Mark A. Bacon

Biomass allocation in tomato (Lycopersicon esculentum) plants grown under partial rootzone drying: enhancement of root growth

Tomato (Lycopersicon esculentum Mill.) plants were grown in either a glasshouse (GH) or a controlled environment cabinet (CEC) to assess the effects of partial rootzone drying (PRD) on biomass allocation. Control and PRD plants received the same amounts of water. In control plants, water was equally distributed between two compartments of a split-root system. In PRD plants, only one compartment was watered while the other was allowed to dry. At the end of each drying cycle, wet and dry compartments were alternated. In the GH, total biomass did not differ between PRD and control plants after four cycles of PRD, but PRD increased root biomass by 55% as resources were partitioned away from shoot organs. In the CEC, leaf water potential did not differ between treatments at the end of either of two cycles of PRD, but stomatal conductance of PRD plants was 20% less at the end of the first cycle than at the beginning. After two cycles of PRD in the CEC, biomass did not differ between PRD and control plants, but PRD increased root biomass by 19% over the control plants. The promotion of root biomass in PRD plants was associated with the alternation of wet and dry compartments, with increased root biomass occurring in the re-watered compartment after previous exposure to soil drying. Promotion of root biomass in field-grown PRD plants may allow the root system to access resources (water and nutrients) that would otherwise be unavailable to control plants. This may contribute to the ability of PRD plants to maintain similar leaf water potentials to conventionally irrigated plants, even when smaller irrigation volumes are supplied.

Alternation of wet and dry sides during partial rootzone drying irrigation alters root-to-shoot signalling of abscisic acid.

Partial rootzone drying (PRD) is an irrigation technique where water is distributed unevenly to the root system such that part is irrigated while the remainder is allowed to dry the soil. Tomato (Lycopersicon esculentum Mill.) plants were grown with their roots in two soil columns to compare the physiological consequences of alternation of wet and dry columns during PRD irrigation (alternate PRD, PRD-A) with retention of the same wet and dry columns (fixed PRD, PRD-F). When PRD plants received 50% less water than well-watered (WW) plants, xylem ABA concentration ([X-ABA]) increased and stomatal conductance decreased relative to WW plants. Although both sets of PRD plants received the same amount of water, [X-ABA] of PRD-A plants increased up to 2-fold above that of PRD-F plants, which further decreased stomatal conductance. Differences in [X-ABA] were detected within an hour of alternation, but did not persist beyond the photoperiod of alternation. [X-ABA] increased linearly as whole-pot soil water content (θpot) and leaf water potential (Ψleaf) declined, but the difference in [X-ABA] between the two sets of PRD plants was not due to differences in either θpot or Ψleaf. In PRD-F plants, the unwatered part of the root system contributes proportionally less to the transpiration stream as the soil progressively dries (Yao et al. 2001, Plant, Cell & Environment 24, 227-235). In PRD-A plants, we hypothesise that re-watering the dry part of the root system allows these roots to contribute proportionally more to total sap flux, thus liberating a pulse of ABA to the transpiration stream as the root ABA pool accumulated during soil drying is depleted. Since the enhancement of [X-ABA] caused by PRD-A increased as θpot and Ψleaf declined, an optimal frequency of alternation to maximise the cumulative physiological effects of this ABA pulse must consider possible negative impacts of leaf water deficit as soil water status declines.

Adaptation of Roots to Drought

In this chapter we examine the morphological, physiological and biochemical adaptations of roots to drought and discuss how roots perceive soil drying and communicate such information to the shoots.

The biochemical control of leaf expansion during drought

Over the past decade it has become clear that we cannot always explain the observed reduction in leaf expansion rates during drought by measuring the plants water relations. This has led us to question the possibility of a role for the cell wall and its biochemical machinery in controlling the rate of leaf expansion during drought. However, if we are to reject or modify previous assumptions regarding the control of leaf expansion during drought, then we must offer alternative explanations. This article addresses recent work from this laboratory and in the literature, concerning the involvement of cell wall-enzymes, pH and abscisic acid (ABA) in regulating leaf expansion during water deficit.