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Functional analysis of a GSP1/Ran ortholog,  CpRan1 , from  Cryphonectria parasitica  was conducted. Genotype analysis revealed that the putative  CpRan1 -null mutant was a heterokaryotic transformant harboring two different types of nuclei, one with the wild-type  CpRan1  allele and the other with the  CpRan1 -null mutant allele. The mycelial growth and colony morphology of the heterokaryotic transformant was normal. Microscopic analysis of the resulting conidia (aseptate and monokaryotic asexual spores) demonstrated that although normal germinating spores were observed from conidia harboring a nucleus with the wild-type  CpRan1  allele, a number of residual conidia that did not germinate existed. Complementation analysis using protoplasts from the heterokaryon with the wild-type  CpRan1  allele confirmed that the  CpRan1  gene is essential to  C. parasitica . Complementation analysis using the various  CpRan1  chimera constructs allowed us to perform a functional analysis of essential amino acids of the  CpRan1 . Among the four suggested essential amino acids, Lys-97 for ubiquitination was determined to not be an essential residue. Moreover, the  CpRan1 -null mutant allele was successfully complemented with mouse Ran gene, which suggested that the biological function of Ran gene is evolutionary conserved and that our heterokaryon rescue can be applied for the functional analysis of heterologous genes.

Functional Analysis of an Essential GSP1/Ran Ortholog Gene, CpRan1, from the Chestnut Blight Fungus Cryphonectria parasitica Using a Heterokaryon