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Summary Under salinity, Vitis spp. rootstocks can mediate salt (NaCl) exclusion from grafted V. vinifera scions enabling higher grapevine yields and production of superior wines with lower salt content. Until now, the genetic and mechanistic elements controlling sodium (Na+) exclusion in grapevine were unknown. Using a cross between two Vitis interspecific hybrid rootstocks, we mapped a dominant quantitative trait locus (QTL) associated with leaf Na+ exclusion (NaE) under salinity stress. The NaE locus encodes six high-affinity potassium transporters (HKT). Transcript profiling and functional characterization in heterologous systems identified VisHKT1;1 as the best candidate gene for controlling leaf Na+ exclusion. We characterized four proteins encoded by unique VisHKT1;1 alleles from the parents, and revealed that the dominant HKT variants exhibit greater Na+ conductance with less rectification than the recessive variants. Mutagenesis of VisHKT1;1 and TaHKT1.5-D from bread wheat, demonstrated that charged amino acid residues in the eighth predicted transmembrane domain of HKT proteins reduces inward Na+ conductance, and causes inward rectification of Na+ transport. The origin of the recessive VisHKT1;1 alleles was traced to V. champinii and V. rupestris. We propose that the genetic and functional data presented here will assist with breeding Na+-tolerant grapevine rootstocks.

Potted grapevines of 140 Ruggeri (Vitis berlandieri × Vitis rupestris), a good Cl(-) excluder, and K 51-40 (Vitis champinii × Vitis riparia 'Gloire'), a poor Cl(-) excluder, and of a family obtained by crossing the two genotypes, were used to examine the inheritance of Cl(-) exclusion. Rooted leaves were then used to further investigate the mechanism for Cl(-) exclusion in 140 Ruggeri. In both a potting mix trial (plants watered with 50 mM Cl(-)) and a solution culture trial (plants grown in 25 mM Cl(-)), the variation in Cl(-) accumulation was continuous, indicating multiple rather than single gene control for Cl(-) exclusion between hybrids within the family. Upper limits of 42% and 35% of the phenotypic variation in Cl(-) concentration could be attributed to heritable sources in the potting mix and solution culture trials, respectively. Chloride transport in roots of rooted leaves of both genotypes appeared to be via the symplastic pathway, since addition of 8-hydroxy-1,3,6-pyrenetrisulphonic acid (PTS), an apoplastic tracer, revealed no obvious PTS fluorescence in the laminae of either genotype, despite significant accumulation of Cl(-) in laminae of K 51-40 during the PTS uptake period. There was no significant difference in either unidirectional (36)Cl(-) flux (10 min) or (36)Cl(-) uptake (3 h) into roots of rooted leaves exposed to 5, 10, or 25 mM Cl(-). However, the percentage of (36)Cl(-) transported to the lamina (3 h) was significantly lower in 140 Ruggeri than in K 51-40, supporting reduced Cl(-) loading into xylem and implicating the root stele in the Cl(-) exclusion mechanism.

Background and Aims: Rootstocks influence the inorganic ion and organic acid composition of grapes of the scion variety. The aim was to investigate the impact of rootstocks on the inter-relationship of inorganic ions and organic acid anions in the skin and pulp of grapes and in resultant wine. Methods and Results: Vines were irrigated with water having electrical conductivities in the range 1.6–2.1 dS/m. Chloride, sodium, potassium, malic and tartaric acid concentrations were higher in almost all cases in skin than in pulp. Significant positive correlations existed between chloride and sodium concentrations in both pulp and skin. A significant negative linear regression existed between malic acid and both chloride and sodium concentrations in skin of Chardonnay berries. There were positive linear regressions in chloride concentration between berry (pulp and skin) and resultant wine chloride in both Chardonnay and Shiraz. Conclusion: The higher malic acid and lower chloride concentrations in skin of most grafted Chardonnay and Shiraz vines, and vice versa for own rooted vines, may indicate competition for similar transporter proteins involved in loading into skins. Alternatively, higher salt concentrations in skins may be associated with accelerated malic acid catabolism. Significance of the Study: Chloride-excluding rootstocks demonstrated advantages through reduced chloride (but not sodium) in pulp and skin of grape berries and in resultant wines. Where rootstocks reduced chloride concentrations in skin of grape berries, there is potential for higher malic acid in skin and in the resultant red wines.

Shiraz and Chardonnay grapevines that had been initially assessed for Cl- exclusion in the 1996 and 1997 harvest seasons at both Merbein (inland region of Victoria) and at Padthaway (sub-coastal region of South Australia), were re-evaluated in 2003 and 2004 at those same two locations. Both scion varieties were grown either on their own roots or were grafted to one of the following rootstocks: Ramsey, 1103 Paulsen, 140 Ruggeri, K51-40, Schwarzmann, 101-14, Rupestris St George and 1202 Couderc. Both sites had been irrigated with saline water since 1994. The salinity of irrigation water at Merbein was approximately 2.1 dS/m; at Padthaway irrigation salinity varied between approximately 1.6 dS/m and 2.5 dS/m during the survey period. Changes in the Cl--excluding ability of all rootstock/scion combinations between 1996, 1997 and 2003, 2004 were based upon analysis of Cl- concentrations in grape juice and in laminae at harvest, as well as Electrical Conductivity (EC), pH and Sodium Adsorption Ratio (SAR) of the 1:5 soil:water extract. Taking juice Cl- as indicative of rootstock effectiveness for Cl- exclusion, and considering data from Merbein, our analyses demonstrated that overall means for Shiraz juice Cl- increased from around 190 to 427 mg/L over the survey period (7 years). In Chardonnay grapevines at Merbein, overall mean concentrations of Cl- in juice increased from around 70 to around 225 mg/L over 7 years. Significant differences between rootstocks were evident, with some rootstocks at Merbein showing a diminished capacity for Cl- exclusion in 2003 and 2004. By contrast, in Padthaway there was no consistent deterioration in Cl--excluding capacity by rootstocks supporting either Shiraz or Chardonnay as scions. In 2004 there were significant differences between grapevines at Merbein and Padthaway in the concentration of Cl- accumulated, even though the mean soil EC1:5 after harvest for 0–90 cm depth was the same at both sites, namely 0.4 dS/m. Containment of grapevine salinity at Padthaway (relative to Merbein) between 1996/97 and 2003/04 was most likely due to factors such as a lower volume of saline irrigation, double the rainfall and 27% lower pan evaporation.

This paper describes variation in transpiration efficiency 'W' (where W = dry matter produced/water transpired) among grapevine genotypes grown under well-watered conditions in either a glasshouse or a vineyard. Nineteen genotypes were grown in a glasshouse where growth and transpiration were measured. W ranged from 2.5 to 3.4 g dm/kg H2O transpired. Carbon-isotope discrimination () of laminae dry matter ranged from 20.8 to 22.7?x2030; and there was a negative relationship (R2 = 0.58) between W and . A large proportion of variation in W could be attributed to variation in stomatal conductance. Genotypic variation in photosynthetic capacity was also an important component of variation in W. In a second experiment, lamina was measured for mature field-grown Shiraz and Chardonnay, grown either on their own roots or grafted to five different rootstocks, and maintained at three sites under well-watered conditions. At all sites and regardless of rootstock, the laminae of Chardonnay had values 1 to 2?x2030; lower than Shiraz. There was also a 1 to 2?x2030; variation among the sites. Rootstock variety affected values inconsistently and by a maximum of 0.5?x2030;. Leaf gas exchange measurements were performed at a single site on sun-exposed leaves of Chardonnay and Shiraz on either their own roots or 1103 Paulsen, a moderate to high vigour rootstock. There was no significant effect of rootstock on leaf gas exchange and photosynthetic rates did not differ between scion varieties. However, Chardonnay had a 20% lower stomatal conductance and a 1.4-fold higher ratio of CO2 assimilation/H2O transpiration (A/T) indicating a potentially higher W, at a leaf level, for Chardonnay compared with Shiraz. We conclude that photosynthetic capacity was also higher for Chardonnay. values, predicted from the Ci/Ca ratio calculated from leaf gas exchange measurements, did not differ significantly from measured values for laminae . This similarity for , in conjunction with the fact that the lower of Chardonnay was reflected in a higher A/T ratio, suggests that may be a reliable predictor of comparative W under vineyard conditions.

Seasonal evapotranspiration (ET) was determined for Sultana grapevines grown on their own roots (Own-rooted) or grafted onto Ramsey rootstock (Grafted), and irrigated with water of three salinity levels – low (0.4 dS m–1), medium (1.8 dS m–1) and high (3.6 dS m–1) – during the 1994/1995 growing season in south-eastern Australia. Transpiration (T) was determined from sap flux, soil evaporation (E s) with a model, and soil water (S) with a neutron probe. Total ET for the season was similar for both Own-rooted and Grafted, averaging 382 mm. However, Grafted partitioned a mean of 193.5 mm (50.8%) of the ET through T compared to 146.7 mm (38.4%) by Own-rooted. Daily rates of T were generally low and attained peaks of 1.2 mm (9.9 l per vine) for Grafted and 0.9 mm (7.5 l) for Own-rooted in late November, and changed very little until after harvest in February. In contrast to T, the E s rate was consistently higher for Own-rooted than for Grafted from November onwards, and at the end of the season totalled 237 mm for Own-rooted compared to 187 mm for Grafted. Differences between Own-rooted and Grafted in their partitioning of ET into T and E s were associated with their canopy development. Grafted had a higher rate of canopy development than Own-rooted, and in mid-season, the former intercepted about 50% more incident radiation than Own-rooted. The crop factors, i. e. ratio of water use to evaporative demand, based on ET were similar for both vine types with an average seasonal value of 0.25, but when based on T were 0.12 for Grafted and 0.10 for Own-rooted. The ratio of fresh fruit weight to total water used at harvest, i. e. crop water use efficiency (CWUE), based on ET, had a mean of 86 kg mm–1 ha–1 for Grafted and 43 kg mm–1 ha–1 for Own-rooted, and when based on T, was 165 and 115 kg mm–1 ha–1, respectively; however, supplementary data obtained during the 1993/1994 season, indicated a CWUE based on T of 294 and 266 kg mm–1 ha–1 for Grafted and Own-rooted, respectively. Salinity did not have significant effects on canopy development and water use for most of the 1994/1995 growing season. The study shows ET and crop factors for the drip-irrigated grapevines to be much lower than previously reported for this district.

Plant cation-chloride cotransporters (CCCs) have been implicated in conferring salt tolerance. They are predicted to improve shoot salt exclusion by directly catalyzing the retrieval of sodium (Na(+)) and chloride (Cl(-)) ions from the root xylem. We investigated whether grapevine (Vitis vinifera [Vvi]) CCC has a role in salt tolerance by cloning and functionally characterizing the gene from the cultivar Cabernet Sauvignon. Amino acid sequence analysis revealed that VviCCC shares a high degree of similarity with other plant CCCs. A VviCCC-yellow fluorescent protein translational fusion protein localized to the Golgi and the trans-Golgi network and not the plasma membrane when expressed transiently in tobacco (Nicotiana benthamiana) leaves and Arabidopsis (Arabidopsis thaliana) mesophyll protoplasts. AtCCC-green fluorescent protein from Arabidopsis also localized to the Golgi and the trans-Golgi network. In Xenopus laevis oocytes, VviCCC targeted to the plasma membrane, where it catalyzed bumetanide-sensitive (36)Cl(-), (22)Na(+), and (86)Rb(+) uptake, suggesting that VviCCC (like AtCCC) belongs to the Na(+)-K(+)-2Cl(-) cotransporter class of CCCs. Expression of VviCCC in an Arabidopsis ccc knockout mutant abolished the mutant's stunted growth phenotypes and reduced shoot Cl(-) and Na(+) content to wild-type levels after growing plants in 50 mm NaCl. In grapevine roots, VviCCC transcript abundance was not regulated by Cl(-) treatment and was present at similar levels in both the root stele and cortex of three Vitis spp. genotypes that exhibit differential shoot salt exclusion. Our findings indicate that CCC function is conserved between grapevine and Arabidopsis, but neither protein is likely to directly mediate ion transfer with the xylem or have a direct role in salt tolerance.