Morphogenesis of coremia and rhizomorphs in the AscomyceteSphaerostilbe repens

Abstract

The differentiating stages of coremia and rhizomorphs inSphaerostilbe repens were studied by transmission electron microscopy. Vegetative mycelium is characterized by highly cytoplasmic cells rich in ribosomes and mitochondria and with few vacuoles as well as endoplasmic reticulum. Cell walls are thin attaining a maximum thickness of 0.10 μm. During the aggregating phase a prosenchymatous mass of randomly oriented cells is produced by localized elongation and branching of the filaments. The hyphae in this region have the appearance of actively metabolising cells. In the course of the differentiating phase, numerous hyphae of the median zone of the aggregate grow upward and downward to give rise to coremium and rhizomorph primordia respectively. The individual hyphal tips lay parallel to each other and cells of the growing apices retain their meristematic characteristics. At the periphery of the aggregate and to a lesser extent in the subapical rhizomorphic zone, cells reduce their cytoplasmic density as a consequence of a decrease in the number of ribosomes. These cells also increase in size and become isodiametric and vacuolated. During cellular differentiation walls increase steadily in thickness and at the elongating phase they reach 0.30 μm in the rhizomorphic cortex. Mucilaginous material is progressively deposited around hyphae and in the most differentiated zones, coalesce to fill interhyphal spaces. This extracellular matrix seems to play a role in maintaining cohesiveness of the aggregated organs. The tissue in the process of differentiation is scattered with cells highly enriched in mitochondria and with cells virtually undifferentiated. Accumulation of microfilaments takes place in the differentiating zone localized behind the immersed meristematic apex. These structures might be involved in wall synthesis. Glycogen rosettes accumulate in the vegetative mycelium surrounding the aggregating centers, suggesting the possibility of supplying energy during the differentiating processes. The vacuolar system, represented by autophagic vacuoles which are present until the differentiation phase, presumably may also participate in the biochemical changes that occur during aggregation. Coremial cells are characterized by an increase in wall thickness, a highly sinuous plasma-membrane as well as large amounts of mucilaginous compounds accumulated between hyphae, but in all other respects they resemble the cells of actively growing vegetative hyphae.