4 Septation and Cytokinesis in Pathogenic Fungi

The control of cytokinesis and septation by pathogenic fungi is critical to the developmental changes associated with host invasion and fungal pathogenesis. Pathogenic processes, such as growth within host tissue, often require changes between yeast-like,

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Septation and Cytokinesis in Pathogenic Fungi

M´IRIAM OSE´S-RUIZ1, WASIN SAKULKOO1, NICHOLAS J. TALBOT1

CONTENTS I. Introduction to Septation and Cytokinesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . II. The Rice Blast Fungus Magnaporthe oryzae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . III. Septation and Cytokinesis in Magnaporthe oryzae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . IV. Septin Ring Formation at the Base of the Appressorium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. Dynamics of Septin Ring Formation in M. oryzae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. Cell Cycle Regulation of Septin Ring Formation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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I. Introduction to Septation and Cytokinesis Cell division in all organisms requires the coordination of all cellular processes to allow the formation of two new individual cells. During the eukaryotic cell cycle, correct duplication of genetic material and its subsequent even division, together with the contents of the cytoplasm, are necessary for successful generation of daughter cells. In order to ensure this outcome, the cell possesses cell cycle checkpoints at crucial stages to ensure fidelity and temporal coordination of cell division. In eukaryotes the mechanism that allows final coordination of cell division and compartmentalisation of new cellular components is called cytokinesis. In fungi, cytokinesis adopts different forms. In 1 Biosciences, College of Life and Environmental Sciences, University of Exeter, Geoffrey Pope Building, Stocker Road, Exeter EX4 4QD, UK; e-mail: [email protected]

Saccharomyces cerevisiae the process of cytokinesis is called budding, resulting in a new daughter cell from a bud formed by the mother, while in the other major model yeast species Schizosaccharomyces pombe, it is called fission and results in two daughter cells of equal size. In filamentous fungi, cytokinesis within a hypha is known as septation and results in discrete hyphal compartments within the multicellular fungal hyphal filament. The processes of budding and fission lead to separation of a daughter cell from its mother cell, allowing formation of new individuals. However, the equivalent process in filamentous fungi, septation, results in new hyphal compartments in which daughter cells remain attached to form a continuous, multicellular organism. This important principle means there are major mechanistic differences in cytokinesis among fungi. In both budding and fission yeast, for example, when a septum is being formed, septum-related proteins are recruited to the site creating a septum and establishing the plane of cell division. It is only after cytokinesis that the septum is degrad