Eight wild Hordeum species: H. bogdanii, H. intercedens, H. jubatum, H. lechleri, H. marinum, H. murinum, H. patagonicum, and H. secalinum, and cultivated barley (H. vulgare) were grown in nutrient solution containing 0.2 (control), 150, 300, or 450 mol m(-3) NaCl. In saline conditions, the wild Hordeum species (except H. murinum) had better Na+ and Cl- 'exclusion', and maintained higher leaf K+, compared with H. vulgare. For example, at 150 mol m(-3) NaCl, the K+:Na+ in the youngest, fully expanded leaf blades of the wild Hordeum species was, on average, 5.2 compared with 0.8 in H. vulgare. In H. marinum grown in 300 mol m(-3) NaCl, K+ contributed 35% to leaf psi(pi), whereas Na+ and Cl- accounted for only 6% and 10%, respectively. By comparison, in H. vulgare grown at 300 mol m(-3) NaCl, K+ accounted for 19% and Na+ and Cl- made up 21% and 25% of leaf psi(pi), respectively. At 300 mol m(-3) NaCl, glycinebetaine and proline together contributed almost 15% to psi(pi) in the expanding leaf blades of H. marinum, compared with 8% in H. vulgare. Decreased tissue water content under saline conditions made a substantial contribution to declines in leaf psi(pi) in the wild Hordeum species, but not in H. vulgare. A number of the wild Hordeum species were markedly more salt tolerant than H. vulgare. H. marinum and H. intercedens, as examples, had relative growth rates 30% higher than H. vulgare in 450 mol m(-3) NaCl. Hordeum vulgare also suffered up to 6-fold more dead leaf material (as a proportion of shoot dry mass) than the wild Hordeum species. Thus, several salt-tolerant wild Hordeum species were identified, and these showed an exceptional capacity to 'exclude' Na+ and Cl- from their shoots.

Growth, root aerenchyma, and profiles of radial O2 loss (ROL) along adventitious roots were evaluated in 35 'wild' Hordeum accessions and cultivated barley (H. vulgare L. ssp. vulgare) when grown in stagnant nutrient solution (deoxygenated and containing 0.1% agar). When grown in stagnant solution, accessions from wetland and 'intermediate' habitats were superior, compared with accessions from non-wetland habitats, in maintaining relative growth rate, tillering, and adventitious root mass. Constitutive aerenchyma formation in adventitious roots was ≥�10% in 22 accessions (cf. H. vulgare at 2%). When grown in stagnant solution, aerenchyma was ≥� 20% in the adventitious roots of 14 accessions (cf. H. vulgare at 12%). Variation among the accessions in the volume of aerenchyma formed when grown in aerated or stagnant solution was not determined by the waterlogging regime of the species' natural habitat. However, the genus Hordeum comprises four genomes and when grown in stagnant solution accessions with the X genome formed, on average, 22% aerenchyma in adventitious roots (50 mm behind apex), whereas those with the H genome averaged 19%, and those with the Y or I genomes averaged 16 and 15%, respectively. Sixteen accessions formed a barrier to ROL in the basal region of adventitious roots when grown in stagnant solution. The formation of a barrier to radial O2 loss was predominant in accessions from wet habitats, and absent in accessions from non-wetland habitats. In addition, this trait was only present in accessions with the X or H genomes. The combination of aerenchyma and a barrier to ROL enhances the longitudinal diffusion of O2 within roots towards the apex. The possibility of a link between having a barrier to ROL and the X or H genomes in Hordeum species might, in future studies, enable a genetic analysis of this important trait.

Summary• The subapical regions of wetland plant roots can develop a barrier to radial O 2 loss (ROL), but barrier resistance has rarely been quantified in terms of its O 2 diffusivity.• Barrier resistance in adventitious roots of the waterlogging-tolerant Hordeum marinum was assessed from measurements of ROL using cylindrical platinum electrodes while either varying shoot O 2 partial pressures or cooling the rooting medium. Anatomical features were examined using fluorescence microscopy.• When grown in stagnant agar nutrient solution, a barrier to ROL was induced over the basal half of 100-120-mm-long roots. Autofluorescence in hypodermal cell walls indicated that putative suberin deposition was coincident with barrier expression. Root cooling revealed a significant respiratory component in barrier resistance. Eliminating the respiration effect by manipulating shoot O 2 partial pressures revealed an O 2 diffusivity for the barrier of 5.96 × 10 −7 cm 2 s −1 , 96% less than that at the apex, which was ≥ 1.59 × 10 −5 cm 2 s −1 .• It is concluded that the ROL barrier is a manifestation of two components acting synergistically: a physical resistance caused principally by secondary cell-wall deposits in the outer hypodermal layer; and respiratory activity in the hypodermal/epidermal layers; with physical resistance being the dominant component.

The adventitious roots of Hordeum marinum grown in stagnant deoxygenated solution contain a barrier to radial O2 loss (ROL) in basal zones, whereas roots of plants grown in aerated solution do not. The present experiments assessed whether induction of the barrier to ROL influences root hydraulic conductivity (Lpr). Wheat (Triticum aestivum) was also studied since, like H. marinum, this species forms aerenchyma in stagnant conditions, but does not form a barrier to ROL. Plants were grown in either aerated or stagnant, deoxygenated nutrient solution for 21-28 d. Root-sleeving O2 electrodes were used to assess patterns of ROL along adventitious roots, and a root-pressure probe and a pressure chamber to measure Lpr for individual adventitious roots and whole root systems, respectively. Lpr, measured under a hydrostatic pressure gradient, was 1.8-fold higher for individual roots, and 5.6-fold higher for whole roots systems, in T. aestivum than H. marinum. However, there was no difference in Lpr between the two species when measured under an osmotic driving force, when water moved from cell to cell rather than apoplastically. Root-zone O2 treatments during growth had no effect on Lpr for either species (measured in aerobic solution). It is concluded that induction of the barrier to ROL in H. marinum did not significantly affect the hydraulic conductivity of either individual adventitious roots or of the whole root system.