Hydraulic lift is the passive movement of water from roots into soil layers with lower water potential, while other parts of the root system in moister soil layers, usually at depth, are absorbing water. Here, we review the brief history of laboratory and field evidence supporting this phenomenon and discuss some of the consequences of this below-ground behavior for the ecology of plants. Hydraulic lift has been shown in a relatively small number of species (27 species of herbs, grasses, shrubs, and trees), but there is no fundamental reason why it should not be more common as long as active root systems are spanning a gradient in soil water potential (Ψ) and that the resistance to water loss from roots is low. While the majority of documented cases of hydraulic lift in the field are for semiarid and arid land species inhabiting desert and steppe environments, recent studies indicate that hydraulic lift is not restricted to these species or regions. Large quantities of water, amounting to an appreciable fraction of daily transpiration, are lifted at night. This temporary partial rehydration of upper soil layers provides a source of water, along with soil moisture deeper in the profile, for transpiration the following day and, under conditions of high atmospheric demand, can substantially facilitate water movement through the soil-plant-atmosphere system. Release of water into the upper soil layers has been shown to afford the opportunity for neighboring plants to utilize this source of water. Also, because soils tend to dry from the surface downward and nutrients are usually most plentiful in the upper soil layers, lifted water may provide moisture that facilitates favorable biogeochemical conditions for enhancing mineral nutrient availability, microbial processes, and the acquisition of nutrients by roots. Hydraulic lift may also prolong or enhance fine-root activity by keeping them hydrated. Such indirect benefits of hydraulic lift may have been the primary selective force in the evolution of this process. Alternatively, hydraulic lift may simply be the consequence of roots not possessing true rectifying properties (i.e., roots are leaky to water). Finally, the direction of water movement may also be downward or horizontal if the prevailing Ψ gradient so dictates, i.e., inverse, or lateral, hydraulic lift. Such downward movement through the root system may allow growth of roots in otherwise dry soil at depth, permitting the establishment of many phreatophytic species.

We examined patterns of genetic variance and covariance in two traits (i) carbon stable isotope ratio delta13C (dehydration avoidance) and (ii) time to flowering (drought escape), both of which are putative adaptations to local water availability. Greenhouse screening of 39 genotypes of Arabidopsis thaliana native to habitats spanning a wide range of climatic conditions, revealed a highly significant positive genetic correlation between delta13C and flowering time. Studies in a range of C3 annuals have also reported large positive correlations, suggesting the presence of a genetically based trade-off between mechanisms of dehydration avoidance (delta13C) and drought escape (early flowering). We examined the contribution of pleiotropy by using a combination of mutant and near-isogenic lines to test for positive mutational covariance between delta13C and flowering time. Ecophysiological mutants generally showed variation in delta13C but not flowering time. However, flowering time mutants generally demonstrated pleiotropic effects consistent with natural variation. Mutations that caused later flowering also typically resulted in less negative delta13C and thus probably higher water use efficiency. We found strong evidence for pleiotropy using near-isogenic lines of Frigida and Flowering locus C, cloned loci known to be responsible for natural variation in flowering time. These data suggest the correlated evolution of delta13C and flowering time is explained in part by the fixation of pleiotropic alleles that alter both delta13C and time to flowering.

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Diel soil water potential fluctuations reflected daytime depletion and nocturnal resupply of water in upper soil layers. Transpiration suppression experiments demonstrated that water absorption by roots caused the daytime depletion. The soil water potential data and experimental results suggest that at night water absorbed from moist soil by deeper roots is transported to and lost from roots into drier upper soil layers. The deeper roots appear to absorb and transport water both day and night. Implications for the efficiency of deep roots and water storage, nutrient uptake and water parasitism in upper soil layers are discussed.