ABA in the leaves and stomatal closure.

Salt stress reduces root water uptake in barley (Hordeum vulgare L.) through modification of the transcellular transport path.

The aim of the study was to understand the hydraulic response to salt stress of the root system of the comparatively salt-tolerant crop barley (Hordeum vulgare L.). We focused on the transcellular path of water movement across the root cylinder that involves the crossing of membranes. This path allows for selective water uptake, while excluding salt ions. Hydroponically grown plants were exposed to 100 mM NaCl for 5-7 days and analysed when 15-17 days old. A range of complementary and novel approaches was used to determine hydraulic conductivity (Lp). This included analyses at cell, root and plant level and modelling of water flow. Apoplastic barrier formation and gene expression level of aquaporins (AQPs) was analysed. Salt stress reduced the Lp of root system through reducing water flow along the transcellular path. This involved changes in the activity and gene expression level of AQPs. Modelling of root-Lp showed that the reduction in root-Lp did not require added hydraulic resistances through apoplastic barriers at the endodermis. The bulk of data points to a near-perfect semi-permeability of roots of control plants (solute reflection coefficient σ ~1.0). Roots of salt-stressed plants are almost as semi-permeable (σ > 0.8).

Energy costs of salinity tolerance in crop plants: night-time transpiration and growth.

Plants grow and transpire during the night. The aim of the present work was to assess the relative flows of carbon, water and solutes, and the energy involved, in sustaining night-time transpiration and leaf expansive growth under control and salt-stress conditions. Published and unpublished data were used, for barley plants grown in presence of 0.5-1 mM NaCl (control) and 100 mM NaCl. Night-time leaf growth presents a more efficient use of taken-up water compared with day-time growth. This efficiency increases several-fold with salt stress. Night-time transpiration cannot be supported entirely through osmotically driven uptake of water through roots under salt stress. Using a simple three- (root medium/cytosol/vacuole) compartment approach, the energy required to support cell expansion during the night is in the lower percentage region (0.03-5.5%) of the energy available through respiration, under both, control and salt-stress conditions. Use of organic (e.g. hexose equivalents) rather than inorganic (e.g. Na+ , Cl- , K+ ) solutes for generation of osmotic pressure in growing cells, increases the energy demand by orders of magnitude, yet requires only a small portion of carbon assimilated during the day. Night-time transpiration and leaf expansive growth should be considered as a potential acclimation mechanism to salinity.

Energy costs of salt tolerance in crop plants.

Agriculture is expanding into regions that are affected by salinity. This review considers the energetic costs of salinity tolerance in crop plants and provides a framework for a quantitative assessment of costs. Different sources of energy, and modifications of root system architecture that would maximize water vs ion uptake are addressed. Energy requirements for transport of salt (NaCl) to leaf vacuoles for osmotic adjustment could be small if there are no substantial leaks back across plasma membrane and tonoplast in root and leaf. The coupling ratio of the H+ -ATPase also is a critical component. One proposed leak, that of Na+ influx across the plasma membrane through certain aquaporin channels, might be coupled to water flow, thus conserving energy. For the tonoplast, control of two types of cation channels is required for energy efficiency. Transporters controlling the Na+ and Cl- concentrations in mitochondria and chloroplasts are largely unknown and could be a major energy cost. The complexity of the system will require a sophisticated modelling approach to identify critical transporters, apoplastic barriers and root structures. This modelling approach will inform experimentation and allow a quantitative assessment of the energy costs of NaCl tolerance to guide breeding and engineering of molecular components.

Photosynthetically active radiation impacts significantly on root and cell hydraulics in barley (Hordeum vulgare L.).

Photosynthetically active radiation (PAR) affects transpirational water loss, yet we do not know through which mechanisms root water uptake is adjusted in parallel. Here, we exposed hydroponically grown barley plants to three levels of PAR [Normal (control), Low, High] and focused on the role which aquaporins (AQPs), apoplastic barriers (Casparian bands, suberin lamellae) and root morphology play in the adjustment of root hydraulic conductivity (Lp). Plants were analyzed when they were 14-18 days (d) old. Root and cell Lp, which involves AQP activity, was determined through exudation and cell pressure probe measurements, respectively. Gene expression of AQPs was analyzed through qPCR. The formation of apoplastic barriers was studied through staining of cross-sections. The rate of transpirational water loss per plant and unit leaf area increased in response to high-PAR and decreased in response to low-PAR treatments, both during day and night. Hydraulic conductivity in roots decreased significantly at organ and cell level in response to Low-PAR, and increased (organ) or did not change (cell level) in response to High-PAR. The formation of apoplastic barriers was little affected by PAR. Gene expression of AQPs tended to be highest in the Low-PAR treatment. Lateral roots, showing few apoplastic barriers, contributed the least in Low- and the most to root surface area in High-PAR plants. It is concluded that barley plants which experience changes in shoot transpirational water loss in response to PAR adjust root water uptake through changes in root Lp, and that these changes are mediated through altered AQP activity and root morphology.

Apoplastic barriers, aquaporin gene expression and root and cell hydraulic conductivity in phosphate-limited sheepgrass plants.

Mineral nutrient supply can affect the hydraulic property of roots. The aim of the present work on sheepgrass (Leymus chinensis L.) plants was to test whether any changes in root hydraulic conductivity (Lp; exudation analyses) in response to a growth-limiting supply of phosphate (P) are accompanied by changes in (1) cell Lp via measuring the cell pressure, (2) the aquaporin (AQP) gene expression by performing qPCR and (3) the formation of apoplastic barriers, by analyzing suberin lamella and Casparian bands via cross-sectional analyses in roots. Plants were grown hydroponically on complete nutrient solution, containing 250 µM P, until they were 31-36 days old, and then kept for 2-3 weeks on either complete solution, or transferred on solution containing 2.5 µM (low-P) or no added P (no-P). Phosphate treatments caused significant decreases in root and cell-Lp and AQP gene expression, while the formation of apoplastic barriers increased, particularly in lateral roots. Experiments using the AQP inhibitor mercury (Hg) suggested that a significant portion of radial root water uptake in sheepgrass occurs along a path involving AQPs, and that the Lp of this path is reduced under low- and no-P. It is concluded that a growth-limiting supply of phosphate causes parallel changes in (1) cell Lp and aquaporin gene expression (decrease) and (2) apoplastic barrier formation (increase), and that the two may combine to reduce root Lp. The reduction in root Lp, in turn, facilitates an increased root-to-shoot surface area ratio, which allocates resources to the root, sourcing the limiting nutrient.

Cortex cell hydraulic conductivity, endodermal apoplastic barriers and root hydraulics change in barley (Hordeum vulgare L.) in response to a low supply of N and P.

BACKGROUND: Mineral nutrient limitation affects the water flow through plants. We wanted to test on barley whether any change in root-to-shoot ratio in response to low supply of nitrogen and phosphate is accompanied by changes in root and cell hydraulic properties and involves changes in aquaporin (AQP) gene expression and root apoplastic barriers (suberin lamellae, Casparian bands). METHODS: Plants were grown hydroponically on complete nutrient solution or on solution containing only 3.3 % or 2.5 % of the control level of nutrient. Plants were analysed when they were 14-18 d old. RESULTS: Nutrient-limited plants adjusted water flow to an increased root-to-shoot surface area ratio through a reduction in root hydraulic conductivity (Lp) as determined through exudation analyses. Cortex cell Lp (cell pressure probe analyses) decreased in the immature but not the mature region of the main axis of seminal roots and in primary lateral roots. The aquaporin inhibitor HgCl2 reduced root Lp most in nutrient-sufficient control plants. Exchange of low-nutrient for control media caused a rapid (20-80 min) and partial recovery in Lp, though cortex cell Lp did not increase in any of the root regions analysed. The gene expression level (qPCR analyses) of five plasma membrane-localized AQP isoforms did not change in bulk root extracts, while the formation of apoplastic barriers increased considerably along the main axis of root and lateral roots in low-nutrient treatments. CONCLUSIONS: Decrease in root and cortex cell Lp enables the adjustment of root water uptake to increased root-to-shoot area ratio in nutrient-limited plants. Aquaporins are the prime candidate to play a key role in this response. Modelling of water flow suggests that some of the reduction in root Lp is due to increased formation of apoplastic barriers.

Root and cell hydraulic conductivity, apoplastic barriers and aquaporin gene expression in barley (Hordeum vulgare L.) grown with low supply of potassium.

Background and Aims: Limited supply of mineral nutrients often reduces plant growth and transpirational water flow while increasing the ratio of water-absorbing root to water-losing shoot surface. This could potentially lead to an imbalance between water uptake (too much) and water loss (too little). The aim of the present study was to test whether, as a countermeasure, the hydraulic properties (hydraulic conductivity, Lp) of roots decrease at organ and cell level and whether any decreases in Lp are accompanied by decreases in the gene expression level of aquaporins (AQPs) or increases in apoplastic barriers to radial water movement. Methods: Barley plants were grown hydroponically on complete nutrient solution, containing 2 mm K+ (100 %), or on low-K solution (0.05 mm K+; 2.5 %), and analysed when they were 15-18 d old. Transpiration, fresh weight, surface area, shoot water potential (ψ), K and Ca concentrations, root (exudation) and cortex cell Lp (cell pressure probe), root anatomy (cross-sections) and AQP gene expression (qPCR) were analysed. Key Results: The surface area ratio of root to shoot increased significantly in response to low K. This was accompanied by a small decrease in the rate of water loss per unit shoot surface area, but a large (~50 %) and significant decrease in Lp at root and cortex cell levels. Aquaporin gene expression in roots did not change significantly, due to some considerable batch-to-batch variation in expression response, though HvPIP2;5 expression decreased on average by almost 50 %. Apoplastic barriers in the endodermis did not increase in response to low K. Conclusions: Barley plants that are exposed to low K adjust to an increased ratio of root (water uptake) to shoot (water loss) surface primarily through a decrease in root and cell Lp. Reduced gene expression of HvPIP2;5 may contribute to the decrease in Lp.

Night-time transpiration in barley (Hordeum vulgare) facilitates respiratory carbon dioxide release and is regulated during salt stress.

Background and Aims: Night-time transpiration accounts for a considerable amount of water loss in crop plants. Despite this, there remain many questions concerning night-time transpiration - its biological function, regulation and response to stresses such as salinity. The aim of the present study was to address these questions on 14- to 18-d-old, hydroponically grown barley plants. Methods: Plants were either stressed for the last 4-7 d prior to, and during subsequent continuous (24 h), diurnal gravimetric transpiration analyses; or subjected to salt stress just before analyses; or stressed for 4-7 d and then transferred to control medium before analyses. The idea behind this experimental setup was to distinguish between a longer- (cuticle, stomata) and shorter-term (stomata) response of transpiration to treatments. Cuticular conductance was assessed through residual transpiration measurements in detached leaves. Cuticle wax load and dark respiration rate of leaves were determined. Leaf conductance to CO2 was calculated. Key Results: Night-time and daytime transpiration rates were highly, and positively, correlated with each other, across all treatments. Night-time transpiration rates accounted for 9-17 % of daytime rates (average: 13.8 %). Despite minor changes in the ratio of night- to daytime transpiration rates, the contribution of cuticular and stomatal conductance to leaf (epidermal) conductance to water vapour differed considerably between treatments. Salt stress did not affect cuticle wax load. The conductance for CO2 of the cuticle was insufficient to support rates of dark respiratory CO2 release. Conclusions: The main biological function of night-time transpiration is the release of respiratory CO2 from leaves. Night-time transpiration is regulated in the short and long term, also under salt stress. Stomata play a key role in this process. We propose to refer, in analogy to water use efficiency (WUE) during the day, to a CO2 release efficiency ('CORE') during the night.

Zinc treatment of hydroponically grown barley plants causes a reduction in root and cell hydraulic conductivity and isoform-dependent decrease in aquaporin gene expression.

The cellular and molecular basis of a reduction in root water uptake in plants exposed to heavy metals such as zinc (Zn) is poorly studied. The aim of the present study on hydroponically grown barley (Hordeum vulgare) was to test whether any reduction in root hydraulic conductivity (Lp) in response to Zn treatment is accompanied by a reduction in cell Lp and gene expression level of aquaporin (AQP) isoforms. Plants were grown in the presence of 0.25 µM, (control), 0.1 and 1 mM Zn in the root medium and analysed when they were 16-18 days old. Root and cell Lp was determined through exudation and cell pressure probe analyses, respectively, and gene expression of five candidate AQPs was analysed [real time quantitative polymerase chain reaction (PCR)]. Zinc treatments caused significant reductions (25-83%) in transpiration rate, root and shoot fresh weight, surface area and stomatal conductance. Zinc concentrations in tissues increased more than 100-fold. Root Lp decreased by 24% (0.1 mM Zn) and 58% (1 mM Zn), while cell Lp decreased by 45 and 71%, respectively. Gene expression of AQPs decreased by 14-80%; decreases were statistically significant for HvPIP1;3, HvPIP2;4 and HvPIP2;5. Turgor in root cortex cells was not reduced by Zn treatments. It is concluded that reductions in plant water flow in response to Zn treatment are facilitated through decreases in root (Lp) and shoot (stomata) hydraulics. The decrease in root Lp is facilitated through reductions in cell Lp and AQP gene expression and may also reflect increased suberization in the endodermis.

Water transport and energy.

Water transport in plants occurs along various paths and is driven by gradients in its free energy. It is generally considered that the mode of transport, being either diffusion or bulk flow, is a passive process, although energy may be required to sustain the forces driving water flow. This review aims at putting water flow at the various organisational levels (cell, organ, plant) in the context of the energy that is required to maintain these flows. In addition, the question is addressed (1) whether water can be transported against a difference in its chemical free energy, 'water potential' (Ψ), through, directly or indirectly, active processes; and (2) whether the energy released when water is flowing down a gradient in its energy, for example during day-time transpiration and cell expansive growth, is significant compared to the energy budget of plant and cell. The overall aim of review is not so much to provide a definite 'Yes' and 'No' to these questions, but rather to stimulate discussion and raise awareness that water transport in plants has its real, associated, energy costs and potential energy gains.

Rapid changes in root hydraulic conductivity and aquaporin expression in rice (Oryza sativa L.) in response to shoot removal - xylem tension as a possible signal.

Background and Aims It is not clear how plants adjust the rate of root water uptake to that of shoot water loss. The aim of this study on rice was to test the idea that root aquaporins (AQPs) and xylem tension play a role in this adjustment. Methods Three-week-old rice (Oryza sativa L.) plants, which were grown hydroponically, had their entire shoot system removed, and root hydraulic conductivity (exudation analyses) and gene expression (quantitative real-time PCR) of root plasma membrane intrinsic aquaporin proteins (PIPs) was followed within 60 min after shoot excision. Key Results All three PIP1 genes (OsPIP1;1, OsPIP1;2 and OsPIP1;3) and three of the six PIP2 genes tested (OsPIP2;1, OsPIP2;4 and OsPIP2;5) showed a rapid (5 min) and lasting (60 min) decrease in gene expression. Expression decreased by up to 85 % within 60 min. The other three PIP2 genes tested (OsPIP2;2, OsPIP2;3 and OsPIP2;6) showed a varied response, with expression decreasing either only initially (5 min) or after 60 min, or not changing at all. In a follow-up experiment, plants had their shoot system removed and the detached root system immediately connected to a vacuum pump through which some tension (80 kPa) was applied. This application of tension prevented any significant decrease in PIP expression. Conclusions Shoot removal leads to a rapid decrease in expression of all PIP1s and some PIP2s in roots of rice. Xylem tension plays some role in this process.

Exogenous application of abscisic acid (ABA) increases root and cell hydraulic conductivity and abundance of some aquaporin isoforms in the ABA-deficient barley mutant Az34.

Background and Aims Regulation of water channel aquaporins (AQPs) provides another mechanism by which abscisic acid (ABA) may influence water flow through plants. To the best of our knowledge, no studies have addressed the changes in ABA levels, the abundance of AQPs and root cell hydraulic conductivity (LpCell) in the same tissues. Thus, we followed the mechanisms by which ABA affects root hydraulics in an ABA-deficient barley mutant Az34 and its parental line 'Steptoe'. We compared the abundance of AQPs and ABA in cells to determine spatial correlations between AQP abundance and local ABA concentrations in different root tissues. In addition, abundance of AQPs and ABA in cortex cells was related to LpCell. Methods Root hydraulic conductivity (LpRoot) was measured by means of root exudation analyses and LpCell using a cell pressure probe. The abundance of ABA and AQPs in root tissues was assessed through immunohistochemical analyses. Isoform-specific antibodies raised against HvPIP2;1, HvPIP2;2 and HvPIP2;5 were used. Key Results Immunolocalization revealed lower ABA levels in root tissues of Az34 compared with 'Steptoe'. Root hydraulic conductivity (LpRoot) was lower in Az34, yet the abundance of HvPIPs in root tissues was similar in the two genotypes. Root hair formation occurred closer to the tip, while the length of the root hair zone was shorter in Az34 than in 'Steptoe'. Application of external ABA to the root medium of Az34 and 'Steptoe' increased the immunostaining of root cells for ABA and for HvPIP2;1 and HvPIP2;2 especially in root epidermal cells and the cortical cell layer located beneath, parallel to an increase in LpRoot and LpCell. Treatment of roots with Fenton reagent, which inhibits AQP activity, prevented the ABA-induced increase in root hydraulic conductivity. Conclusion Shortly after (<2 h) ABA application to the roots of ABA-deficient barley, increased tissue ABA concentrations and AQP abundance (especially the plasma-membrane localized isoforms HvPIP2;1 and HvPIP2;2) were spatially correlated in root epidermal cells and the cortical cell layer located beneath, in conjunction with increased LpCell of the cortical cells. In contrast, long-term ABA deficiency throughout seedling development affects root hydraulics through other mechanisms, in particular the developmental timing of the formation of root hairs closer to the root tip and the length of the root hair zone.

Limitation of Cell Elongation in Barley (Hordeum vulgare L.) Leaves Through Mechanical and Tissue-Hydraulic Properties.

The aim of the present study was to assess the mechanical and hydraulic limitation of growth in leaf epidermal cells of barley (Hordeum vulgare L.) in response to agents which affect cellular water (mercuric chloride, HgCl(2)) and potassium (cesium chloride, CsCl; tetraethylammonium, TEA) transport, pump activity of plasma membrane H(+)-ATPase and wall acidification (fusicoccin, FC). Cell turgor (P) was measured with the cell pressure probe, and cell osmotic pressure (π) was analyzed through picoliter osmometry of single-cell extracts. A wall extensibility coefficient (M) and tissue hydraulic conductance coefficient (L) were derived using the Lockhart equation. There was a significant positive linear relationship between relative elemental growth rate and P, which fit all treatments, with an overall apparent yield threshold of 0.368 MPa. Differences in growth between treatments could be explained through differences in P. A comparison of L and M showed that growth in all except the FC treatment was co-limited through hydraulic and mechanical properties, though to various extents. This was accompanied by significant (0.17-0.24 MPa) differences in water potential (ΔΨ) between xylem and epidermal cells in the leaf elongation zone. In contrast, FC-treated leaves showed ΔΨ close to zero and a 10-fold increase in L.

The significance of water co-transport for sustaining transpirational water flow in plants: a quantitative approach.

In a recent Opinion paper, Wegner (Journal of Experimental Botany 65, 381-392, 2014) adapts a concept developed for water flow in animal tissues to propose a model, which can explain the loading of water into the root xylem against a difference in water potential (Ψ) between the xylem parenchyma cell (more negative Ψ) and the xylem vessel (less negative Ψ). In this model, the transport of water is energized through the co-transport of ions such as K(+) and Cl(-) through plasma membrane-located transporters. The emphasis of the model is on the thermodynamic feasibility of the co-transport mechanism per se. However, what is lacking is a quantitative evaluation of the energy input required at the organismal level to sustain such a co-transport mechanism in the face of considerable net (transpirational) flows of water through the system. Here, we use a ratio of 500 water molecules being co-transported for every pair of K(+) and Cl(-) ions, as proposed for the animal system, to calculate the energy required to sustain daytime and night-time transpirational water flow in barley plants through a water co-transport mechanism. We compare this energy with the total daily net input of energy through photosynthetic carbon assimilation. Water co-transport can facilitate the filling of xylem against a difference in Ψ of 1.0MPa and puts a minor drain on the energy budget of the plant. Based on these findings it cannot be excluded that water co-transport in plants contributes significantly to xylem filling during night-time and possibly also daytime transpiration.