Journal Article

Evolution of mixed-linkage (1 → 3, 1 → 4)-β-<span class="smallCaps">d</span>-glucan (MLG) and xyloglucan in <i>Equisetum</i> (horsetails) and other monilophytes

Xinxin Xue and Stephen C. Fry

in Annals of Botany

Published on behalf of The Annals of Botany Company

Volume 109, issue 5, pages 873-886
Published in print April 2012 | ISSN: 0305-7364
Published online February 2012 | e-ISSN: 1095-8290 | DOI:
Evolution of mixed-linkage (1 → 3, 1 → 4)-β-d-glucan (MLG) and xyloglucan in Equisetum (horsetails) and other monilophytes

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  • Ecology and Conservation
  • Evolutionary Biology
  • Plant Sciences and Forestry


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Background and Aims

Horsetails (Equisetopsida) diverged from other extant eusporangiate monilophytes in the Upper Palaeozoic. They are the only monilophytes known to contain the hemicellulose mixed-linkage (1 → 3, 1 → 4)-β-d-glucan (MLG), whereas all land plants possess xyloglucan. It has been reported that changes in cell-wall chemistry often accompanied major evolutionary steps. We explored changes in hemicelluloses occurring during Equisetum evolution.


Hemicellulose from numerous monilophytes was treated with lichenase and xyloglucan endoglucanase. Lichenase digests MLG to di-, tri- and tetrasaccharide repeat-units, resolvable by thin-layer chromatography.

Key Results

Among monilophytes, MLG was confined to horsetails. Our analyses support a basal trichotomy of extant horsetails: MLG was more abundant in subgenus Equisetum than in subgenus Hippochaete, and uniquely the sister group E. bogotense yielded almost solely the tetrasaccharide repeat-unit (G4G4G3G). Other species also gave the disaccharide, whereas the trisaccharide was consistently very scarce. Tetrasaccharide : disaccharide ratios varied interspecifically, but with no consistent difference between subgenera. Xyloglucan was scarce in Psilotum and subgenus Equisetum, but abundant in subgenus Hippochaete and in the eusporangiate ferns Marattia and Angiopteris; leptosporangiate ferns varied widely. All monilophytes shared a core pattern of xyloglucan repeat-units, major XEG products co-chromatographing on thin-layer chromatography with non-fucosylated hepta-, octa- and nonasaccharides and fucose-containing nona- and decasaccharides.


G4G4G3G is the ancestral repeat-unit of horsetail MLG. Horsetail evolution was accompanied by quantitative and qualitative modification of MLG; variation within subgenus Hippochaete suggests that the structure and biosynthesis of MLG is evolutionarily plastic. Xyloglucan quantity correlates negatively with abundance of other hemicelluloses; but qualitatively, all monilophyte xyloglucans conform to a core pattern of repeat-unit sizes.

Keywords: Equisetum; Hippochaete; Equisetum bogotense; ferns; eusporangiate monilophytes; leptosporangiate monilophytes; evolution; cell wall (primary); hemicellulose; mixed-linkage beta-glucan; xyloglucan

Journal Article.  6850 words.  Illustrated.

Subjects: Ecology and Conservation ; Evolutionary Biology ; Plant Sciences and Forestry

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