Summarized data of lichen polysaccharides and their origin
1. Introduction
Lichens are symbiotic organisms composed of a fungal partner, the mycobiont, and one or more photosynthetic partners, the photobiont, which may be either a green alga or a cyanobacterium (Nash III, 1996; Wolseley and Aguirre-Hudson, 1994 and Yoshimura, 2004). The mycobiont in this combination is an ascomycete or a basidiomycete and the photobiont a green alga or a cyanobacterium. Some lichen species can contain more than one green algal species as photobionts (Friedl and Gärtner, 1988; Friedl, 1989; Ihda et al., 1993). At present, about 13,500 fungal species have been recognized to be involved in lichen symbiosis (Hawksworth et al. 1995; Kirk et al., 2001). However, Sipman and Aptroot (2001) stated that this number could be as high as 20,000 after including “orphaned” species; according to Lumbsch
The hyphae of mycobionts are septate, branched, thin or thick walled, and the walls are colourless or variously coloured. It is necessary that the photobiont, which is essentially aquatic in nature, remains protected from desiccation in a lichenized terrestrial condition. The protection is provided by the mycobiont which forms the bulk of the thallus, by the development of specialized hyphal tissues in the form of a cover or cortex over the stratum of the photobiont. The development of the cortex is assumed to be stimulated by the photobiont (Ahmadjian, 1987). In some gelatinous lichens with cyanobacteria, the polysaccharidic sheath produced by the photobiont (cyanobacteria) contributes to water retention (Prieto et al., 2008). Lichens have been used for ecological, medicinal and other economic purposes for over 100 years and these beneficial effects have been correlated to some extent with their polysaccharide content. Amongst the identified lichens so far, about 100 species have been studied for their polysaccharides and their composition (Cordeiro et al., 2005).
2. Separation and characterization
The cell wall of fungi is composed mainly of polysaccharides such as lichenan, isolichenan, and galactomannan (fig.1) (Elix and Stocker-Wörgötter, 2008). Bernard and Latgé (2001) described that the fungal wall is a complex structure composed typically of chitin, 1,3-- and 1,6--glucan, mannan and proteins, although wall composition frequently varies markedly between the species of fungi.
For the separation and isolation of lichen polysaccharides, traditional methods and modern techniques are used. Traditional methods basically involved freezing and thawing of the material originally extracted with hot water. Dialysis and ethanol precipitation has been employed for further purification. Also, alkali solutions have been used to extract these compounds. The polysaccharides present in the thalli of
3. Biological source of the polysaccharide fraction
Whether these polysaccharides are produced by the mycobiont or photobiont separately or in symbiosis has been debated for a long time. Takahashi et al. (1979) showed that aqueous extracts from cultivated myco and photobiont had different monosaccharide composition and physical properties. It was also found that while the extracts of the mycobiont had a similar composition to that of the parent intact lichen, the photobiont fractions were different from those of the symbiotic thalli and its mycobiont. Complete structural analysis by Cordeiro et al. (2004) confirmed Takahashi’s results, and showed that the nigeran, laminaran and the galactomannan, found previously in the symbiotic thalli of
Three different polysaccharide structural types: β-glucans, α-glucans (linear or lightly substituted), and galactomannans (branched) (Carbonero et al, 2005a) are present in the fungal cell wall. According to Olafsdottir and Ingólfsdóttir (2001), all the polymers present in lichen thalli are categorized into glucan type [β-(1→3)(1→4)], lichenan type [α-(1→3)(1→4)] and pustulan [β-(1→6)]. Some of these types are depicted in Table 1. However, the recent discovery of a few additional complex heteroglycans, such as rhamnogalactofuranan (Olafsdottir, 1999), galactomannoglucans (Woranovicz-Barreira, 1999 and Carbonero et al., 2002) and thamnolan (Carbonero
In the past, lichen polysaccharides have been extracted from the whole thallus without giving consideration to the origin of components such as the fungal partner or the photobiont (Gorin and Iacomini, 1984, 1985; Gorin
|
|
|
|
|
|
Lichenan | Homoglucan with β(1→3) (1→4) linkage |
|
Whole thallus |
Shibata, 1973 | |
Isolichenan | Homoglucan with α(1→3) (1→4) linkage | -do- | -do- | Shibata, 1973 | |
Pustulan | Glucan with β(1→6) linkage |
|
-do- | Shibata, 1973 | |
Everniin | Glucan with α(1→3) (1→4) linkage |
|
-do- | Shibata, 1973 | |
Acroscyphan | Homoglucan with α(1→3) (1→4) (1→6) linkage |
|
-do- | Shibata, 1973 | |
Galacto-glucomannans | a (1→6)-linked main chain of -Manp |
O-2 and O-4 by α-Galp and β-Galp nonreducing end-units |
Parmotrema austrosinense, P. delicatulum, P. mantiqueirense, P. schindlerii, P. tinctorum and Rimelia (R. cetrata and R. reticulata) |
mycobiont |
Carbonero |
Xylorhamno-galactofuranan | (1→3)-linked galactofuranosyl |
galactofuranosyl units 5- |
|
Photobiont |
Cordeiro |
laminaran and pustulan and galacto- furanomannan |
(1→3)-and (1→6)-linked β-glucans | (1→6)-linked α-mannopyranosyl |
|
Mycobiont |
Carbonero |
Hetero- polysaccharide |
(1→5)-linked galactofuranosyl units | Complex |
|
Photobiont |
Cordeiro |
O-methylated mannogalactan | (1→6)-linked β-galactopyranose |
at O-3 by β-Gal |
|
Photobiont |
Cordeiro |
Colleman like | Complex heteroglycan |
2-OMe Man |
|
Whole thallus (cyanobacteria) |
Jensen |
(1→4)-linked β-D-xylan |
|
Mycobiont |
Ruthes |
4. Polysaccharides as a taxonomic tool
The identification and classification is generally based on morphology of the organism. Taxonomy of lichen species have been corroborated by phylogenetic applications with the advances in DNA technology. Lichen polysaccharides have been used as a taxonomic tool and chemotaxonomic classification has resulted in clarification of conflicting taxonomic data.
The lichen-forming ascomycete order Lichinales comprises around 250 species and is distributed among 52 genera and four families (Eriksson, 2006). Earlier molecular studies (Wedin et al., 2005) did not confirm its phylogenetic relationships, although the order was treated as a separate class,
Investigation of mannose containing polysaccharides as a taxonomic tool centers around the structural diversity of the galactomannans isolated from several lichenized fungi. The taxonomic value of these galactomannans depends on the side-chain substituents on (1 →6)-linked α-D-mannopyranosyl main chains (Gorin and Lacomini, 1985).
Although classical taxonomy regarded
Lichen polysaccharides, can also be used as a taxonomic tool to differentiate some lichen species, since some of the heteropolysaccharides and their chemical characters are unique to certain groups of lichens (Carbonero et al., 2006 and Cordeiro et al., 2007). Further, the polysaccharides content of the lichen photobiont may be used as a marker in algal symbiont taxonomy (Cordeiro et al., 2007).
5. Biological activities of lichen polysaccharides
Many lichens are known to have immunomodulating properties, potent antibiotic, antitumour, antiviral as well as antioxidant properties which are mostly attributed to the secondary metabolites (Malhotra et al.,2008, Behera, et al., 2007). According to Yanaki et al., (1986) and Bohn and BeMiller, (1995), functional activity of polysaccharides mainly depends on molecular weight, degree of branching, water solubility, structure and configuration. Hence they have different uses in different fields. The biological activities of lichen polysaccharides reported have been limited to anti-tumor, anti-inflamatory or immunomodulatory activity (Omarsdottir et al., 2007).
Cordeiro et al., (2008) reported that one of the β-galactofuranan polysaccharides isolated from
The cytotoxic activity, phagocytic activity and antitumor activity of an α-D glucan from
According to Nishikawa et al., (1970), O-acetylated pustulan isolated from three species of
Behera et al., (2007) showed correlation between lichen protein/polysaccharide ratio and their antioxidant properties. Since cultured lichen extracts were used in the study, the effect of secondary metabolites as well as polyphenols present in the extract have not been evaluated. Thus the antioxidant activity cannot be solely attributed to the polysaccharide specially since polyphenols are known antioxidants. In addition, the mechanism of scavenging activity of polysaccharides on free radicals is not fully understood yet.
6. Conclusion
Polysaccharides from lichens unlike bacterial polysaccharides do not show a wide range of variation in sugar content. The predominant sugars are limited to glucose, galactose and mannose, with arabinose and xylose present in minor proportions in addition to others such as rhamnose. These polysaccharides however have been useful in chemotaxonomic studies due to this conservation of sugar structures and regular structural patterns. As far as biological activity is concerned, very few studies have been reported and it would be worthwhile further investigating the immunomodulating effects of the polysaccharides.
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