First report on polysaccharides of Asterochloris and their potential role in the lichen symbiosis

A structural study of the carbohydrates from the aposymbiotically cultured Asterochloris sp., the algal symbiont of the lichen Cladina confusa was carried out for the first time. A xylorhamnogalactofuranan was purified and was predominated by (1-->3)-linked galactofuranosyl units with sidechains in position 6 on approximately 6.4% of the units. The sidechains have galactofuranosyl units 5-O and 6-O-substituted, as well rhamnopyranosyl units 2-O, 3-O and 2,3-di-O-substituted. Xylose was detected only as nonreducing end units, together with galactofuranosyl units. Amylose and a beta-(1-->4)-xylan were also present. These polysaccharides have not been found in the symbiotic thallus of C. confusa, which contained only glucans, galactomannoglucan and galactoglucomannan. A potential role of these carbohydrates in lichen recognition proccess is also discussed.     

Characteristics of yeast lectins and their role in cell-cell interactions

Lectins are ubiquitous proteins with the ability to induce cell agglutination and, mediate cellular and molecular recognition processes in a variety of biological interactions. Fungi display exquisite specificity for target tissues and attach to host glycoconjugates via these sugar-binding proteins. Although only few reports are available on lectin activity of yeasts, these sugar binding proteins have been embraced for their role in cell flocculation, a commercially beneficial property, that simplifies downstream recovery operations in yeast fermentations. The lectins bind to cell wall mannans of the neighboring cells via hydrogen bonds leading to the formation of cell aggregates which get interrupted in the presence of specific sugars. Attachment of pathogenic yeasts to host cell surface is also a consequence of lectin-mediated recognition process. This review provides a brief overview of yeast lectins, with an insight to lectin-mediated cellular recognition phenomenon in yeasts.     

Bacterial communities associated with the lichen symbiosis

Lichens are commonly described as a mutualistic symbiosis between fungi and "algae" (Chlorophyta or Cyanobacteria); however, they also have internal bacterial communities. Recent research suggests that lichen-associated microbes are an integral component of lichen thalli and that the classical view of this symbiotic relationship should be expanded to include bacteria. However, we still have a limited understanding of the phylogenetic structure of these communities and their variability across lichen species. To address these knowledge gaps, we used bar-coded pyrosequencing to survey the bacterial communities associated with lichens. Bacterial sequences obtained from four lichen species at multiple locations on rock outcrops suggested that each lichen species harbored a distinct community and that all communities were dominated by Alphaproteobacteria. Across all samples, we recovered numerous bacterial phylotypes that were closely related to sequences isolated from lichens in prior investigations, including those from a lichen-associated Rhizobiales lineage (LAR1; putative N(2) fixers). LAR1-related phylotypes were relatively abundant and were found in all four lichen species, and many sequences closely related to other known N(2) fixers (e.g., Azospirillum, Bradyrhizobium, and Frankia) were recovered. Our findings confirm the presence of highly structured bacterial communities within lichens and provide additional evidence that these bacteria may serve distinct functional roles within lichen symbioses.     

Twenty-seven modes of reproduction in the obligate lichen symbiosis

Fungi exhibit some of the greatest reproductive diversity across Eukaryotes. In addition to sexual and asexual reproduction, fungi engage in parasexual (mitotic recombinatorial) processes to acquire new genetic variation. Reproduction has been studied extensively in numerous free-living fungi but comparatively less knowledge exists for lichenized fungi, which are assumed to reproduce only through sexual spores, asexual conidia, and specialized asexual lichen propagules. We present a new conceptual framework describing reproductive modes in lichens that includes sexual and asexual processes as well as accommodating the possibility of parasexual reproduction. To support the plausibility of some of these modes of reproduction, we reviewed data spanning more than 200 years of anatomical investigation. We recovered evidence supporting the possibility of 22 of 27 possible modes of reproduction and found no counter-evidence to suggest the remaining five do not occur in nature. This conceptual framework allows for a greater plurality of reproductive processes than previously acknowledged in lichens, exceeding that of their non-lichenized relatives.     

Compatibility and thigmotropism in the lichen symbiosis: A reappraisal

The development of many complex stratified lichen thalli is made through stages of complex phenotypic interactions between a filamentous fungus (the mycobiont), and a trebouxioid alga (the photobiont). Typically, the second stage of this symbiotic development is marked by the envelopment of the photobiont by the mycobiont through increased lateral hyphal branching and the formation of appressoria. Previously, the mycobiont’s envelopment of photobiont cells was considered thigmotropic (a growth response due to shape) as a mycobiont can envelop algal sized objects in its environment. However, after growing the mycobiontCladonia grayi with various phototrophs and glass beads, we conclude that the mycobiont does not show this characteristic second stage morphological response when grown in non-compatible pairings. Instead,C. grayi displays a distinctive morphological growth response only in compatible symbiotic pairings, such as with its natural photobiontAsterochlor’is sp.     

Novel lectins in the endoplasmic reticulum of wheat germ and their possible role

Novel lectins were isolated from the endoplasmic reticulum fractionof wheat germ by affinity chromatography. Their binding specificitywas similar to that of concanavalin A. However, the lectinsdid not cross-react with the antiserum against concanavalinA. Gel filtration indicated that there were at least two kindsof lectin molecules, one with molecular weight about 4,300 daltonsand the other with more than 300,000 daltons. By the differencein their binding specificity and molecular weights, the lectinsare distinguished from wheat germ agglutinin. Their possiblerole in the binding of ribosomes to endoplasmic reticulum isdiscussed. ¹ This work was reported at the Annual Meeting of the JapaneseSociety of Plant Physiologists (1978)(18). (Received April 10, 1978; )     

GalNAc/Gal-specific rat liver lectins: their role in cellular recognition

By investigating the presence and distribution of GalNAc/Gal-specific receptors on liver cells in vitro and in vivo, we provided evidence that the hepatocyte is not the only liver cell expressing receptor activity but that receptors of similar specificity are found on liver macrophages and also on endothelial cells. The receptor distribution in the plasma membrane is strinkingly different between the three cell types, as judged from the binding pattern of colloidal gold particles coated with asialofetuin or lactosylated serum albumin. Binding to hepatocytes occurs as single particles statistically distributed, binding to liver macrophages in a clustered arrangement all over the cell membrane and binding to endothelial cells also in a clustered arrangement but restricted to coated pits only. The different receptor distribution results in different binding and uptake abilities. Whereas hepatocytes bind and take up molecules and small particles (5 nm) only, the clustered receptor arrangement of endothelial cells and macrophages enables them to effectively bind and ingest larger particles. Ligands larger than 35 nm can be taken up by the macrophages only. The different receptor arrangement results also in different capacities of cell contact formation. Although in vitro liver macrophages and hepatocytes can both bind desialylated cells the macrophage needs much less galactosyl groups exposed on erythrocytes to establish stable contacts than the hepatocyte. The contacts formed by hepatocytes stay reversible for 30 min at 37 degrees C, whereas the contacts formed by the liver macrophages become irreversible after 10 min at 37 degrees C.(ABSTRACT TRUNCATED AT 250 WORDS)     

Vertical and horizontal photobiont transmission within populations of a lichen symbiosis

Lichens are widespread symbioses and play important roles in many terrestrial ecosystems. The genetic structure of lichens is the result of the association between fungal and algal populations constituting the lichen thallus. Using eight fungus- and seven alga-specific highly variable microsatellite markers on within-population spatial genetic data from 62 replicate populations across Europe, North America, Asia and Africa, we investigated the contributions of vertical and horizontal transmission of the photobiont to the genetic structure of the epiphytic lichen Lobaria pulmonaria. Based on pairwise comparisons of multilocus genotypes defined separately for the mycobiont and for the photobiont, we inferred the transmission mode of the photobiont and the relative contribution of somatic mutation and recombination. After constraining the analysis of one symbiont to pairs of individuals with genetically identical symbiotic partners, we found that 77% of fungal and 70% of algal pairs were represented by clones. Thus, the predominant dispersal mode was by means of symbiotic vegetative propagules (vertical transmission), which dispersed fungal and algal clones co-dependently over a short distance, thus shaping the spatial genetic structure up to distances of 20m. Evidence for somatic mutation generating genetic diversity was found in both symbionts, accounting for 30% of pairwise comparisons in the alga and 15% in the fungus. While the alga did not show statistically significant evidence of recombination, recombination accounted for 7.7% of fungal pairs with identical algae. This implies that, even in a mostly vegetatively reproducing species, horizontal transmission plays a role in shaping the symbiotic association, as shown in many coral and other symbioses in nature.     

Plant lectins: the ties that bind in root symbiosis and plant defense

Lectins are a diverse group of carbohydrate-binding proteins that are found within and associated with organisms from all kingdoms of life. Several different classes of plant lectins serve a diverse array of functions. The most prominent of these include participation in plant defense against predators and pathogens and involvement in symbiotic interactions between host plants and symbiotic microbes, including mycorrhizal fungi and nitrogen-fixing rhizobia. Extensive biological, biochemical, and molecular studies have shed light on the functions of plant lectins, and a plethora of uncharacterized lectin genes are being revealed at the genomic scale, suggesting unexplored and novel diversity in plant lectin structure and function. Integration of the results from these different types of research is beginning to yield a more detailed understanding of the function of lectins in symbiosis, defense, and plant biology in general.     

Microbial consortia of bacteria and fungi with focus on the lichen symbiosis

The investigation of fungal–bacterial interactions is an emerging field of research applying tools of modern microbial ecology. Studies have previously focused on the mycorrhizosphere, but in past decade, the role of bacteria in other fungal niches has been increasingly evaluated. This review presents recent progress in the understanding of fungal–bacterial interactions and contains a special focus on lichen symbioses. Lichens are traditionally considered as mutualisms between fungi and photoautotrophic species, but recent molecular approaches have revealed that lichens also harbour diverse microbial communities. Using modern DNA/RNA-based and microscopic techniques (e.g. FISH and confocal laser scanning microscopy) we are now able to analyse the abundance, composition, and structure of microbial communities in the lichen holobiont. Lichen-associated microbial communities consist of diverse taxonomic groups; the majority of bacteria belong to Alphaproteobacteria. Microbial communities can form biofilm-like structures on specific parts of the lichen thallus. Until now, the function and interaction within the microbial consortia is not fully understood. The functions displayed mainly by culturable strains suggest that bacteria have lytic activities, complement the nitrogen budget and produce bioactive substances, including hormones and antibiotics. Bacterial contribution to the lichen symbiosis is perhaps not restricted to one particular function in the lichen system, but supports a complex functional network which remains to be studied in greater detail.     

The role of lectins in plant defence

Summary Recent progress in the search for the physiological role of plant lectins supports the idea that some of these proteins are involved in the defence mechanisms of the plant. To place the evidence in favour of such a defensive role in a broad perspective, a short overview is given of the most important plant pathogens and predators. In addition, the solutions that plants have developed to resist the continuous threat of a hostile environment are briefly discussed in relation to the protective role of proteins in general. The presumed involvement of plant lectins in defence mechanisms is first inferred from an analysis of the biochemical, physiological, cellular biological and molecular biological properties of plant lectins. Subsequently, the available experimental evidence for the involvement of lectins in the plant's defence against viruses, bacteria, fungi and herbivorous invertebrates and vertebrates is discussed in some detail. Since the defensive role of plant lectins is determined largely by their ability to recognize and bind foreign glycans, a brief discussion is given of how the basically protective properties of these proteins can be exploited for histochemical applications in biological and biomedical research.     

Possible role of lectins in mycoparasitism.

Lectin activity in a host-mycoparasite relationship was demonstrated with Rhizoctonia solani and Trichoderma harzianum. Attachment of O but not A and B erythrocytes to hyphae occurred on R. solani but not on its mycoparasite. This phenomenon, which was Ca2+ and Mn2+ dependent, was prevented by galactose, present in T. harzianum cell walls, and by fucose.     

Cyanobacterial lichen symbiosis: the fungal partner as an optimal harvester

Lichen symbiosis has been traditionally treated as a model case of mutualism in which both partners, the fungus and the photobiont, gain benefits reciprocally. Some recent evidence, however, supports an alternative view that lichen symbiosis may represent an association largely controlled by the commensal or even parasitic fungal partner. The latter gains photosynthates from the photobiont (algae and/or cyanobacteria) which may not always substantially benefit from the symbiosis. We analyze from this perspective how a lichen fungus may maximize photosynthetic gains in bipartite and tripartite associations. We treat the frequency of nitrogen-fixing cells called heterocysts in cyanobacteria and the relative proportion of green algal cells vs. that of cyanobacteria per unit fungus as the variables to be manipulated for maximal carbon gain. The model predicts that even with a negligible cost of cephalodia (compartments containing cyanobacteria) it is in the interest of the tripartite lichen, first, to increase the heterocyst frequency, and second, keep the relative number of cyanobacteria considerably lower than that of green algae. Hence, the lichen fungus achieves higher fitness by making the cyanobacterial partner to specialize on N fixation. The available empirical data support these predictions as the reported heterocyst frequencies in bipartite lichens range from 2 to 8%, and in tripartite lichens between 10 and 55%. It is concluded that interaction asymmetry (i.e. commensalism or parasitism rather than mutualism) provides a sound basis to understand the high phenotypic plasticity expressed by fungi-forming bipartite and tripartite associations with cyanobacteria and green algae.     

Cell surface carbohydrates of symbiotic dinoflagellates and their role in the establishment of cnidarian–dinoflagellate symbiosis

Symbiodiniaceae algae are often photosymbionts of reef-building corals. The establishment of their symbiosis resembles a microbial infection where eukaryotic pattern recognition receptors (e.g. lectins) are thought to recognize a specific range of taxon-specific microbial-associated molecular patterns (e.g. glycans). The present study used the sea anemone, Exaiptasia diaphana and three species of Symbiodiniaceae (the homologous Breviolum minutum, the heterologous-compatible Cladocopium goreaui and the heterologous-incompatible Fugacium kawagutii) to compare the surface glycomes of three symbionts and explore the role of glycan–lectin interactions in host–symbiont recognition and establishment of symbiosis. We identified the nucleotide sugars of the algal cells, then examined glycans on the cell wall of the three symbiont species with monosaccharide analysis, lectin array technology and fluorescence microscopy of the algal cell decorated with fluorescently tagged lectins. Armed with this inventory of possible glycan moieties, we then assayed the ability of the three Symbiodiniaceae to colonize aposymbiotic E. diaphana after modifying the surface of one of the two partners. The Symbiodiniaceae cell-surface glycome varies among algal species. Trypsin treatment of the alga changed the rate of B. minutum and C. goreaui uptake, suggesting that a protein-based moiety is an essential part of compatible symbiont recognition. Our data strongly support the importance of D-galactose (in particular β-D-galactose) residues in the establishment of the cnidarian–dinoflagellate symbiosis, and we propose a potential involvement of L-fucose, D-xylose and D-galacturonic acid in the early steps of this mutualism.Subject terms: Cellular microbiology, Molecular biology