Reduction In Root Hydraulic Conductivity Research Articles

Transpiration difference under high evaporative demand in chickpea (Cicer arietinum L.) may be explained by differences in the water transport pathway in the root cylinder.

Terminal drought substantially reduces chickpea yield. Reducing water use at vegetative stage by reducing transpiration under high vapor pressure deficit (VPD), i.e. under dry/hot conditions, contributes to drought adaptation. We hypothesized that this trait could relate to differences in a genotype's dependence on root water transport pathways and hydraulics. Transpiration rate responses in conservative and profligate chickpea genotypes were evaluated under increasing VPD in the presence/absence of apoplastic and cell-to-cell transport inhibitors. Conservative genotypes ICC 4958 and ICC 8058 restricted transpiration under high VPD compared to the profligate genotypes ICC 14799 and ICC 867. Profligate genotypes were more affected by aquaporin inhibition of the cell-to-cell pathway than conservative genotypes, as measured by the root hydraulic conductance and transpiration under high VPD. Aquaporin inhibitor treatment also led to a larger reduction in root hydraulic conductivity in profligate than in conservative genotypes. In contrast, blockage of the apoplastic pathway in roots decreased transpiration more in conservative than in profligate genotypes. Interestingly, conservative genotypes had high early vigour, whereas profligate genotypes had low early vigour. In conclusion, profligate genotypes depend more on the cell-to-cell pathway, which might explain their higher root hydraulic conductivity, whereas water-saving by restricting transpiration led to higher dependence on the apoplastic pathway. This opens the possibility to screen for conservative or profligate chickpea phenotypes using inhibitors, itself opening to the search of the genetic basis of these differences.

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.

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). 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. 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. 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.

Surveillance of cell wall diffusion barrier integrity modulates water and solute transport in plants

The endodermis is a key cell layer in plant roots that contributes to the controlled uptake of water and mineral nutrients into plants. In order to provide such functionality the endodermal cell wall has specific chemical modifications consisting of lignin bands (Casparian strips) that encircle each cell, and deposition of a waxy-like substance (suberin) between the wall and the plasma membrane. These two extracellular deposits provide control of diffusion enabling the endodermis to direct the movement of water and solutes into and out of the vascular system in roots. Loss of integrity of the Casparian strip-based apoplastic barrier is sensed by the leakage of a small peptide from the stele into the cortex. Here, we report that such sensing of barrier integrity leads to the rebalancing of water and mineral nutrient uptake, compensating for breakage of Casparian strips. This rebalancing involves both a reduction in root hydraulic conductivity driven by deactivation of aquaporins, and downstream limitation of ion leakage through deposition of suberin. These responses in the root are also coupled to a reduction in water demand in the shoot mediated by ABA-dependent stomatal closure.

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 1mM 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% (0.1mM Zn) and 58% (1mM 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.

Phosphorus deprivation effects on water relations of Nicotiana tabacum plant via reducing plasma membrane permeability

Plants grown in phosphorus-deprived solutions often exhibit disruption of water transport due to reduction in root hydraulic conductivity (Lpr). To uncover the relationship between root Lpr and water permeability coefficient (Pf) of plasma membrane and the role of aquaporins, we evaluated Pf of plasma membrane and also PIP-type aquaporin gene expression in tobacco (Nicotiana tabacum L.) plant roots after seven days P-deprivation. The results showed significant reduction in sap flow rate (Jv) and osmotic root hydraulic conductivity (Lpr-o) in P-deprived roots. These effects were reversed 24 h after P-resupplying. Interestingly, the Pf of root protoplasts was 57% lower in P-deprived plants compared with P-sufficient ones. The expression of NtPIP1;1 and NtPIP2;1 aquaporins did not change significantly in P-deprived plants compared with P-sufficient ones, but the copy number of NtAQP1 increased significantly in P-deprived plants. P-deprivation did not change Lpr-o significantly in antisense NtAQP1 plants. Taken together, these findings suggest that P-deprivation may play an important role in modulation of root hydraulic conductivity by affecting Pf in transcellular pathway of water flow across roots and aquaporins. Finally, we concluded that dominant water transport pathway under P-deprivation was transcellular one.

Mild Salt Stress Conditions Induce Different Responses in Root Hydraulic Conductivity of Phaseolus vulgaris Over-Time

Plants respond to salinity by altering their physiological parameters in order to maintain their water balance. The reduction in root hydraulic conductivity is one of the first responses of plants to the presence of salt in order to minimize water stress. Although its regulation has been commonly attributed to aquaporins activity, osmotic adjustment and the toxic effect of Na+ and Cl− have also a main role in the whole process. We studied the effects of 30 mM NaCl on Phaseolus vulgaris plants after 9 days and found different responses in root hydraulic conductivity over-time. An initial and final reduction of root hydraulic conductivity, stomatal conductance, and leaf water potential in response to NaCl was attributed to an initial osmotic shock after 1 day of treatment, and to the initial symptoms of salt accumulation within the plant tissues after 9 days of treatment. After 6 days of NaCl treatment, the increase in root hydraulic conductivity to the levels of control plants was accompanied by an increase in root fructose content, and with the intracellular localization of root plasma membrane aquaporins (PIP) to cortex cells close to the epidermis and to cells surrounding xylem vessels. Thus, the different responses of bean plants to mild salt stress over time may be connected with root fructose accumulation, and intracellular localization of PIP aquaporins.

Phosphorus deficiency-induced reduction in root hydraulic conductivity in Medicago falcata is associated with ethylene production

Plants grown in phosphorus-deficient solutions often exhibit disruption of water transport due to reduction in root hydraulic conductivity (Lpr) and enhanced ethylene production. To uncover the relationship between the reduction in Lpr and increase in ethylene production, we investigated effect of phosphorus (P) deficiency on ethylene production and Lpr in legume plants of Medicago falcata L. There was an increase in ethylene production and a reduction of Lpr of M. falcata roots when M. falcata seedlings grown in P sufficient solutions (0.5 mM H 2PO 4 2−) were transferred to P-deficient solutions (5 μM H 2PO 4 2−). Antagonists of ethylene biosynthesis, CoCl 2 and aminoethoxyvinylglycine (AVG), abolished the P deficiency-induced ethylene production. Root hydraulic conductivity of M. falcata seedlings grown in P-sufficient solutions was insensitive to CoCl 2 and AVG, while the two chemicals enhanced Lpr for those grown in P-deficient solutions, suggesting that P deficiency-induced decrease in Lpr can be reversed by inhibiting ethylene production. Ethylene precursor 1-amino cyclopropane-1-carboxylic acid (ACC) and ethylene donor ethephon had greater inhibitory effect on Lpr of P-sufficient seedlings than that of P-deficient seedlings. Root hydraulic conductivity of P-sufficient seedlings was more sensitive to HgCl 2 than that of P-deficient seedlings. Taken together, these findings suggest that ethylene induced by P deficiency may play an important role in modulation of root hydraulic conductivity by affecting aquaporins in plants.

Intracellular ROS

Intracellular localization of stress induced reactive oxygen species (ROS) has emerged as an important aspect towards understanding of cellular responses to environmental stimuli. Our recent study published in the PNAS (103:18008–13)1 shows that NaCl-induced ROS appear within endosomes on the way to tonoplast as part of the vacuolar vesicle trafficking. In addition to showing ROS damage to the tonoplast, this finding may shed light upon recently reported aspects of root water relations during salt stress, suggesting a new signaling role for intracellular ROS in Arabidopsis root cells, during salt stress: ROS that are compartmentalized in endosomes are delivered by the vacuolar vesicle trafficking pathway to the tonoplast, resulting in oxidative gating of TIPs water channels. The closure of the tonoplast aquaporins contributes to the observed reduction in root hydraulic conductivity during salt stress.

Cycloheximide inhibits root water flow and stomatal conductance in aspen (Populus tremuloides) seedlings

ABSTRACTAspen (Populus tremuloides Michx.) roots were treated with cycloheximide, a protein synthesis inhibitor, to examine the role of protein synthesis in root water transport and plant water relations. Within less than 30 min following root application, cycloheximide inhibited steady‐state root water flow rates and 1 h after the application of 1 mm cycloheximide, root hydraulic conductivity had decreased by 85% compared with control roots. However, stomatal conductance showed a significant inhibition only after 2 h following cycloheximide treatment. The reduction in root hydraulic conductivity was accompanied by an almost three‐fold increase in the apoplastic water flow ratio as determined by the trisodium 3‐hydroxy‐5,8,10‐pyrenesulphonate tracer dye. Cycloheximide‐treated roots showed a decrease in the immunostaining intensity of a 32 kDa microsomal protein band that immunoreacted with the AnthPIP1; 1 antibody suggesting a decrease in the membrane aquaporin expression. These changes occurred without severe metabolic disruptions as measured by root respiration. The results point to the importance of protein‐mediated transport in roots and the rapidity of response suggests that protein synthesis may be used as a principal regulatory mechanism in root water transport in aspen.