SummaryPlant 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.

In previous work, we provided evidence from sap flow measurements that when root systems span soil layers of different moisture content, water is redistributed by roots in the direction of the difference in water potential. In addition to the phenomenon termed "hydraulic lift", where water is redistributed from depth to dry topsoil, the process of "hydraulic redistribution" includes downward transfer of water when the surface layers of soils with low permeability become wet after rainfall. In this paper, we support our previous findings with evidence from measurements of soil water and estimate the quantities of water transferred to depth following rain. Amounts of water stored at depth are not likely to be significant for drought avoidance by plants. However, downward transfer of water may be important to plant establishment and the reduction of waterlogging in certain soil types.

In 1986, four allopatric Eucalyptus species (E. camaldulensis Dehnh, E. saligna Smith, E. leucoxylon F. Muell and E. platypus Hook.) were planted together in a 480-mm rainfall zone, in 8-m wide contour belts as part of a plan to minimize waterlogging and secondary salinization. Throughout 1997, 1998 and 1999, there was significant inter-specific variation in predawn leaf water potential (Psi(pd)); however, maximum stomatal conductance (g(sm)) only differed significantly between species in mid to late summer. Relationships between g(sm) and Psi(pd) were significant and showed that stomata of E. camaldulensis were significantly more sensitive to Psi(pd), and presumably soil water potential, than stomata of E. leucoxylon or E. platypus. When applied to the Psi(pd) data, these relationships predicted that g(sm), and by inference transpiration, varied much less between species than Psi(pd). Diurnal measurements throughout the season confirmed this prediction, and showed that E. camaldulensis and E. saligna avoided drought by gaining access to deeper water, whereas E. leucoxylon and E. platypus maintained greater g(sm) at a given water stress than E. camaldulensis or E. saligna. Osmotic potentials measured after rehydration and water release curves of the leaves indicated that different mechanisms accounted for the apparent drought tolerance of E. leucoxylon and E. platypus. In summer, E. leucoxylon reduced osmotic potential at full and zero turgor by similar amounts compared with winter. In summer, E. platypus had a significantly lower bulk elastic modulus and relative water content at turgor loss point than E. camaldulensis, E. saligna or E. leucoxylon. This elastic adjustment resulted in a larger difference between osmotic potential at full and zero turgor in summer than in winter. The inherently low osmotic potential in E. leucoxylon and elastic adjustment in E. platypus resulted in turgor loss at a similar and significantly lower water potential than in E. camaldulensis or E. saligna. These results have implications for species selection for planting to manage groundwater recharge in areas prone to waterlogging and secondary salinization.

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