Medicago sativa var. Gabes is a perennial glycophyte that develops new shoots even in high salinity (150 mM NaCl). In the upper exporting leaves, K(+) is high and Na(+) is low by comparison with the lower leaves, where Na(+) accumulation induces chlorosis after 4 weeks of NaCl treatment. By secondary ion mass spectroscopy, a low Na(+)/K(+) ratio was detected in the phloem complex of blade veins in these lower leaves. By transmission electron microscopy, the ultrastructural features were observed in the phloem complex. In the upper leaves of both control and NaCl-treated plants, companion cells in minor veins were found to be transfer cells. These cells may well be involved in the intravenous recycling of ions and in Na(+) flowing out of exporting leaves. Under the effect of NaCl, companion cells in the main veins develop transfer cell features, which may favor the rate of assimilate transport from exporting leaves toward meristems, allowing the positive balance necessary for the survival in salt conditions. These features no longer assist the lower leaves when transfer cells are necrotized in both minor and main veins of NaCl-treated plants. As transfer cells are the only degenerating phloem constituent, our observations emphasize their role in controlling nutrient (in particular, Na(+)) fluxes associated with the stress response.
The perennial Medicago sativa cv. Gabs is widely grown on saline soils in Tunisian oases. The mechanisms by which this NaCl-tolerant cultivar maintains a positive growth balance were analyzed. In this plant of considerable agronomic interest, biochemical analyses were conducted in order to study the effects of salinity on mature leaves. Free-radical detoxification mechanisms and changes induced by the accumulation of reactive oxygen species (ROS) in response to the NaCl stress were compared between the upper (young) and lower (old) carbohydrate source leaves. Long-term NaCl (150 mM) treatment significantly reduced the size of source leaves supporting growth. Salinity damage was greater in the lower than in the upper leaves. This damage was associated with a high Na + : K + ratio and a decrease in the activity of H 2 O 2 -scavenging enzymes, leading to lipid peroxidation. In lower source leaves that were mainly affected by ionic stress, superoxide dismutase (SOD) was overexpressed and guaiacol peroxidase (GPX) activity increased. In contrast, in upper source leaves that were mainly exposed to water deficit, catalase and ascorbate peroxidase (APX) activities increased whereas GPX activity was unchanged. The upper source leaves maintained adequate ionic and water status and an efficient ROS detoxification, allowing sinks to be supplied with photoassimilates and maintaining a positive growth balance in this cultivar of alfalfa.
We studied the distribution of wall ingrowth (WI) polymers by probing thin sections of companion cells specialized as transfer cells in minor veins of Medicago sativa cv Gabès blade with affinity probes and antibodies specific to polysaccharides and glycoproteins. The wall polymers in the controls were similar in WIs and in the primary wall but differently distributed. The extent of labeling in these papillate WIs differed for JIM5 and JIM7 homogalacturonans but was in the same range for LM5 and LM6 rhamnogalacturonans and xyloglucans. These data show that WI enhancement probably requires arabinogalactan proteins (JIM8) mainly localized on the outer part of the primary wall and WIs. By comparison, NaCl-treated plants exhibited cell wall polysaccharide modifications indicating (1) an increase in unesterified homogalacturonans (JIM5), probably implicated in Na(+) binding and/or polysaccharide network interaction for limiting turgor variations in mesophyll cells; (2) enhancement of the xyloglucan network with an accumulation of fucosylated xyloglucans (CCRC-M1) known to increase the capacity of cellulose binding; and (3) specific recognition of JIM8 arabinogalactan proteins that could participate in both wall enlargement and cohesion by increasing the number of molecular interactions with the other polymers. In conclusion, the cell wall polysaccharide distribution in enlarged WIs might (1) participate in wall resistance to sequestration of Na(+), allowing a better control of hydric homeostasis in mesophyll cells to maintain metabolic activity in source leaves, and (2) maintain tolerance of M. sativa to NaCl.
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