THE MORPHOGENESIS AND POSSIBLE EVOLUTIONARY ORIGINS OF FUNGAL SCLEROTIA

1. Fungal sclerotia are able to survive adverse conditions for long periods and they are formed by many important plant pathogens. An understanding of the factors involved in their initiation and development may lead to a method of repressing their formation in nature, thereby reducing the chances of survival of fungi that depend on them as persistent resting stages in their life-cycles. Also, data on sclerotial morphogenesis may be applicable to other multihyphal fungal structures. 2. There are three types of sclerotial development. The most primitive and least common is the loose type, which is illustrated by Rhizoctonia solani. The sclerotium forms by irregular branching of the mycelium followed by intercalary septation and hyphal swelling. When mature, it consists of loosely interwoven hyphae that are rich in food reserves and darkly pigmented. The main types of development are terminal and lateral. The former develops from the coalescence of initials that are produced by a well-defined pattern of branching at the tip of a hypha or tips of closely associated hyphae, e.g. Botrytis cinerea. Lateral sclerotia are formed by the interweaving of side branches of one or several main hyphae. When only one main hypha is involved the sclerotium is of the lateral, simple type, e.g. Sclerotinia gladioli. If several main hyphae give rise to a sclerotium, the term strand type has been used. Sclerotium rolfsii is the classical example. 3. There is a considerable literature on the effects of environmental conditions on the initiation, development and maturation of sclerotia but few attempts have been made to interpret the data. Phenolics and/or polyphenol oxidases have been found to be connected with morphogenesis of the protoperithecium of Neurospora crassa, the perithecium of Podospora anserina and of Hypomyces sp. and the basidiocarp of Schixophyllum commune. A close correlation has been shown between melanin synthesis and microsclerotial development by Verticillium but there appears to be no literature on the role of phenolics and polyphenol oxidases in the morphogenesis of sclerotia. Possibly these substances may inhibit growth of the apices of main hyphae by changing the permeability of the membrane, by inducing a thickening of the cell wall at the tip or by reducing the plasticity of the wall. Such a check in growth could trigger-off the formation of initials close to the margin of the colony or elsewhere in the culture. Sulphydryl groups and disulphide bonds are of great significance in morphogenesis of organisms and are probably involved in sclerotial initiation. The formation of a large number of hyphal branches is a prerequisite for sclerotial initiation and mycelial branching is possible only if there is plasticity of hyphal walls. The ability of the wall to be moulded is possibly related to changes in the sulphur linkages of the protein of the protein-carbohydrate complexes of the cell wall and could be influenced by sulphur availability or the activity of specific enzymes. 4. After a sclerotial primordium has been initiated, further increase in size will depend on the continued, active translocation of nutrients to the site of development. Movement of nutrients to sclerotia is through a few translocatory hyphae. Presumably, nutrients will continue to move into the young sclerotium as long as a concentration or pressure gradient is maintained. Energy and substances for the formation of new branches are supplied in this way and as the requirements for hyphal branches are reduced, excess nutrients become available for conversion to inactive or insoluble reserves and for exudation. The exudates are often complex, consisting of proteins, including enzymes, lipids and carbohydrates. Many sclerotia have a mucilaginous matrix in which the medullary hyphae are embedded. Sclerotium-forming, fungal species that are not regarded as having such a matrix appear to secrete a layer of mucilage over the surface of sclerotial hyphae. This mucilage could have a morphogenetic function and serve as an adhesive which loosely binds hyphae together. More permanent unions are by hyphal fusions or anastomoses. 5. The sclerotium matures within a few days of attaining its maximum size. The rind effectively seals off the medullary hyphae from the surroundings and the translocatory hyphae cease to function. Thus the sclerotium is isolated both physiologically and nutritionally. The endogenous reserves enable the structure to exist in the absence of exogenous nutrients and then, when conditions become suitable, to germinate. 6. The sclerotium appears to provide an example of convergent evolution whereby analogous structures, which have become adapted to resist adverse conditions, have evolved. Data are available mainly for Typhula spp. and ScZerotinia spp. Sclerotia may be degenerate sexual reproductive structures, hyphal aggregates that have developed from closely interwoven conidiophores and undifferentiated conidia or they may be modified vegetative structures.