Abstract
The presence of sulfated polysaccharides in cell walls of seaweeds is considered to be a consequence of the physiological adaptation to the high salinity of the marine environment. Recently, it was found that sulfated polysaccharides were present in certain freshwater Cladophora species and some vascular plants. Cladophora (Ulvophyceae, Chlorophyta) is one of the largest genera of green algae that are able to grow in both, seas and freshwater courses. Previous studies carried out on the water-soluble polysaccharides of the marine species C. falklandica established the presence of sulfated xylogalactoarabinans constituted by a backbone of 4-linked β-L-arabinopyranose units partially sulfated mainly on C3 and also on C2 with partial glycosylation, mostly on C2, with terminal β-D-xylopyranose or β-D-galactofuranose units. Besides, minor amounts of 3-, 6- and/or 3,6-linked β-D-galactan structures, with galactose in the pyranosic form were detected. In this work, the main water soluble cell wall polysaccharides from the freshwater alga Cladophora surera were characterized. It was found that this green alga biosynthesizes sulfated polysaccharides, with a structure similar to those found in marine species of this genus. Calibration of molecular clock with fossil data suggests that colonization of freshwater environments occurred during the Miocene by its ancestor. Therefore, the presence of sulfated polysaccharides in the freshwater green macroalga C. surera could be, in this case, an adaptation to transient desiccation and changes in ionic strength. Retention of sulfated polysaccharides at the cell walls may represent a snapshot of an evolutionary event, and, thus constitutes an excellent model for further studies on the mechanisms of sulfation on cell wall polysaccharides and environmental stress co-evolution.
Highlights
It has been proposed that the presence of sulfated polysaccharides in cell walls of seaweeds and marine angiosperms, absent in terrestrial plants, is a consequence of the physiological adaptation to the marine environment (Kloareg and Quatrano, 1988; Aquino et al, 2005) due to a strong environmental pressure
Specimens of the green alga were collected in a freshwater environment (Quequén Grande river, 38◦27 39 S 58◦45 39 W) located in Buenos Aires Province, Argentina (Supplementary Figure S1)
Calibration of molecular clock with fossil data suggests that colonization of freshwater environments probably occurred 11.4 million years ago (MYA; 4–25 MYA 95% Posterior Probabilities (PP)) by the ancestor of the four freshwater species closely related to C. surera (Figure 1B); or even before, at about 65.5 MYA (35–115 MYA 95%PP), during the diversification of the more basal species Rhizoclonium hieroglyphicum
Summary
It has been proposed that the presence of sulfated polysaccharides in cell walls of seaweeds and marine angiosperms, absent in terrestrial plants, is a consequence of the physiological adaptation to the marine environment (Kloareg and Quatrano, 1988; Aquino et al, 2005) due to a strong environmental pressure. Most of sulfated polysaccharides present in seaweed (e.g., carrageenans) are well known to exhibit the solubility characteristics typical of hydrophilic colloids due to the presence of hydroxyl and sulfate groups in their backbones Under certain conditions, they can form hydrogels, three-dimensional networks capable of maintaining a large amount of water. The glycophyte Oryza sativa Linnaeus, when exposed to salt stress (200 mM NaCl) did not induce the biosynthesis of sulfated polysaccharides, but increased the concentration of carboxylated polysaccharides of the pectin type (Aquino et al, 2011) These data suggested that the presence of sulfated polysaccharides in marine plants is an adaptation to high-salinity environments, which may have been conserved during plant evolution from marine green algae (Aquino et al, 2011)
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