Hydraulic control of stomatal conductance in Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] and alder [Alnus rubra (Bong)] seedlings

Experiments were conducted on 1-year-old Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] and 2- to 3-month-old alder [Alnus rubra (Bong)] seedlings growing in drying soils to determine the relative influence of root and leaf water status on stomatal conductance (g_c). The water status of shoots was manipulated independently of that of the roots using a pressure chamber that enclosed the root system. Pressurizing the chamber increases the turgor of cells in the shoot but not in the roots. Seedling shoots were enclosed in a whole-plant cuvette and transpiration and net photosynthesis rates measured continuously. In both species, stomatal closure in response to soil drying was progressively reversed with increasing pressurization. Responses occurred within minutes of pressurization and measurements almost immediately returned to pre-pressurization levels when the pressure was released. Even in wet soils there was a significant increase in g_c with pressurization. In Douglas fir, the stomatal response to pressurization was the same for seedlings grown in dry soils for up to 120 d as for those subjected to drought stress over 40 to 60 d. The stomatal conductance of both Douglas fir and alder seedlings was less sensitive to root chamber pressure at higher vapour pressure deficits (D), and stomatal closure in response to increasing D from 1.04 to 2.06 kPa was only partially reversed by pressurization. Our results are in contrast to those of other studies on herbaceous species, even though we followed the same experimental approach. They suggest that it is not always appropriate to invoke a ‘feedforward’ model of short-term stomatal response to soil drying, whereby chemical messengers from the roots bring about stomatal closure.     

Carbon isotope fractionation in tree ring early and late wood in relation to intra-growing season water balance

We determined the stable carbon isotope composition (δ^1.3C) of cellulose extracted from early and late wood in Douglas fir [Pseudotsuga menziexii (Mirb.) Franco] tree rings. Data were obtained for the period 1962 to 1981, at the start of which the trees were 20 years old. A water balance model was used to calculate daily stand transpiration and water deficit. The model incorporates site factors (soil water availability, slope and aspect) and environmental variables (solar radiation, air temperature and rainfall). There was far greater variability in late wood than in early wood δ^1.3C. In wet years, late wood δ^1.3C was significantly lighter (by as much as 2δ) than early wood δ^1.3C but in dry years this difference was reversed. Differences between spring and summer cumulative transpiration accounted for almost 60δ of the variability in differences between early and late wood δ^1.3C. We found excellent correspondence between summer cumulative transpiration and late wood δ^1.3C, with estimates of transpiration accounting for up to 93% of the variability in δ^1.3C. Correlations between early wood δ^1.3C and spring transpiration were generally poor (r2<0.4), but we were able to identify those exceptional years in which there had been a very dry spring. Our results indicate that, while tree ring δ^1.3C correlates reasonably well with basal area increment, it is a far better indicator of inter- and intra-annual variability in water availability than radial growth.     

Sedimentation and pedogenesis in a Central Amazonian Black water basin

Sedimentation rates were estimated in a Central Amazonian Black-water inundation forest. Sediment deposition on the forest ground, remote from the river bed, during an annual flood period, is of the order of 1 to 10 tons per hectare, depending on water depth and duration of flooding. The sediments consisted of fine organic matter, kaolinite, quartz sands and biogenic particles of silica. Their genesis and deposition depend on the interplay between pedogenic, limnological and biological processes. Sediments derive primarily from the materials leached from the soils. Clay soils are the main source of dissolved silica, and the sandy soils are the main sources of organic coumpounds and mineral particles. The physical sedimentation of particles as quartz sand grains only occurs in the upper reaches of the studied river. In the flood plain, the sedimentation is due to the coagulation and deposition of combined mineral particles and humic substances, and to the biological precipitation of the silica leached from the soil by sponges.     

Cold hardiness and water relations parameters in Rhododendron cv. Catawbiense Boursault subjected to drought episodes

We determined whether increase in cold hardiness of Rhododendron cv. Catawbiense Boursault induced by water stress was correlated with changes in tissue water relations. Water content of the growing medium was either maintained near field capacity for the duration of the study or plants were subjected to drought episodes at different times between 15 July and 19 February. Watering during a drought episode was delayed until soil water content decreased below 0.4 m³ m⁻³ then watering was resumed at a level to maintain soil water content between 0.3 and 0.4 m³ m⁻³. Cold hardiness was evaluated in the laboratory with freeze tolerance tests on detached leaves. Water relations parameters were determined using pressure-volume analysis. Exposure to drought episodes increased cold hardiness during the cold acclimation stage in late summer and fall but not during the winter. When water-stressed plants were re-watered to field capacity, the previous gain in cold hardiness gradually disappeared. Water relations parameters correlating with seasonal changes of cold hardiness included dry matter content (r =−0.67). apoplastic water content (r =−0.60), and water potential at the turgor loss point (r = 0.40). Changes of cold hardiness in water-stressed plants in reference to well-watered plants were correlated with changes of all water relations parameters, except for osmotic potential at full turgor (r = 0.13). It is proposed that water stress reduced the hydration of cell walls, thereby increasing their rigidity. Increased rigidity of cell walls could result in a development of greater negative turgor pressures at subfreezing temperatures and therefore increased resistance to freeze dehydration.     

CAM variations in the leaf-succulent Delosperma tradescantioides (Mesembryanthemaceae), native to southern Africa

Drought responses of diurnal gas exchange, malic acid accumulation and water status were examined in Delosperma tradescantioides , a succulent that grows in drought-prone microenvironments in summer rainfall and all-year rainfall regions of southern Africa. When well-watered, this species exhibited Crassulacean acid metabolism (CAM)-cycling, but its carbon fixation pattern changed during the development of drought, shifting to either low-level CAM or to CAM-idling. The rate and pattern of this change depended on environmental conditions, duration of water stress and leaf age. At the onset of drought, diurnal malate fluctuation increased, but was strongly depressed (by ca 70%) as drought continued, and when leaf water content and water potential were low (ca 35 and 50% of the initial levels, respectively). When rewatered, rates of growth and photosynthesis, gas exchange and water status recovered fully to pre-stressed values within two days. Whole-shoot carbon uptake rates suggested that leaf growth had continued unabated during a short-term (∼ one week) drought. This emphasises that CAM-idling allows the maintenance of active metabolism with negligible gas exchange when soil water is limiting. It is possible that old or senescent leaves may provide water for the expansion of developing leaves during initial periods of drought. Regardless of the water regime and environmental conditions, leaf nocturnal malate accumulation and water content were positively correlated and increased with leaf age. Thus the gradual loss of water from older mature leaves may induce CAM-idling, which reduces water loss. An important ecological consequence of this combination of CAM modes is the potential to switch rapidly between fast growth via C₃ gas exchanges when well-watered to water-conserving CAM-idling during drought.     

Physiological aspects of Taxus brevifolia seeds in relation to seed storage characteristics

Water relations, desiccation tolerance and longevity of Taxus brevifolia (Nutt.) seeds were studied to determine the optimal stage of development and storage conditions for seeds of this species. Seeds equilibrated to a range of relative humidities (RHs) had unusually low water contents which can be accounted for by the high lipid content of gametophyte tissues (71% of the dry mass). Water relations of embryonic tissue were more typical of those reported for other seed species. The water content below which freezing transitions were not observable in the embryo was ca 0.24 g H₂O (g dry weight)⁻¹ (g g⁻¹) for all maturity classes studied. Embryos did not achieve significant levels of desiccation tolerance (survival to water contents less than 0.5 g g⁻¹) until the latter stages of development when dry matter was maximal. Mature embryos could be dried to 0.025 g g⁻¹ (seed water content of 0.010 g g⁻¹) with no loss of viability. Thus, at the latter stages of development, embryo water content could be optimized to avoid both desiccation and freezing damage. Survival of mature seeds declined over a 2-year period when seeds were stored at temperatures between 5 and 35°C and RHs between 14 and 75%, corresponding to seed water contents between 0.015 and 0.07 g g⁻¹. The deterioration rate was slowest for seeds stored at the lowest RH and temperature. Our data indicate that seeds of Taxus brevifolia show orthodox rather than recalcitrant storage characteristics, but that the optimum water content for storage was extremely low. The results suggest that even if stored at optimal water contents and low temperatures, T. brevifolia seeds will be relatively short lived. The high quantity of lipids or reducing sugars may be contributing factors in the poor storage characteristics.     

Relative growth rate, biomass allocation pattern and water use efficiency of three wheat cultivars during early ontogeny as dependent on water availability

We have investigated the water use efficiency of whole plants and selected leaves and allocation patterns of three wheat cultivars (Mexipak, Nesser and Katya) to explore how variation in these traits can contribute to the ability to grow in dry environments. The cultivars exhibited considerable differences in biomass allocation and water use efficiency. Cultivars with higher growth rates of roots and higher proportions of biomass in roots (Nesser and Katya) also had higher leaf growth rates, higher proportions of their biomass as leaves and higher leaf area ratios. These same cultivars had lower rates of transpiration per unit leaf area or unit root weight and higher biomass production per unit water use. They also had higher ratios of photosynthesis to transpiration, and lower ratios of intercellular to external CO₂ partial pressure. The latter resulted from large differences in stomatal conductance associated with relatively small differences in rates of photosynthesis. There was little variation between cultivars in response to drought, and differences in allocation pattern and plant water use efficiency between cultivars as found under well-watered conditions persisted under dry conditions. At the end of the non-watered treatment, relative growth rates and transpiration rates decreased to similar values for all cultivars. High ratios of photosynthesis to transpiration, and accordingly high biomass production per unit of transpiration, is regarded as a favourable trait for dry environments, since more efficient use of water postpones the decrease in plant water status.     

Inositol 1,4,5-trisphosphate formation in leaves of winter oilseed rape plants in response to freezing, tissue water potential aed abscisic acid

Time courses of formation of inositol 1,4,5-trisphosphate (IP₃) were followed in the leaves of non-acclimated and cold (2°C)-acclimated winter oilseed rape ( Brassica napus L. var. oleifera ) plants, subjected to different freezing temperatures or to polyethylene glycol 8000 (PEG) and abscisic acid (ABA) treatments. Changes in water potential (Ψ_w) and in ABA level in the frost- and PEG-treated tissues were also determined. Results obtained indicate that temperatures sligthly higher than LT₅₀ induced a transient and substantial increase in IP₃ level, both in non-acclimated and cold-acclimated tissues. At comparable freezing temperature (–5°C) the response of cold-acclimated leaves was lower than that of non-acclimated ones. The PEG-depedent decrease in Ψ_w to –0.9 MPa or ABA (0.1 m M ) treatment gave rise to a transient increase in IP₃ content in non-acclimated tissues only. Collectively, the data indicate that cold acclimation of plants may lead to lower cell responsiveness to the factors studied in terms of induction of IP₃ formation. Changes in the IP₃ content, observed in the present experiments, support our previous suggestion that non-killing freezing temperatures may induce the phosphoinositide pathway, both in non-acclimated and cold-acclimated tissues. Lowering of tissue water potential to some threshold value or a high exogenous ABA supply may mimic the freezing-dependent reaction in the non-acclimated leaves.