Stable carbon isotopes as indicators of increased water use efficiency and productivity in white spruce (Picea glauca (Moench) Voss) seedlings

The relationship among water use efficiency (WUE), productivity and carbon isotopic composition (δ¹³C) in white spruce (Picea glauca (Moench) Voss) seedlings was investigated. Sixteen hundred seedlings representing 10 controlled crosses were planted in the field in individual buried sand-filled cylinders. The soil water content in the cylinders was measured using time domain reflectometry over two growing seasons and seedling water use determined by water balance. Two watering treatments were imposed: irrigation and dry land. There was significant (1.6–2.0%c) genetic variation in needle δ¹³C. Ranking of crosses in terms of δ¹³C was generally maintained over watering treatments and there was not a significant genetic versus environmental interaction. There was a positive correlation between δ¹³C and both intrinsic and long-term WUE (more positive δ¹³C with increased WUE) and between δ¹³C and productivity, suggesting a correlation due to variation in photosynthetic capacity. Root to shoot ratios did not increase in water-stressed plants, indicating that responses to drought were primarily at the level of gas exchange, rather than through morphological changes. Our results indicate that it should be possible to use δ¹³C as a surrogate for WUE and to select white spruce genotypes for high WUE without compromising yield.     

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.     

Rates of stomatal opening in conifer seedlings in relation to air temperature and daily carbon gain

Experiments were conducted on well watered 1-year-old Douglas fir [Pseudotsuga menziesii (Mirb.) Franco], western hemlock [Tsuga heterophylla (Raf.) Sarg.] and western redcedar (Thuja plicata Donn) seedlings to determine the effects of temperature on whole-plant photosynthetic and stomatal responses to short-term fluctuations in irradiance (Q). Following a step change in Q, time constants (τ, the period over which 63% of the total change occurs) for stomatal conductance (g_s) and assimilation rate (A) decreased linearly with increasing air temperature (T_air). For example, in western redcedar τ_A decreased from 30 ± 4 min at 5 °C to 10 ± 1 min at 25 °C. In all cases, τ_A was within 10–15% of τ_gs. There was considerable variation in τ among individuals within a given species. Differences between species became more pronounced with decreasing temperature. Multiplicative models that included functions for τ accounted for 99% of the diurnal variability in A and g_s for seedlings exposed to varying air temperature, irradiance and vapour pressure deficit. Estimates of daily A were within 2% of those measured. Intermittent cloud cover and understory shading were approximated by exposing seedlings to 3–4 episodes (≥1 h) of shade (200 or 500 μmol m^?2 s^?1) or complete darkness during the day. In such cases, daily A was overestimated by up to 4 and 21%, respectively, if a function for τ was excluded from the models. Our results suggest that there is scope for selecting seedling stock for increased carbon assimilation on the basis of reduced time constants. For example, in western redcedar, a 40% reduction in τ could lead to increases in daily carbon gains of almost 5% depending on the frequency and degree of shading. If these daily gains were translated into increased dry matter production and compounded, seasonal gains would be even larger.     

Carbon isotope discrimination and water-use efficiency of six crops grown under wet and dryland conditions

Mustard (Sinapis alba L.), Argentine canola (Brassica napus L. cv. Westar), Polish canola (Brassica campestris L. cv. Tobin), pea (Pisum sativum L.), durum wheat (Triticum durum L. cv. Kyle) and soft wheat (Triticum aestivum L. cv. Fielder) were grown at Outlook, Saskatchewan, Canada, under irrigated and dryland conditions. Carbon isotope discrimination (Δ) and water-use efficiency (W), defined as grams of above ground dry matter produced per kilogram water used, were negatively correlated in the six field-grown crops. In irrigated plants Δ remained relatively constant (20–21‰) throughout the growing season. However, in dryland plants, Δ declined in response to the progressive depletion of stored soil water (Polish canola, 20-2-18-8‰; mustard, 19.9–18 5‰; pea, 19.9–17 2‰ durum wheat, 19.7–16.4‰; Argentine canola, 19.4–17.6‰; soft wheat, 19.0–17.4‰). Although there were genetic differences in Δ among the species, water availability was the major factor controlling Δ.