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The transcriptional regulator  HAP4 , induced by respiratory substrates, is involved in the balance between fermentation and respiration in  S. cerevisiae . We identified putative orthologues of the Hap4 protein in all ascomycetes, based only on a conserved sixteen amino acid-long motif. In addition to this motif, some of these proteins contain a DNA-binding motif of the bZIP type, while being nonetheless globally highly divergent. The genome of the yeast  Hansenula polymorpha  contains two  HAP4 -like genes encoding the protein HpHap4-A which, like ScHap4, is devoid of a bZIP motif, and HpHap4-B which contains it. This species has been chosen for a detailed examination of their respective properties. Based mostly on global gene expression studies performed in the  S. cerevisiae HAP4  disruption mutant ( ScΔhap4 ), we show here that HpHap4-A is functionally equivalent to ScHap4, whereas HpHap4-B is not. Moreover  HpHAP4-B  is able to complement the H 2 O 2  hypersensitivity of the  ScYap1  deletant,  YAP1  being, in  S. cerevisiae , the main regulator of oxidative stress. Finally, a transcriptomic analysis performed in the  ScΔyap1  strain overexpressing  HpHAP4-B  shows that HpHap4-B acts both on oxidative stress response and carbohydrate metabolism in a manner different from both ScYap1 and ScHap4. Deletion of these two genes in their natural host,  H. polymorpha , confirms that  HpHAP4-A  participates in the control of the fermentation/respiration balance, while  HpHAP4-B  is involved in oxidative stress since its deletion leads to hypersensitivity to H 2 O 2 . These data, placed in an evolutionary context, raise new questions concerning the evolution of the  HAP4  transcriptional regulation function and suggest that Yap1 and Hap4 have diverged from a unique regulatory protein in the fungal ancestor.

plos one

Functional Study of the Hap4-Like Genes Suggests That the Key Regulators of Carbon Metabolism HAP4 and Oxidative Stress Response YAP1 in Yeast Diverged from a Common Ancestor