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J Exp Bot
2016 Aug 01;6715:4495-505. doi: 10.1093/jxb/erw237.
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SLAH1, a homologue of the slow type anion channel SLAC1, modulates shoot Cl- accumulation and salt tolerance in Arabidopsis thaliana.
Qiu J
,
Henderson SW
,
Tester M
,
Roy SJ
,
Gilliham M
.
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Salinity tolerance is correlated with shoot chloride (Cl(-)) exclusion in multiple crops, but the molecular mechanisms of long-distance Cl(-) transport are poorly defined. Here, we characterize the in planta role of AtSLAH1 (a homologue of the slow type anion channel-associated 1 (SLAC1)). This protein, localized to the plasma membrane of root stelar cells, has its expression reduced by salt or ABA, which are key predictions for a protein involved with loading Cl(-) into the root xylem. Artificial microRNA knockdown mutants of AtSLAH1 had significantly reduced shoot Cl(-) accumulation when grown under low Cl(-), whereas shoot Cl(-) increased and the shoot nitrate/chloride ratio decreased following AtSLAH1 constitutive or stelar-specific overexpression when grown in high Cl(-) In both sets of overexpression lines a significant reduction in shoot biomass over the null segregants was observed under high Cl(-) supply, but not low Cl(-) supply. Further in planta data showed AtSLAH3 overexpression increased the shoot nitrate/chloride ratio, consistent with AtSLAH3 favouring nitrate transport. Heterologous expression of AtSLAH1 in Xenopus laevis oocytes led to no detectible transport, suggesting the need for post-translational modifications for AtSLAH1 to be active. Our in planta data are consistent with AtSLAH1 having a role in controlling root-to-shoot Cl(-) transport.
Fig. 1. Expression level of AtSLAH1 (A) and AtSLAH3 (B) treated with control (2mM NaCl), 50mM and 100mM NaCl for 7 days, or 20 μM ± cis-trans-ABA for 4 or 16h. Arabidopsis (Col-0) were grown in hydroponics for 5 weeks and exposed to NaCl treatment for 7 days. The ABA was applied 4 or 16h before harvest. Transcripts were detected in the whole root cDNA. Results are presented as means+SEM, n=5. The expression levels were normalized to four control genes (AtGAPDH, AtActin2, AtTubulin and AtCyclophilin). Statistical significance was determined by one-way analysis of variance (ANOVA) and Tukeyâs test (Pâ¤0.05); a, b and c represent data groups that are statistically different from each other.
Fig. 2. Under low Clâ conditions, amiRNA:AtSLAH1 mutants had significantly reduced expression levels of AtSLAH1 and reduced shoot Clâ compared with null segregants. Plants were grown hydroponically for 6 weeks in BNS containing 2mM NaCl (low Clâ conditions). (A) AtSLAH1 expression in roots of all amiRNA-AtSLAH1 mutants (amiRNA:AtSLAH1_1, 2, 3 and 4) and null segregants (nulls). (B) Shoot Clâ accumulation of amiRNA-AtSLAH1 mutants and nulls under low Clâ conditions. (C) Correlation between transcript level of AtSLAH1 and shoot Clâ concentration. (D) The shoot NO3
â/Clâ ratio in all amiRNA:AtSLAH1 mutant and null segregant lines grown under low Clâ conditions. Results are mean+SEM (n>8), except (C), which is ±SEM. Statistical differences determined by one-way ANOVA and Tukeyâs test (Pâ¤0.005); a and b represent statistically significant differences between data groups.
Fig. 3. Under high Clâ conditions, 35S:AtSLAH1 overexpression lines accumulated higher shoot Clâ and showed reduced NO3
â/Clâ ratio compared with null segregants (nulls). Plants were grown hydroponically in BNS until 6 weeks old and then exposed to BNS containing 75mM NaCl (high Clâ conditions) for 7 days. (A) Semi-quantitative RT-PCR of 35S:AtSLAH1 overexpression lines and nulls. (B) Shoot Clâ concentration under high Clâ conditions. (C) Shoot NO3
â/Clâ ratio. (D) Whole shoot biomass (fresh weight) as measured after high Clâ treatment. Results are mean+SEM (n>6). Statistical differences determined by one-way ANOVA and Tukeyâs test (Pâ¤0.05); a and b represent statistically significant differences between data groups.
Fig. 4. Stelar cell type-specific overexpression of AtSLAH1 in E2586 significantly increased shoot Clâ accumulation and reduced shoot biomass under high Clâ conditions. (A) Correlation between shoot Clâ accumulation and relative expression of AtSLAH1 in GAL4:AtSLAH1 overexpression lines under high Clâ (75mM NaCl) supply. Open circles and solid line: GAL4:AtSLAH1_1; open triangle and dash line: GAL4:AtSLAH1_2. Plants were grown hydroponically in BNS for 6 weeks and then exposed to BNS containing 75mM NaCl (high Clâ conditions) for 7 days. Correlation between shoot Clâ concentration and the abundance of AtSLAH1 in GAL4:AtSLAH1_1 (R
2=0.5805, P ⤠0.005, significant deviation from zero), GAL4:AtSLAH1_2 (R
2=0.5395, P ⤠0.005, significant deviation from zero). (B) Whole shoot biomass (fresh weight) as measured after high Clâ treatment. Results are mean+SEM (n>6). Statistical differences determined by one-way ANOVA and Tukeyâs test (Pâ¤0.05); a and b represent statistically significant differences between data groups.
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