Drought response strategies define the relative contributions of hydraulic dysfunction and carbohydrate depletion during tree mortality.

Plant survival during drought requires adequate hydration in living tissues and carbohydrate reserves for maintenance and recovery. We hypothesized that tree growth and hydraulic strategy determines the intensity and duration of the 'physiological drought', thereby affecting the relative contributions of loss of hydraulic function and carbohydrate depletion during mortality. We compared patterns in growth rate, water relations, gas exchange and carbohydrate dynamics in three tree species subjected to prolonged drought. Two Eucalyptus species (E. globulus, E. smithii) exhibited high growth rates and water-use resulting in rapid declines in water status and hydraulic conductance. In contrast, conservative growth and water relations in Pinus radiata resulted in longer periods of negative carbon balance and significant depletion of stored carbohydrates in all organs. The ongoing demand for carbohydrates from sustained respiration highlighted the role that duration of drought plays in facilitating carbohydrate consumption. Two drought strategies were revealed, differentiated by plant regulation of water status: plants maximized gas exchange, but were exposed to low water potentials and rapid hydraulic dysfunction; and tight regulation of gas exchange at the cost of carbohydrate depletion. These findings provide evidence for a relationship between hydraulic regulation of water status and carbohydrate depletion during terminal drought.

The fate of hydraulically redistributed water in a semi-arid zone eucalyptus species.

Although hydraulic redistribution has been observed for a range of tree species, including Eucalyptus kochii subsp. borealis (C. Gardner) D. Nicolle, there is limited direct evidence that water taken up by deep roots in moist soil is in fact exuded by shallow roots in dry soil. This paper reports an experiment designed to test this hypothesis. Water enriched with deuterium was added to the groundwater via a slotted tube at 4.5 m depth below 5-year-old E. kochii subsp. borealis trees. Nocturnal sap flow increased markedly immediately after deep irrigation, indicating that the trees were using water from this depth. Two weeks later, samples of surface soil and xylem water were found to contain levels of deuterium up to 30% higher than soils and xylem water from a control plot upslope of the main treatment plot. This is strong evidence that trees used groundwater and that efflux of important amounts of hydraulically redistributed water occurred via the roots of E. kochii subsp. borealis.

Observed and modelled leaf area index in Eucalyptus globulus plantations: tests of optimality and equilibrium hypotheses.

This paper reports on variation in leaf area index (L) in five Eucalyptus globulus Labill. plantations in response to application of nitrogen, thinning at age 2 years and variation in climate wetness index (the ratio of rainfall to potential evaporation). Observed L is compared with: (i) L predicted to optimize net primary productivity for a given average annual temperature, annual water use and potential evaporation (L(opt)) and (ii) L calculated as a linear function of climate wetness index (L(eq)). L peaked in fertilized plots at between 4 and 5 years of age or immediately after canopy closure. The value of L from canopy closure to age 8 years was not strongly related to annual rainfall or climate wetness index. At two sites with total soil nitrogen <1.2 mg g(-)(1), L in fertilized plots was about two units greater than in unfertilized plots. This difference persisted until measurements ended in 2004 when the trees were 8 years old. The L of plots thinned to 300 and 600 stems ha(-)(1) at age 2 years recovered quickly and was not significantly different from L in unthinned plots when the trees were 8 years old. L(opt) was a good predictor of the leaf area index of 8-year-old plots of E. globulus when nitrogen and phosphorus were non-limiting (model efficiency (EF) was 0.5). For the same plots, L(eq) underestimated observed L by an average of two units, and the model efficiency was low (-3.25). Data from two nitrogen-limited sites demonstrated that for fertilized plots L(opt) (EF = 0.6) was a much better predictor of L than L(eq) (EF = -3.36). At the same sites, L(eq) (EF = 0.42) was a better model for predicting L of unfertilized plots than L(opt) (-3.59). These results provide evidence that comparing observed L with L(opt) can identify stands limited by factors other than growing climate.

Plasticity in the Huber value contributes to homeostasis in leaf water relations of a mallee Eucalypt with variation to groundwater depth.

Information on how vegetation adapts to differences in water supply is critical for predicting vegetation survival, growth and water use, which, in turn, has important impacts on site hydrology. Many field studies assess adaptation to water stress by comparing between disparate sites, which makes it difficult to distinguish between physiological or morphological changes and long-term genetic adaptation. When planting trees into new environments, the phenotypic adaptations of a species to water stress will be of primary interest. This study examined the response to water availability of Eucalyptus kochii ssp. borealis (C. Gardner) D. Nicolle, commonly integrated with agriculture in south-western Australia for environmental and economic benefits. By choosing a site where the groundwater depth varied but where climate and soil type were the same, we were able to isolate tree response to water supply. Tree growth, leaf area and stand water use were much larger for trees over shallow groundwater than for trees over a deep water table below a silcrete hardpan. However, water use on a leaf area basis was similar in trees over deep and shallow groundwater, as were the minimum leaf water potential observed over different seasons and the turgor loss point. We conclude that homeostasis in leaf water use and water relations was maintained through a combination of stomatal control and adjustment of sapwood-to-leaf area ratios (Huber value). Differences in the Huber value with groundwater depth were associated with different sapwood-specific conductivity and water use on a sapwood area basis. Knowledge of the coordination between water supply, leaf area, sapwood area and leaf transpiration rate for different species will be important when predicting stand water use.

Growth efficiency increases as relative growth rate increases in shoots and roots of Eucalyptus globulus deprived of nitrogen or treated with salt.

We used calorimetry to test whether there is a single general relationship between growth and respiration in shoots and roots of Eucalyptus globulus Labill. seedlings when stressed, irrespective of the type or severity of stress. We found that nitrogen (N) deprivation and salt treatment had no effect on the relationship between growth and respiration and little effect on absolute rates of respiration. Carbon-conversion efficiency (epsilonC) ranged from 0.7 to 0.9 for specific growth rates (R(SG)) greater than 0.3 day(-1). Above an R(SG) of 0.1 day(-1), epsilonC decreased gradually with decreasing R(SG) and between an R(SG) of 0- 0.1 day(-1), epsilonC decreased rapidly. We conclude that the relationship between epsilonC and R(SG) is not greatly affected by salt or N-deprivation stresses. Relationships between gross productivity and epsilonC may be generally applicable, in which case they could improve on the "flat-tax" approach to modeling net primary productivity from gross productivity while avoiding the complexity of more explicit models of plant respiration. However, the relationship between gross productivity and epsilonC was sensitive to temperature and the effect of temperature on epsilonC thus requires further investigation. Nitrogen deprivation caused large decreases in leaf area and shoot to root ratio, and mature leaves of N-deprived plants had lower intrinsic water-use efficiencies than leaves of plants well supplied with nutrients. Nitrogen deprivation increased apical dominance and most of the reduction in leaf area was the result of fewer secondary branches, although leaf size was also reduced. Our results suggest that N deprivation reduces productivity primarily by reducing sink size, rather than sink activity, and that apical dominance may be an important mechanism for maintaining adequate epsilonC in resource-limited conditions.