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Genetics, Vol. 157, 1169-1177, March 2001, Copyright © 2001

Regulatory Mechanisms Controlling Expression of the DAN/TIR Mannoprotein Genes During Anaerobic Remodeling of the Cell Wall in Saccharomyces cerevisiae

Natalia E. Abramovaa, Brian D. Cohenb, Odeniel Sertila, Rachna Kapoora, Kelvin J. A. Daviesc, and Charles V. Lowrya
a Center for Immunology and Microbial Disease, Albany Medical College, Albany, New York 12203,
b Laboratory of Reproductive & Metabolic Disorders, Wadsworth Center, Albany, New York 12201-2002
c The Andrus Gerontology Center, University of Southern California, Los Angeles, California 90089-0191

Corresponding author: Charles V. Lowry, Center for Immunology and Microbial Disease, Albany Medical College MC-151, 47 New Scotland Ave., Albany, NY 12203., cvlowry{at}aol.com (E-mail)

Communicating editor: M. JOHNSTON

The DAN/TIR genes of Saccharomyces cerevisiae encode homologous mannoproteins, some of which are essential for anaerobic growth. Expression of these genes is induced during anaerobiosis and in some cases during cold shock. We show that several heme-responsive mechanisms combine to regulate DAN/TIR gene expression. The first mechanism employs two repression factors, Mox1 and Mox2, and an activation factor, Mox4 (for mannoprotein regulation by oxygen). The genes encoding these proteins were identified by selecting for recessive mutants with altered regulation of a dan1::ura3 fusion. MOX4 is identical to UPC2, encoding a binucleate zinc cluster protein controlling expression of an anaerobic sterol transport system. Mox4/Upc2 is required for expression of all the DAN/TIR genes. It appears to act through a consensus sequence termed the AR1 site, as does Mox2. The noninducible mox4{Delta} allele was epistatic to the constitutive mox1 and mox2 mutations, suggesting that Mox1 and Mox2 modulate activation by Mox4 in a heme-dependent fashion. Mutations in a putative repression domain in Mox4 caused constitutive expression of the DAN/TIR genes, indicating a role for this domain in heme repression. MOX4 expression is induced both in anaerobic and cold-shocked cells, so heme may also regulate DAN/TIR expression through inhibition of expression of MOX4. Indeed, ectopic expression of MOX4 in aerobic cells resulted in partially constitutive expression of DAN1. Heme also regulates expression of some of the DAN/TIR genes through the Rox7 repressor, which also controls expression of the hypoxic gene ANB1. In addition Rox1, another heme-responsive repressor, and the global repressors Tup1 and Ssn6 are also required for full aerobic repression of these genes.




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