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Improved salt tolerance of mycorrhizal plants is commonly attributed to their better mineral nutrition, particularly phosphorus. However, the effect of arbuscular-mycorrhizal (AM) fungi on salt tolerance may not be limited to this mechanism. We investigated the possibility that non-nutritional effects of AM fungi, based on proline accumulation or increased photosynthesis and related parameters, can influence the tolerance of lettuce (Lactuca saliva L.) to salinity. Three levels of salt (3, 4 and 5 g NaCl kg(-1) dry soil) were applied and plants were maintained under these conditions for 7 weeks. The salt-treated AM plants produced greater root and shoot dry weights than unfertilized or P-fertilized non-AM controls. With increasing salinity, both shoot and root dry weights were reduced, but this decrease was greater in uninoculated plants. In particular, shoot dry weight was not reduced in G. fasciculatum-colonized plants as a consequence of salt, whereas in uninoculated plants it was reduced by about 35% at the highest salt level. Proline accumulation was considerably lower for P-amended non-AM and for AM plants except for G. mosseae-colonized plants than was the case for unamended plants. Transpiration, carbon dioxide exchange rate (CER), stomatal conductance and water use efficiency (WUE) were higher in mycorrhizal plants. At 5 g NaCl kg(-1), both photosynthesis and WUE increased by more than 100% in mycorrhizal treatment relative to uninoculated plants. The contents of phosphorus of P-fertilized non-AM plants was similar to or higher than those of G. mosseae- and G. fasciculatum-colonized plants. Plants colonized by G. deserticola had the highest P-content regardless of salt level. Hence, the effect of G. mosseae and G. fasciculatum on salt tolerance in this experiment could not be attributed to a difference in the P content. The mechanisms by which these two fungi alleviated salt stress appeared to be based on physiological processes (increased CER, transpiration, stomatal conductance and WUE) rather than on nutrient uptake (N or P).
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