Progress in understanding the network of mechanisms involved in maize primary root growth maintenance under water deficits is reviewed. These include the adjustment of growth zone dimensions, turgor maintenance by osmotic adjustment, and enhanced cell wall loosening. The role of the hormone abscisic acid (ABA) in maintaining root growth under water deficits is also addressed. The research has taken advantage of kinematic analysis, i.e. characterization of spatial and temporal patterns of cell expansion within the root growth zone. This approach revealed different growth responses to water deficits and ABA deficiency in distinct regions of the root tip. In the apical 3 mm region, elongation is maintained at well-watered rates under severe water deficit, although only in ABA-sufficient roots, whereas the region from 3-7 mm from the apex exhibits maximum elongation in well-watered roots, but progressive inhibition of elongation in roots under water deficit. This knowledge has greatly facilitated discovery of the mechanisms involved in regulating the responses. The spatial resolution with which this system has been characterized and the physiological knowledge gained to date provide a unique and powerful underpinning for functional genomics studies. Characterization of water deficit-induced changes in transcript populations and cell wall protein profiles within the growth zone of the maize primary root is in progress. Initial results from EST and unigene analyses in the tips of well-watered and water-stressed roots highlight the strength of the kinematic approach to transcript profiling.

Seedlings of maize (Zea mays L. cv WF9 x Mo 17) were grown in vermiculite at various water potentials. The primary root continued slow rates of elongation at water potentials which completely inhibited shoot growth. To gain an increased understanding of the root growth response, we examined the spatial distribution of growth at various water potentials. Time lapse photography of the growth of marked roots revealed that inhibition of root elongation at low water potentials was not explained by a proportional decrease in growth along the length of the growing zone. Instead, longitudinal growth was insensitive to water potentials as low as -1.6 megapascal close to the root apex, but was inhibited increasingly in more basal locations such that the length of the growing zone decreased progressively as the water potential decreased. Cessation of longitudinal growth occurred in tissue of approximately the same age regardless of spatial location or water status, however. Roots growing at low water potentials were also thinner, and analysis revealed that radial growth rates were decreased throughout the elongation zone, resulting in greatly decreased rates of volume expansion.anism by which osmotic adjustment occurs in roots growing at low q. Specifically, our aim is to determine the extent to which the maintenance of lower 4i, in the growing zone under water deficits can be attributed to increased rates of solute deposition, or to reduced growth and hence slower rate of osmoticum dilution by volume expansion. In this paper, attention is focused on the effects of low qf,y on the spatial distribution of expansive growth rate. Although the spatial growth pattern at high q,, for roots of maize and other species has been well characterized for many years (8,11), the extent to which the pattern may change at low q is not known. Indeed, despite early recognition that knowledge of how plant growth patterns may be altered by environmental variation facilitates the opportunities to understand the regulation of the growth response (11), relatively little information of this kind is available. Here, we show that both longitudinal and radial growth patterns are altered markedly in roots growing at low qi,. In a forthcoming paper (RE Sharp, TC Hsiao, WK Silk, unpublished data), we combine this information with profiles of qi5 and component solutes to determine effects of low q on solute deposition rates in the root growing zone, and evaluate the relationship of the growth and solute deposition responses to osmotic adjustment.Plant growth is generally decreased when soil water is limited. Root growth is often less inhibited than shoot growth (2, 21), however. A recent study of maize has shown that root growth is intrinsically less sensitive than growth of the aerial plant parts to low water potentials (q') of the growing region (30), indicating some form of internal regulation. Root elongation is of obvious advantage to plants in drying soil, and may be particularly important for seedling establishment because of the vulnerabi...

Shoot and root growth are differentially sensitive to water stress. Interest in the involvement of hormones in regulating these responses has focused on abscisic acid (ABA) because it accumulates in shoot and root tissues under water-limited conditions, and because it usually inhibits growth when applied to well-watered plants. However, the effects of ABA can differ in stressed and non-stressed plants, and it is therefore advantageous to manipulate endogenous ABA levels under water-stressed conditions. Studies utilizing ABA-deficient mutants and inhibitors of ABA synthesis to decrease endogenous ABA levels, and experimental strategies to circumvent variation in plant water status with ABA deficiency, are changing the view of the role of ABA from the traditional idea that the hormone is generally involved in growth inhibition. In particular, studies of several species indicate that an important role of endogenous ABA is to limit ethylene production, and that as a result of this interaction ABA may often function to maintain rather than inhibit shoot and root growth. Despite early speculation that interaction between these hormones may influence many of the effects of water deficit, this topic has received little attention until recently.

Potted maize seedlings were subjected to a single period of water stress. As the severity of water stress increased, measurements were made of leaf and root solute and water potentials, leaf diffusive conductance and leaf and root growth. After day four of the drying cycle, the rate of leaf extension and the development of leaf area were reduced. This reduction correlated well with a reduction in leaf turgor which occurred at this time. A significant accumulation of solutes in the root tips of the unwatered plants resulted in the maintenance of root turgor for the duration of the water stress treatment. Root growth of the unwatered plants was also maintained as the severity of water stress increased. A mild degree of water stress resulted in a net increase in root growth compared to the situation in well-watered plants. The significance of solute regulation and continued root growth for plants growing in drying soil is discussed.

Roots of maize (Zea mays L.) seedlings continue to grow at low water potentials that cause complete inhibition of shoot growth. In this study, we have investigated the role of abscisic acid (ABA) in this differential growth sensitivity by manipulating endogenous ABA levels as an altemative to extemal applications of the hormone. An inhibitor of carotenoid biosynthesis (fluridone) and a mutant deficient in carotenoid biosynthesis (vp 5) were used to reduce the endogenous ABA content in the growing zones of the primary root and shoot at low water potentials. Experiments were performed on 30 to 60 hour old seedlings that were transplanted into vermiculite which had been preadjusted to water potentials of approximately -1.6 megapascals (roots) or -0.3 megapascals (shoots). Growth occurred in the dark at nearsaturation humidity. Results of experiments using the inhibitor and mutant approaches were very similar. Reduced ABA content by either method was associated with inhibition of root elongation and promotion of shoot elongation at low water potentials, compared to untreated and wild-type seedlings at the same water potential. Elongation rates and ABA contents at high water potential were little affected. The inhibition of shoot elongation at low water potential was completely prevented in fluridone-treated seedlings during the first five hours after transplanting. The results indicate that ABA accumulation plays direct roles in both the maintenance of primary root elongation and the inhibition of shoot elongation at low water potentials.A major reason behind the slow progress in the area of crop adaptation to drought is the insufficient basic understanding of the regulation of growth responses to water stress. When water is limited, shoot growth in many species is more inhibited than root growth (24), and in some cases, the absolute biomass of roots has been shown to increase relative to wellwatered controls (12,23). In maize, roots continue to grow at low water potentials that cause complete inhibition of shoot growth (25,30). The role of the hormone ABA in the differential growth responses of the primary root and shoot of maize to low water potentials is the subject of this paper.ABA accumulates to high concentrations in tissues ofplants