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Background and Aims The study compared the viticultural performance and wine sensory attributes of Shiraz grafted on eight rootstocks with a range in vine vigour and grown under drip‐irrigation with moderately saline water. Methods and Results The rootstocks were 140 Ruggeri, 1103 Paulsen, 110 Richter, Ramsey, 101‐14, M5489, M5512 and M6262. Ramsey and 140 Ruggeri had the highest leaf area index and M6262 the lowest. Ramsey had highest yield, followed by 140 Ruggeri > M5489 = 110 Richter > M5512 > 1103 Paulsen = 101‐14 > M6262. Rootstock M6262 had highest wine chloride concentration of 482 and 384 mg/L in 2012 and 2013, respectively, but salty taste and a thickness mouthfeel (viscosity) were significant only in 2012. Wines from the other seven rootstocks had a chloride and sodium concentration under 175 and 40 mg/L, respectively. Colour density was highest in wine from 101‐14 in 2012 and from 101‐14, 1103 Paulsen and M5489 in 2013. A salt tolerance index (STI) for the eight rootstocks was calculated based on yield, leaf area index, grape juice chloride and sodium concentration and wine colour density. Conclusions Rootstock M6262 had lowest STI in both years. M5489 had highest STI in 2013 followed by 110 Richter = M5512 = 101‐14 = 140 Ruggeri. The recognition threshold for salty taste may lie between a chloride concentration in wine of between 384 and 482 mg/L. Significance of the Study The STI, based on five wine grape performance parameters, provides a promising measure of rootstock salt tolerance compared with the traditional parameters of yield and/or capacity for salt exclusion.

Sodium (Na+) exclusion is an important rootstock trait, which contributes to the overall salt tolerance of a grafted vine by limiting Na+ accumulation in both leaves and berries. A population of 68 hybrid Vitis genotypes, presumed to be from a cross between the rootstocks ‘K51-40’ and ‘140 Ruggeri’, have previously been investigated for their ability to exclude Na+. Here we report that the genotyping of this population with eight short sequence repeat (SSR) markers showed that ‘140 Ruggeri’ is the male parent of only 40 individuals. The rootstock ‘Schwarzmann’ is the male parent of 18 genotypes, with the remaining 10 genotypes being derived from unknown pollen sources. The role of HKT1;1 in Na+ exclusion in the 40 ‘K51-40’ × ‘140 Ruggeri’ hybrids has previously been reported, with four unique alleles of this gene present in this family, with each parent heterozygous for a dominant allele (EK or ER) linked with strong exclusion and for a recessive allele (eK or eR) linked with poor exclusion. In the present study, we found ‘Schwarzmann’ is also heterozygous for HKT1;1 alleles, consisting of dominant (EK) and recessive (eK) alleles identical to those of ‘K51-40’. The 18 ‘K51-40’ × ‘Schwarzmann’ hybrids were scored for inheritance of these alleles. When screened for the Na+ exclusion phenotype, two homozygous eKeK genotypes accumulated less Na+ than eKeR hybrids having ‘140 Ruggeri’ as the pollen parent. Thus, it appears that the recessive eR allele from ‘140 Ruggeri’ results in higher Na+ accumulation than the recessive eK allele from ‘K51-40’ or ‘Schwarzmann’, and this is supported by functional Na+ transport rates of the encoded HKT1;1 proteins previously reported.

Background and Aims Rooted leaves were used to analyse the salt exclusion phenotype in grapevines. Genotypes included rootstocks 140 Ruggeri and K51‐40 and cultivar Cabernet Sauvignon – respectively, good, poor and intermediate chloride excluders. Methods and Results We investigated the effect of short‐term salt treatment on the chloride, sodium and potassium concentration of organs and whole rooted leaves and the time course of chloride accumulation in salt‐treated, whole rooted leaves. The effect of Control and salt plus low and high nitrate concentration on chloride and nitrate concentration in organs was assessed. Salt treatment increased chloride and sodium concentration in all organs but had no effect on potassium concentration. The chloride concentration of whole rooted leaves and the sodium concentration of lamina and petiole were similar between salt‐treated rooted leaves of 140 Ruggeri and K51‐40; 140 Ruggeri accumulated more chloride in roots and less in leaf (petiole and lamina) than K51‐40. Roots of salt‐treated 140 Ruggeri and K51‐40 responded to higher external nitrate by decreasing chloride concentration and increasing nitrate concentration. Conclusions Restricted transport of chloride to the leaf and greater storage in roots is a feature of the chloride exclusion phenotype in rooted leaves. The short‐term capacity of 140 Ruggeri for chloride exclusion from the leaf was reduced in the salt plus high nitrate treatment. This was linked to higher nitrate and reduced chloride accumulation in roots. Significance of the Study The chloride exclusion phenotype in rooted leaves was shown to involve greater partitioning to roots and less to the leaf, rather than differences in net chloride accumulation by whole rooted leaves.

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.

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

Background and Aims The effect of rootstock on chloride (Cl−) and sodium (Na+) exclusion capacity and yield of Chardonnay and Shiraz was investigated in the final 2 years (2007/08 and 2008/09) of a long-term (14-year) saline irrigation trial. Methods and Results Chardonnay and Shiraz, on own roots and on nine rootstocks, were irrigated for the first 12 seasons with water of electrical conductivity (ECiw) of 2.1 dS/m, with 7.15 ML/ha the mean total applied. The final two seasons were years of water restrictions, where the vines were irrigated with water of (mean) ECiw of 1.65 dS/m at a seasonal application of 3.04 ML/ha. Vines on own roots had a relatively low yield, for example 2.0 kg/vine (Chardonnay) and 8.8 kg/vine (Shiraz), and a high concentration of both chloride and sodium in grape juice (>1000 mg/L for Chardonnay and >500 mg/L for Shiraz). Ramsey and 1103 Paulsen rootstocks produced a high yield of 11–12 kg/vine with Chardonnay, and Ramsey and C7 produced a high yield of 23.5–24.5 kg/vine with Shiraz. Chardonnay on C5 and Shiraz on C7 rootstocks had the lowest concentration of grape juice chloride and sodium (in each case 10 years) study to compare the effect of rootstocks on the tolerance of Chardonnay and Shiraz to salinity.

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: Within-site comparisons were made of rootstock effects on yield, and chloride and sodium concentrations in petioles, juice and wine of Shiraz and Chardonnay vines at sites with irrigation water salinities (ECiw) ranging from low (0.4 dS/m) to moderate-high (1.8 to 3.3 dS/m). It also compared consistency of yield performance of the various rootstocks with both scions over 8 years at one site with an ECiw of 2.1 dS/m. Methods and Results: Chardonnay and Shiraz on own roots and on Ramsey, 1103 Paulsen, 140 Ruggeri, K51-40, Schwarzmann, 101-14, Rupestris St. George and 1202 Couderc were compared. Ramsey resulted in better yields relative to most of the other rootstocks at three of the four sites for each scion. Exceptions were the low salinity site where Schwarzmann was best with Chardonnay, and Padthaway where 140 Ruggeri was best with Shiraz. Chardonnay wine chloride concentrations were similar to grape juice chloride concentrations, but Shiraz wine chloride concentrations were on average 1.7-fold higher than grape juice chloride. Conclusions: Shiraz on own roots, K51-40 and 1202C rootstocks carry some risk of accumulating unacceptable levels of chloride in grape juice and wine when the salinity of the irrigation water is at moderate to high levels. Rootstocks K51-40 (with Chardonnay and Shiraz) and potentially 101-14 (with Shiraz) should be avoided in situations of long term irrigation with moderate to high salinity water. Significance of the Study: The study identifies rootstocks with acceptable yields and grape juice chloride concentrations for potential use in regions affected by salinity.

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.

Field-grown vines of cv. Sultana on either their own roots or grafted to a range of rootstocks, were drip-irrigated at one of three salinity levels (0.40, 1.75 and 3.50 dS/m) over a five year period. Rootstocks were Ramsey, 1103 Paulsen, J17-69 and 4 hybrids (designated R1, R2, R3 and R4) derived from parentage involving Vitis champini, V. berlandieri and V. vinifera. Grape juice total soluble solids, titratable acidity and pH were measured at harvest, while colour of dried fruit was measured before and after processing and again after six months storage. Damage index (an indicator of skin damage) was measured post-processing; sugar crystal formation in dried grapes and dried grape compaction were measured post-storage. There was a strong salinity x rootstock interaction for grape juice soluble solids concentration, soluble solids yield (the product of soluble solids concentration and fruit yield) and pH, but not for titratable acidity when analysed on the basis of 5 year means. Small increases (< 5%) in juice soluble solids concentration were recorded at medium salinity (1.75 dS/m) for the low vigour genotypes, Sultana on own-roots and on J17-69 rootstock, based on the 5 year means and Fisher's protected (interaction) LSDs. Moderate increases (< 10%) also occurred in years of low crop load e.g. 1993 at high salinity for the high vigour rootstock R2 and in 1995 at medium salinity for Sultana on own roots and at high salinity for Sultana on R1 rootstock. By contrast, decreases in soluble solids concentration occurred with increasing salinity for the high vigour rootstocks (Ramsey, 1103 Paulsen and R2) in years of high crop load e.g. 1992. Small (< 2%) increases in grape juice pH were recorded at high salinity for Sultana on R3 rootstock and moderate increases (< 7%) in grape juice titratable acidity were recorded at high salinity for Sultana on own roots and Sultana on J17-69, R1, R2 and R4 rootstocks. Dried grapes from all treatments achieved a light amber colour (quality grade termed 5 crown light) and were generally of high quality. Sultanas from own-rooted grapevines were redder (higher ‘a-value’) than sultanas from 1103 Paulsen and Ramsey when assessed as unprocessed fruit, after processing (both years) and after 6 weeks storage. While soluble solids yields per vine were 23–31% lower at high salinity for Sultana on own roots and on R1, R3 and R4 rootstocks, they were unaffected by high salinity for Ramsey, 1103 Paulsen and R2 rootstocks. Moreover, soluble solids yields for Sultana on Ramsey, 1103 Paulsen and R2 rootstocks were 1.4 to 2.5-fold higher than for Sultana on the other rootstocks at high salinity. This study has shown that over a 5 year period rootstocks such as Ramsey, 1103 Paulsen and R2 grafted with Sultana were tolerant of salinity, producing dried grapes of generally high quality.

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 study was conducted over three seasons on irrigated Shiraz grapevines growing in a warm climate. We addressed The question of whether differences in berry size (within a population of berries from minimally pruned, own-rooted or Ramsey-grafted vines), would lead to differences in juice composition, wine composition or wine sensory score. Predictably, berry mass was found to increase with seed number, but berries in the smallest mass categories (0.3-0.7 g) still had similar juice soluble solids and pH; and similar concentrations of K+, tartaric acid and malic acid, compared with larger berries (1.4-2.0 g). Only by the very smallest mass category (0.3-0.55 g) was there any indication of better colour density (both by own-rooted and Ramsey-grafted vines) or higher anthocyanin concentration (for own-rooted vines) compared with larger berries (1.4-2.0 g). Concentrations of tartaric acid and K~ in berry skins were highest in the smallest berry mass categories (0.3-0.7 g( and decreased with increasing berry mass (up to 1.4-2.0 g). A strong correlation (R2 = 0.85 )between skin tartaric acid and K+ concentrations was observed across that range. Small-scale wine lost based on small berries (0.8-0.9 g) versus large berries (1.2-1.3 g) showed no differences in measures such as soluble solids, total acids or pH ob juice; nor any differences in pH, total acids, K+, tartrate, malate, spectral characteristics or sensory score of corresponding wines. Moreover, small berries had a similar skin to fruit ratio, and a similar juice yield, compared lo large berries. However, when measured post-fermentation, the ratio of seed weight to skin weight was higher for small berries. The mass range ob berries used here for small-scale winemaking (i.e. from 0.8-0.9 g up to 1.2-1.3 g), covered the range of Shiraz berry mass typically bound in irrigated vineyards (from 0.8 lo 1.5 g(, and thus confirms The relevance of present outcomes to practical winemaking. Finally, our data lot variation in juice and wine composition as a function of berry size, showed consistent trends for all seasons, and thus implies that reported instances of improved wine quality from small berries (often associated with certain pruning treatments nr deficit irrigation strategies), are more likely due to treatment effects that lead to small Fruit, rather than to intrinsic developmental differences between large and small berries

Field vines of cv. Sultana, grown either on their own roots, or grafted to a range of rootstocks, were drip-irrigated at three salinity levels (0.40, 1.75 and 3.50 dS/m) over a five-year period. Rootstocks comprised Ramsey, 1103 Paulsen, J17-69 and 4 hybrids (designated R1, R2, R3 and R4) derived from parentage involving Vitis champini, V. berlandieri and V. vinifera. Concentrations of Cl-, Na+, K+, Ca2+ and Mg2+ were measured in petioles at flowering, and in laminae and grape juice at harvest, in each year of the trial. Vines on all rootstocks accumulated less chloride in either petioles at flowering or in laminae and juice at harvest compared with vines on own roots at all salinity treatments. By inference, all rootstocks behaved as chloride excluders relative to the roots of own-rooted vines. 1103 Paulsen was the best chloride excluder based on lowest concentrations of accumulated Cl- in petioles, laminae and grape juice at high salinity. Sultana on R3 rootstock at high salinity accumulated more Na+ in both laminae and grape juice (at harvest) than did Sultana on own roots or on any of the other rootstocks. Laminae K+ at harvest time was reduced at high salinity in Sultana on own roots and on all rootstocks. Concentrations of both Cl- and Na+ in petioles at flowering and in laminae and grape juice at harvest showed no significant correlation with either yield (as kg of fresh grapes per vine) or vigour (as measured by fresh weight of one-year-old pruning wood per vine) for any salinity treatment. There was however, a strong positive correlation between yield and the subsequent weight of one-year-old pruning wood for all salinity treatments. There was also a negative correlation between Na+ concentrations in petioles at flowering and the subsequent weight of one year-old-pruning wood from the 0.40 dS/m treatment. Similar negative correlations were found between Na+ concentration in both laminae and grape juice at harvest time, and the subsequent weight of one-year-old pruning wood from the 0.40 dS/m treatment (but not from either the 1.75 or 3.50 dS/m treatments). Based on these findings and those from Walker et al. 2002a we conclude that a high innate vigour of a rootstock combined with moderate to high chloride and sodium exclusion ability represents the best combination for salt tolerance in Sultana grapevines as measured by yield at moderate to high salinity.

Vegetative growth and yield of grapevines grown in the field on their own roots or grafted to a range of rootstocks were investigated under drip irrigation with water of three salinity levels (0.40, 1.75 and 3.50 dS/m) over a five-year period. Rootstocks were Ramsey, 1103 Paulsen, J17-69 and 4 hybrids (designated R1, R2, R3 and R4) derived from parentage involving Vitis champini, V. berlandieri and V. vinifera. Of measured yield components (bunches per shoot, bunches per vine, weight per bunch, weight per berry and total yield), only weight per berry was significantly reduced by high salinity (3.50 dS/m) in each year of the trial with the exception of Sultana on 1103 Paulsen and R2 in 1991 and Sultana on Ramsey in 1993. Weights of one-year-old pruning wood were also reduced by high salinity in all years for own roots and all rootstocks, with the exception of R2. Mean yield values at each salinity level over the five-year period of the trial were highest for Sultana on Ramsey, 1103 Paulsen and R2. High salinity had no effect on five-year mean yields of Sultana on Ramsey, 1103 Paulsen and R2. Yield (five-year means) of Sultana on Ramsey and R2 at 1.75 dS/m were significantly higher than at 0.40 dS/m by 14.6% and 13.4% respectively. In contrast, 5-year mean yields of Sultana on J17-69, own roots, R1, R3 and R4 at 3.50 dS/m were reduced by 54, 30, 20, 30 and 30% respectively. Yield of Sultana on J17-69, R1 and R4 rootstocks was reduced by 47, 20 and 24% respectively at 1.75 dS/m. When yield was regressed against bunches per vine and weight per bunch for Sultana on own roots and on Ramsey rootstock, bunches per vine was the main determinant of yield, while weight per berry showed a poor correlation with yield at all salinity levels. Rootstock ranking for salt tolerance based on yield at high salinity was the same as rankings for pruning wood weights at high salinity. The same occurred at medium salinity, demonstrating that vigour imparted by the rootstock was a major factor in Sultana salt tolerance as measured by yield. Mean root weighted soil saturation paste electrical conductivities (RWECe) (determined from soil saturation paste salinities and root length densities) were in the range 2.0-2.6 dS/m for the low salinity treatment, increasing to approximately 5.4 dS/m with increasing salinity of irrigation water. Yield reduction for own-rooted vines for each 1.0 dS/m increase in RWECe above 2.6 dS/m was 9.3%.

Background Salt tolerance in grapevine is associated with chloride (Cl−) exclusion from shoots; the rate-limiting step being the passage of Cl− between the root symplast and xylem apoplast. Despite an understanding of the physiological mechanism of Cl− exclusion in grapevine, the molecular identity of membrane proteins that control this process have remained elusive. To elucidate candidate genes likely to control Cl− exclusion, we compared the root transcriptomes of three Vitis spp. with contrasting shoot Cl− exclusion capacities using a custom microarray. Results When challenged with 50 mM Cl−, transcriptional changes of genotypes 140 Ruggeri (shoot Cl− excluding rootstock), K51-40 (shoot Cl− including rootstock) and Cabernet Sauvignon (intermediate shoot Cl− excluder) differed. The magnitude of salt-induced transcriptional changes in roots correlated with the amount of Cl− accumulated in shoots. Abiotic-stress responsive transcripts (e.g. heat shock proteins) were induced in 140 Ruggeri, respiratory transcripts were repressed in Cabernet Sauvignon, and the expression of hypersensitive response and ROS scavenging transcripts was altered in K51-40. Despite these differences, no obvious Cl− transporters were identified. However, under control conditions where differences in shoot Cl− exclusion between rootstocks were still significant, genes encoding putative ion channels SLAH3, ALMT1 and putative kinases SnRK2.6 and CPKs were differentially expressed between rootstocks, as were members of the NRT1 (NAXT1 and NRT1.4), and CLC families. Conclusions These results suggest that transcriptional events contributing to the Cl− exclusion mechanism in grapevine are not stress-inducible, but constitutively different between contrasting varieties. We have identified individual genes from large families known to have members with roles in anion transport in other plants, as likely candidates for controlling anion homeostasis and Cl− exclusion in Vitis species. We propose these genes as priority candidates for functional characterisation to determine their role in chloride transport in grapevine and other plants. Electronic supplementary material The online version of this article (doi:10.1186/s12870-014-0273-8) contains supplementary material, which is available to authorized users.

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.

The use of a heat-pulse technique to monitor sap flow from which transpiration can be deduced was evaluated in ungrafted grapevines (Vitis vinifera L., cv. Sultana) under glasshouse and field conditions. There was a significant degree of agreement between daily transpiration deduced from heat-pulse velocity (T hp) and that determined directly by gravimetry in the glasshouse and by calibration using the Penman-Monteith equation in the field. Comparison throughout the growing season of T hp to transpiration calculated with the full Penman-Monteith equation produced a high coefficient of determination (r 2=0.69). A similar comparison of T hp with transpiration calculated with only the aerodynamic component of the Penman-Monteith equation produced a non-linear relationship, due to the equation over-estimating transpiration relative to T hp at high vapour pressure deficits (i.e. above 2.5 kPa). Values for total seasonal transpiration measured with heat-pulse sensors or calculated with either the full Penman-Monteith equation or with only the modified aerodynamic component of the equation were within 10% of each other. Transpiration (T hp) in grapevines with an adequate supply of soil water was shown to be coupled to ambient air conditions.

Shiraz grapevines on either their own roots, or on the rootstocks Ramsey, 1103 Paulsen, 140 Ruggeri or 101-14, were grown at two separate sites within the Murray-Darling viticultural region with similar irrigation regimes but with an irrigation water salinity of either 0.43 dS/m (low salinity site) or 2.3 dS/m (high salinity site). Rootstock effects on grape berry development, ion concentrations, soluble solids and acidity were followed during one season. Wines were also made and compared using spectral analysis and sensory evaluation. Rootstock effects that were common across both sites were (1) a close relationship between K+ and soluble solids accumulation in developing grape berries which commenced at the onset of veraison and was indicative of a link between K+ and sucrose transport in the phloem, and (2), higher wine K+, pH and colour hue for all rootstocks with one exception, namely 101-14 at high salinity where 101-14 responded similarly to own roots. Juice K+, pH and loss of K+ from juice during winemaking were highest for grapes from the high salinity site. Mean berry weight was smaller and the range in berry size across rootstocks was narrower at the saline site. The narrower range in berry sizes may have contributed to fewer rootstock effects on wine spectral characteristics at high salinity. There was no effect of rootstock on CO2 assimilation rate or stomatal conductance at either site, although intrinsic leaf-based water-use efficiency measured as A/g was 50% higher at the saline site. All treatments exhibited berry shrivel at maturity, but the extent was smaller at high salinity. Slower development of berry colour during veraison was observed on some rootstocks, for example 101-14, and while unrelated to canopy size per se, a higher leaf-to-fruit ratio for 101-14 may have been a factor. Slower berry colour development during veraison had no bearing on the colour density of wine made from the harvested grapes.

Salinity treatments of 0.43, 1.7 and 3.4 dS/m were applied through a drip‐irrigation system to four‐year‐old vines of own‐rooted Sultana (SO) and Sultana on Ramsey rootstock (Sr) The vines were planted in spring 1987 and established for two years under irrigation with low salinity (0.43 dS/m) water before commencing the range of salinity treatments in spring 1989. The effects of salinity and rootstock on yield, size and composition of berries, canopy size, lamina CO2 assimilation, leaf water relations and lamina ion concentrations were studied between veraison and harvest during the second season of salinity treatments. Mature leaves of SR had higher rates of CO2 assimilation (leaf area basis) and stomatal conductance than comparable leaves of SO. The high salinity treatment (3.4 dS/m) reduced CO2 assimilation rate of SO but not of SR. Medium salinity (1.7 dS/m) had no significant effect on CO2 assimilation rate of either type. Laminae of SO accumulated significantly higher concentrations of chloride than SR vines at all salinity levels. There was a significant negative correlation (r2= 0.44) between CO2 assimilation rate and laminae chloride of SO. Leaf sodium concentrations increased with increasing salinity, but concentrations in laminae at high salinity were similar in SO and SR, with SR showing no reduction in CO2 assimilation. Leaf potassium concentrations were higher in SR, but decreased with increasing salinity, whereas magnesium concentrations were similar in SO and SR vines but increased in both at high salinity. Leaf water potential and relative water content were not significantly affected by salinity in either SO or SR vines. Both vine types had reduced total leaf area and pruning wood weights as salinity increased, but they were greater in SR at all salinity levels because of the higher inherent capacity for biomass production in SR vines. The bigger canopies, lower lamina chloride concentrations and ‘normal’ photosynthesis rates of SR vines at 3.4 dS/m enabled these vines to mature crops with similar berry weights, sugar contents and fruit yield compared with those of SO vines at 0.43 dS/m. The research demonstrated the benefit of using Ramsey rootstock for Sultana under saline field conditions and provided a physiological explanation for their higher salt tolerance.

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.