The role of root temperature T(R) in regulating the water-uptake capability of rice roots and the possible relationship with aquaporins were investigated. The root hydraulic conductivity Lp(r) decreased with decreasing T(R) in a measured temperature range between 10 degrees C and 35 degrees C. A single break point (T(RC) = 15 degrees C) was detected in the Arrhenius plot for steady-state Lp(r). The temperature dependency of Lp(r) represented by activation energy was low (28 kJ mol(-1)) above T(RC), but the value is slightly higher than that for the water viscosity. Addition of an aquaporin inhibitor, HgCl(2), into root medium reduced osmotic exudation by 97% at 25 degrees C, signifying that aquaporins play a major role in regulating water uptake. Below T(RC), Lp(r) declined precipitously with decreasing T(R) (E(a) = 204 kJ mol(-1)). When T(R) is higher than T(RC), the transient time for reaching the steady-state of Lp(r) after the immediate change in T(R) (from 25 degrees C) was estimated as 10 min, while it was prolonged up to 2-3 h when T(R) < T(RC). The Lp(r) was completely recovered to the initial levels when T(R) was returned back to 25 degrees C. Immunoblot analysis using specific antibodies for the major aquaporin members of PIPs and TIPs in rice roots revealed that there were no significant changes in the abundance of aquaporins during 5 h of low temperature treatment. Considering this result and the significant inhibition of water-uptake by the aquaporin inhibitor, we hypothesize that the decrease in Lp(r) when T(R) < T(RC) was regulated by the activity of aquaporins rather than their abundance.
Root temperature is found to be a very important factor for leaves to alter the response and susceptibility to chilling stress. Severe visible damage was observed in the most active leaves of seedlings of a japonica rice (Oryza sativa cv. Akitakomachi), e.g. the third leaf at the third-leaf stage, after the treatment where only leaves but not roots were chilled (L/H). On the other hand, no visible damage was observed after the treatment where both leaves and roots were chilled simultaneously (L/L). The chilling injury induced by L/H, a novel type of chilling injury, required the light either during or after the chilling in order to develop the visible symptoms such as leaf bleaching and tissue necrosis. Chlorophyll fluorescence parameters measured after various lengths of chilling treatments showed that significant changes were induced before the visible injury. The effective quantum yield and photochemical quenching of PSII dropped dramatically within 24 h in both the presence and absence of a 12 h light period. The maximal quantum yield and non-photochemical quenching of PSII decreased significantly only in the presence of light. On the other hand, L/H chilling did not affect the function of PSI, but caused a significant decrease in the electron availability for PSI. These results suggest that the leaf chilling with high root temperature destroys some component between PSII and PSI without the aid of light, which causes the over-reduction of PSII in the light, and thereby the visible injury is induced only in the light.
The effects of low air humidity and low root temperature (LRT) on water uptake, growth and aquaporin gene expression were investigated in rice plants. The daily transpiration of the plants grown at low humidity was 1.5- to 2-fold higher than that at high humidity. LRT at 13°C reduced transpiration, and the extent was larger at lower humidity. LRT also reduced total dry matter production and leaf area expansion, and the extent was again larger at lower humidity. These observations suggest that the suppression of plant growth by LRT is associated with water stress due to decreased water uptake ability of the root. On the other hand, the net assimilation rate was not affected by low humidity and LRT, and water use efficiency was larger for LRT. We found that low humidity induced coordinated up-regulation of many PIP and TIP aquaporin genes in both the leaves and the roots. Expression levels of two root-specific aquaporin genes, OsPIP2;4 and OsPIP2;5, were increased significantly after 6 and 13 d of LRT exposure. Taken together, we discuss the possibility that aquaporins are part of an integrated response of this crop to low air humidity and LRT.
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