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Proteins and traces of polysaccharide are the only polymeric colloids consistently transported in the xylem sap of plants. The hypothesis that such proteins could have physical inhibitory effects on xylem water transport was investigated. Ovalbumin, with a molecular weight of 45 kDa and a molecular diameter of 5.4 nm, is an inert, water-soluble protein that is midway along the size range of endogenous xylem sap proteins. Solutions of ovalbumin conjugated to a fluorescent marker and supplied to transpiring shoot explants of tobacco (Nicotiana tabacum L.) and olive (Olea europaea L.) were shown by confocal laser scanning microscopy to accumulate specifically at wall-based pit membranes that connect neighbouring xylem conduits. In addition, pressure-induced perfusion of micro-filtered ovalbumin solutions, at concentrations similar to those of endogenous xylem sap proteins, through the xylem of tobacco stem or olive twig segments resulted in the retention of c. 40% of the ovalbumin and reductions in the axial hydraulic conductance of the xylem. Smaller molecules such as Texas Red 3000 (MW 3 kDa) and Alexafluor 488-cadaverin conjugates (MW 0.64 kDa) did not show similar characteristics. The partial reduction in xylem hydraulic conductance appeared to be related to the accumulation of ovalbumin at xylem pit membranes and the consequent fouling of trans-membrane water-conducting pores with smaller diameters than those of the ovalbumin molecules. Potential implications of these novel findings for whole-plant water relations are considered.

Background: Drought is a major limitation to plant productivity. Various options are available for increasing water availability and sustaining growth of crop plants in drought-prone environments., Scope: After a general introduction to the problems of water availability, this review focuses on a critical evaluation of recent progress in unravelling mechanisms for modifying plant growth responses to drought., Conclusions: Investigations of key regulatory mechanisms integrating plant growth responses to water deficits at the whole-organism, cellular and genomic levels continue to provide novel and exiting research findings. For example, recent reports contradict the widespread conception that root-derived abscisic acid is necessarily involved in signalling for stomatal and shoot-growth responses to soil water deficits. The findings bring into question the theoretical basis for alternate-side root-irrigation techniques. Similarly, recent reports indicate that increased ABA production or increased aquaporin expression did not lead to improved drought resistance. Other reports have concerned key genes and proteins involved in regulation of flowering (FT), vegetative growth (DELLA), leaf senescence (IPT) and desiccation tolerance (LEA). Introgression of such genes, with suitable promoters, can greatly impact on whole-plant responses to drought. Further developments could facilitate the introduction by breeders of new crop varieties with growth physiologies tailored to improved field performance under drought. Parallel efforts to encourage the introduction of supplementary irrigation with water made available by improved conservation measures and by sea- or brackish-water desalination, will probably provide comprehensive solutions to coping with drought-prone environments.

Several types of small, endogenous signal peptides are now known to induce a wide range of local and systemic responses in plants, but how such signal peptide activity is transported over long distances remains unclear. In particular, the possible occurrence and root-to-shoot transport of signal peptide activity in the xylem does not appear to have been previously investigated. Suspension-cultured cells of wild tomato Lycopersicon peruvanium L. were used in an established bioassay for detecting nanomolar concentrations of signal peptides via the induction of alkalinizing activity. Xylem sap naturally exuded from the cut and washed stem-surfaces of de-topped tomato plants (Lycopersicon esculentum L. cv. Castlemart) was collected, partially purified, concentrated, and shown by the bioassay consistently to contain significant alkalinizing activity. Plant salinity treatment induced further small increases in activity. Subsidiary experiments indicated that the alkalinizing activity found in the xylem-sap had properties similar to those of known plant signal peptides and was root derived. Thus, it was (i) detectable within minutes, (ii) eluted similarly during HPLC chromatography, (iii) destroyed by incubation with proteases and stable in the presence of protease inhibitor cocktail, and (iv) not found in bioassays of simulated xylem sap placed on the cut stem-surfaces of non-exuding roots in order to detect any significant release of wound peptides from the stem. Further investigations of the signal peptide activity in root xylem sap could provide new insights into its identity, genes, receptors, origins, and possible hormonal roles in regulating shoot growth and development.

Water deficit caused by addition of polyethylene glycol 6000 at -0.5 MPa water potential to well-aerated nutrient solution for 48 h inhibited the elongation of maize (Zea mays) seedling primary roots. Segmental growth rates in the root elongation zone were maintained 0 to 3 mm behind the tip, but in comparison with well-watered control roots, progressive growth inhibition was initiated by water deficit as expanding cells crossed the region 3 to 9 mm behind the tip. The mechanical extensibility of the cell walls was also progressively inhibited. We investigated the possible involvement in root growth inhibition by water deficit of alterations in metabolism and accumulation of wall-linked phenolic substances. Water deficit increased expression in the root elongation zone of transcripts of two genes involved in lignin biosynthesis, cinnamoyl-CoA reductase 1 and 2, after only 1 h, i.e. before decreases in wall extensibility. Further increases in transcript expression and increased lignin staining were detected after 48 h. Progressive stress-induced increases in wall-linked phenolics at 3 to 6 and 6 to 9 mm behind the root tip were detected by comparing Fourier transform infrared spectra and UV-fluorescence images of isolated cell walls from water deficit and control roots. Increased UV fluorescence and lignin staining colocated to vascular tissues in the stele. Longitudinal bisection of the elongation zone resulted in inward curvature, suggesting that inner, stelar tissues were also rate limiting for root growth. We suggest that spatially localized changes in wall-phenolic metabolism are involved in the progressive inhibition of wall extensibility and root growth and may facilitate root acclimation to drying environments.

Growth in the apical elongation zone of plant roots is central to the development of functional root systems. Rates of root segmental elongation change from accelerating to decelerating as cell development proceeds from newly formed to fully elongated status. One of the primary variables regulating these changes in elongation rates is the extensibility of the elongating cell walls. To help decipher the complex molecular mechanisms involved in spatially variable root growth, we performed a gene identification study along primary root tips of maize (Zea mays) seedlings using suppression subtractive hybridization (SSH) and candidate gene approaches. Using SSH we isolated 150 non-redundant cDNA clones representing root growth-related genes (RGGs) that were preferentially expressed in the elongation zone. Differential expression patterns were revealed by Northern blot analysis for 41 of the identified genes and several candidate genes. Many of the genes have not been previously reported to be involved in root growth processes in maize. Genes were classified into groups based on the predicted function of the encoded proteins: cell wall metabolism, cytoskeleton, general metabolism, signaling and unknown. In-situ hybridization performed for two selected genes, confirmed the spatial distribution of expression shown by Northern blots and revealed subtle differences in tissue localization. Interestingly, spatial profiles of expression for some cell wall related genes appeared to correlate with the profile of accelerating root elongation and changed appropriately under growth-inhibitory water deficit.

Growth of elongating primary roots of maize (Zea mays) seedlings was approximately 50% inhibited after 48 h in aerated nutrient solution under water deficit induced by polyethylene glycol 6000 at -0.5 MPa water potential. Proton flux along the root elongation zone was assayed by high resolution analyses of images of acid diffusion around roots contacted for 5 min with pH indicator gel. Profiles of root segmental elongation correlated qualitatively and quantitatively (r(2) = 0.74) with proton flux along the surface of the elongation zone from water-deficit and control treatments. Proton flux and segmental elongation in roots under water deficit were remarkably well maintained in the region 0 to 3 mm behind the root tip and were inhibited from 3 to 10 mm behind the tip. Associated changes in apoplastic pH inside epidermal cell walls were measured in three defined regions along the root elongation zone by confocal laser scanning microscopy using a ratiometric method. Finally, external acidification of roots was shown to specifically induce a partial reversal of growth inhibition by water deficit in the central region of the elongation zone. These new findings, plus evidence in the literature concerning increases induced by acid pH in wall-extensibility parameters, lead us to propose that the apparently adaptive maintenance of growth 0 to 3 mm behind the tip in maize primary roots under water deficit and the associated inhibition of growth further behind the tip are related to spatially variable changes in proton pumping into expanding cell walls.