Aquaporins, found in virtually all living organisms, are membrane-intrinsic proteins that form water-permeable complexes. The mammalian aquaporin AQP1 has also shown CO2 permeability when expressed heterologously in Xenopus oocytes, although whether this is a biochemical curiosity or of physiological significance is a matter of debate. Here we report that, in the same expression system, a CO2 permeability comparable to that of the human AQP1 is observed for the tobacco plasma membrane aquaporin NtAQP1. NtAQP1 facilitates CO2 membrane transport in the homologous plant system at the cellular level, and has a significant function in photosynthesis and in stomatal opening. NtAQP1 overexpression heightens membrane permeability for CO2 and water, and increases leaf growth. The results indicate that NtAQP1-related CO2 permeability is of physiological importance under conditions where the CO2 gradient across a membrane is small, as is the case between the atmosphere and the inside of a plant cell.
The molecular functions of several aquaporins are well characterized (e.g., by analysis of aquaporin-expressing Xenopus oocytes). However, their significance in the physiology of water transport in multicellular organisms remains uncertain. The tobacco plasma membrane aquaporin NtAQP1 was used to elucidate this issue. By comparing antisense plants that were inhibited in NtAQP1 expression with control plants, we found evidence for NtAQP1 function in cellular and whole-plant water relations. The consequences of a decrease in cellular water permeability were determined by measurement of transpiration rate and stem and leaf water potential as well as growth experiments under extreme soil water depletion. Plants impaired in NtAQP1 expression showed reduced root hydraulic conductivity and lower water stress resistance. In conclusion, our results emphasize the importance of symplastic aquaporin-mediated water transport in whole-plant water relations.
Young maize seedlings (Zea mays L. cv.Giza 2) were exposed to moderate salinity in hydroponic culture. NADH‐nitrate reductase (NR) activity (E.C. 1·6.6·1), NR activation state, NR‐mRNA‐steady state levels and major solute contents in leaves and roots were investigated. With increasing external salt concentration, Na+, Cl−, sugars, amino acids and quarternary ammonium compounds accumulated in leaves and roots, with concentrations in leaves exceeding those in roots. The nitrate content of leaves decreased, but increased in roots. The diurnal pattern of NR activity and of NR‐mRNA was also changed under salinity, but the NR activation state was not affected. In the first light phase, maximum NR activity increased rapidly in leaves of control plants, but was much slower in leaves from salinized plants. Thus, integrated over the whole day, the NR activity of salt‐stressed plants was lower than in control plants. NR transcript levels of control plants were low in the early night phase, started to increase in the second night phase, followed by a distinct peak at 2 h into the light period. This large ‘early morning peak’ of NR‐mRNA was hardly affected by salinity, whereas the initial slow increase of m‐RNA levels in the early night phase was almost absent in salinized plants. This is considered as one reason for the low NR activity of salinized plants in the first half of the day. It is also suggested that nitrate is a major signal affecting NR expression and activity under salinity. Sugars and amino acids appeared less important.
SummaryEpinastic leaf movement of tobacco is based on differential growth of the upper and lower leaf surface and is distinct from the motor organ-driven mechanism of nyctinastic leaf movement of, for example, mimosa species. The epinastic leaf movement of tobacco is observed not only under diurnal light regimes but also in continuous light, indicating a control by light and the circadian clock. As the transport of water across membranes by aquaporins is an important component of rapid plant cell elongation, the role of the tobacco aquaporin Nt aquaporin (AQP)1 in the epinastic response was studied in detail. In planta NtAQP1-luciferase (LUC) activity studies, Northern and Western blot analyses demonstrated a diurnal and circadian oscillation in the expression of this plasma membrane intrinsic protein (PIP)1-type aquaporin in leaf petioles, exhibiting peaks of expression coinciding with leaf unfolding. Cellular water permeability of protoplasts isolated from leaf petioles was found to be high in the morning, i.e. during the unfolding reaction, and low in the evening. Moreover, diurnal epinastic leaf movement was shown to be reduced in transgenic tobacco lines with an impaired expression of NtAQP1. It is concluded that the cyclic expression of PIP1-aquaporin represents an important component of the leaf movement mechanism.
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