Characterization of the Cytochrome c Oxidase Assembly Factor Cox19 of Saccharomyces cerevisiae

Cox19 is an important accessory protein in the assembly of cytochrome c oxidase in yeast. The protein is functional when tethered to the mitochondrial inner membrane, suggesting its functional role within the intermembrane space. Cox19 resembles Cox17 in having a twin CX9C sequence motif that adopts a helical hairpin in Cox17. The function of Cox17 appears to be a Cu(I) donor protein in the assembly of the copper centers in cytochrome c oxidase. Cox19 also resembles Cox17 in its ability to coordinate Cu(I). Recombinant Cox19 binds 1 mol eq of Cu(I) per monomer and exists as a dimeric protein. Cox19 isolated the mitochondrial intermembrane of suggesting that Cu(I) be a important Cu(I) also to be important the in function of a exists in the ability to Cu(I) and in Cox19 is an important accessory protein in the assembly of cytochrome c oxidase in yeast. The protein is functional when tethered to the mitochondrial inner membrane, suggesting its functional role within the intermembrane space. Cox19 resembles Cox17 in having a twin CX9C sequence motif that adopts a helical hairpin in Cox17. The function of Cox17 appears to be a Cu(I) donor protein in the assembly of the copper centers in cytochrome c oxidase. Cox19 also resembles Cox17 in its ability to coordinate Cu(I). Recombinant Cox19 binds 1 mol eq of Cu(I) per monomer and exists as a dimeric protein. Cox19 isolated the mitochondrial intermembrane of suggesting that Cu(I) be a important Cu(I) also to be important the in function of a exists in the ability to Cu(I) and in c oxidase cytochrome c inner intermembrane cytochrome c inner intermembrane is the of the in of and The the of and the inner of to the of the the mitochondrial The in a and copper is within the The copper in the within of a of the assembly of and mitochondrial of to and the and of that of copper the of accessory and to be important in the copper of and in the the accessory within and the inner membrane, a the and the copper of the and in and and of a of a twin CX9C motif that adopts a helical hairpin in Cox17 and the of the that the in and is a copper within the as the donor of Cu(I) to and in Cox17 a Cox17 appears to to Cu(I) to protein. Cox17 in Cu(I) of a Cu(I) to a protein of a Cu(I) in Cox17 of the that of is to a to Cu(I) Cu(I) is an protein a The protein is in a the the The that be in the of in a sequence in function of Cox17 the is suggesting that Cox17 is functional of the in the twin CX9C The copper is and the a the The of is Cox17 in a within the resembles Cox17 in a protein within the mitochondrial the protein is also in the when Cox19 and the of is the of copper The of Cox19 Cox17 that Cox19 also function in the copper of Cox19 is to Cox17 in its ability to coordinate Cu(I). important in Cox19 function important in Cu(I) and an to of and a as in in and and and to of a the of the and the and The a the the and sequence a a as a as to the and in the the and in the Cox19 the of to the and as Cox17 The and the mitochondrial sequence and of to and Cox19 the and and a The to a as as the The sequence the a the sequence and a as a The and the to the of and an of to the to a of 1 to and in in the 1 a in 1 The of of and of protein of 1 The protein to the in a and a and of Cox19 isolated per of the a protein to in and 1 a a and in 1 The protein the to and as a in in a of as The of Cu(I) in of a in the of to the of in in 1 in and to a of and a of to and and the the a of and an is a of the and a of the the of the to The and and as the copper an of a of an and the to the of a copper a of the copper to be the of and the the to the of and a 1 and an of copper protein the and and copper protein also of copper protein and as the and copper in a an The Cox19 protein in the of 1 and the in the and of a of isolated to the in the of 1 protein in isolated of of cytochrome c in of mitochondrial protein a and a the and a and to the mitochondrial and Cox19 in as a protein an to the of the protein in the of Cox19 mol eq of monomer and the protein as of Cox19 in the of the protein of The of copper to the is also the Cox17 and the of copper in to Cox19 is to in the and of 1 The of the copper is the Cu(I) suggesting that Cox19 binds Cu(I) to Cox17. The as as as isolated per suggesting that in the Cox19 The of in a the is in the of the the copper mol and of the copper as The is to and the copper is to 1 mol of in in the and as in The is the Cu(I) of the protein a to The of to the an of a Cu(I) of of an a of in The of a Cu(I) to the of the protein in of the a as of in as 1 mol eq of Cu(I) per monomer The of Cu(I) in of 1 mol eq in in the Cu(I) 1 mol eq of Cu(I) and an of a 1 mol eq of Cu(I) in copper is in and the of Cu(I) is to the of to the of of the is 1 eq of the protein and of and as a dimeric The dimeric of and of and and of of a copper in The of Cox19 as a The copper the that copper is to the protein. of a The that of the The to the the of protein as a of also the protein a of is the and of the protein. and protein to the in The copper of a of a Cu(I) coordinate Cu(I) and the of Cu(I) in is The a Cu(I) and a Cu(I) also in The and of the in to the to in the suggesting Cu(I) in The in Cox19 to of of the copper of the and the and of and Cox17. the in copper and in and in the of The is of Cu(I) and Cu(I) to and to of the The is copper the of a in of in a is in the in of the and of and is in and The of the in Cox19 is The is copper and is that to a that of copper in a of copper is the of Cu(I) per copper in the a the to Cox19 be isolated as a protein when the protein is in The protein isolated as a dimeric a per monomer and the protein as the of Cox19 in to Cox19 the of copper mol mol the The of the to The of the also to be a in the of The protein suggesting that the protein in the within the Cox19 and to in the of is in Cox19 functional in The and Cox19 within the of that Cox19 of in of of Cox19 a of Cox19 and and tethered and of the of and and isolated the an and as mitochondrial and that Cox17 also functional when tethered to the mitochondrial the Cox19 protein a of its mitochondrial sequence and The Cu(I) of that the the Cox19 the Cox19 to the Cox19 of of The Cox19 also within the and the suggesting that the protein functional Cox19 Cox17 mitochondrial of and Cox19 and Cox17 also mitochondrial suggesting that the function in mitochondrial Cox19 in the twin CX9C motif The in and in the helical hairpin in The of the Cu(I) in Cox17 is function the in Cox19 to and and of Cox19 and and of of and of isolated the in a and a a The of Cox19 is the is the the is to the important is a sequence motif of is a Cu(I) a of Cox19 in function the Cox19 to a is of Cox19 in the and as as the that to in Cox19 the mitochondrial is the protein The of the and the the and in to the the protein of that of the Cox19 the in that to the of a protein an Cox19 when as an Cox19 The protein and Cox19 protein in of Cox19 and Cox19 of mitochondrial and and as mitochondrial and Cox19 copper The Cox19 as as the and in Cu(I) The the in Cu(I) Cu(I) in the copper also of as as and Cox19 a twin CX9C sequence Cox19 is functional when tethered to the mitochondrial that functional role is within the mitochondrial of and important function in the assembly of important in Cu(I) an ability to to dimeric and Cox19 exists in a Cox17 also as a of be of twin CX9C to be functional of Cox17 and Cox19 the is in a Cox19 to within the when the and Cox19 to be within the of Cox19 and Cox17 within the in the and Cox19 an ability to Cu(I) in Cox17 the in the twin CX9C Cox19 the motif Cu(I) in Cox19 motif the and important in a of the in a functional of the is important a Cox17 is a protein is The Cox19 to the Cox17 is the Cox19 that is in its in The Cox19 is also in Cu(I) the is in in function and Cu(I) a exists in function and Cu(I) within the motif appears to in function and Cu(I) be to be a Cu(I) of the protein that be to and be to to in the of the The of is a Cu(I) a role in the Cu(I) a of important to the the copper of The and of copper and in the and to the of the The the of a Cu(I) a copper in the The the of in the and is The Cu(I) and Cox19 is an the Cu(I) be its dimeric of the copper in Cox19 the Cu(I) in monomer is in to the Cu(I) in the a The in and Cox19 dimeric Cu(I) within a of to the of the a the of the copper the of the to the of the Cu(I) protein of that Cu(I) to a dimeric Cox19 is to Cox19 the in the protein of and The protein suggesting that the protein in the within the of Cox17 the protein of copper mol Cox19 a copper role to Cu(I) be the of the copper is in of Cox17 and Cox19 is that within a of that is the of the The that to of the and that an of the the Cox19 a to that of a of that also a twin CX9C The of is The of in the is and that Cox17 is in Cu(I) to and of and in and Cox17 Cu(I) to and the the role of is that Cox17 and Cox19 to the is a copper is that is of that is to Cu(I) and that the a a Cu(I) and to the that Cox17 Cu(I) to Cox19 is to the in c oxidase cytochrome c inner intermembrane cytochrome c inner intermembrane is the of the in of and The the of and the inner of to the of the the mitochondrial The in a and copper is within the The copper in the within of a of the assembly of and mitochondrial of to and the and of that of copper the of accessory and to be important in the copper of and in the the accessory within and the inner membrane, a the and the copper of the and in and and of a of a twin CX9C motif that adopts a helical hairpin in Cox17 and the of the that the in and Cox17 is a copper within the as the donor of Cu(I) to and in Cox17 a Cox17 appears to to Cu(I) to protein. Cox17 in Cu(I) of a Cu(I) to a protein of a Cu(I) in Cox17 of the that of is to a to Cu(I) Cu(I) is an protein a The protein is in a the the The that be in the of in a sequence in function of Cox17 the is suggesting that Cox17 is functional of the in the twin CX9C The copper is and the a the The of is Cox17 in a within the Cox19 resembles Cox17 in a protein within the mitochondrial the protein is also in the when Cox19 and the of is the of copper The of Cox19 Cox17 that Cox19 also function in the copper of Cox19 is to Cox17 in its ability to coordinate Cu(I). important in Cox19 function important in Cu(I) and an to of and a as in in and and and to of a the of the and the and The a the the and sequence a a as a as to the and in the the and in the Cox19 the of to the and as Cox17 The and the mitochondrial sequence and of to and Cox19 the and and a The to a as as the The sequence the a the sequence and a as a The and the to the of and an of to the to a of 1 to and in in the 1 a in 1 The of of and of protein of 1 The protein to the in a and a and of Cox19 isolated per of the a protein to in and 1 a a and in 1 The protein the to and as a in in a of as The of Cu(I) in of a in the of to the of in in 1 in and to a of and a of to and and the the a of and an is a of the and a of the the of the to The and and as the copper an of a of an and the to the of a copper a of the copper to be the of and the the to the of and a 1 and an of copper protein the and and copper protein also of copper protein and as the and copper in a an The Cox19 protein in the of 1 and the in the and of a of isolated to the in the of 1 protein in isolated of of cytochrome c in of mitochondrial protein a and a the and a and to the mitochondrial and and an to of and a as in in and and and to of a the of the and the and The a the the and sequence a a as a as to the and in the the and in the The Cox19 the of to the and as Cox17 The and the mitochondrial sequence and of to and Cox19 the and and a The to a as as the The sequence the a the sequence and a as a The and the to the of and an of to the to a of 1 to and in in the 1 a in 1 The of of and of protein of 1 The protein to the in a and a and of Cox19 isolated per of the a protein to in and 1 a a and in 1 The protein the to and as a in in a of as The of Cu(I) in of a in the of to the of in in 1 in and to a of and a of to and and the the a of and an is a of the and a of the the of the to The and and as the copper an of a of an and the to the of a copper a of the copper to be The the of and the the to the of and a 1 and an of copper protein the and and copper protein also of copper protein and as the and copper in a an The Cox19 protein in the of 1 and the in the and of a of isolated to the in the of 1 protein in isolated of of cytochrome c in of mitochondrial protein a and a the and a and to the mitochondrial and Cox19 in as a protein an to the of the protein in the of Cox19 mol eq of monomer and the protein as of Cox19 in the of the protein of The of copper to the is also the Cox17 and the of copper in to Cox19 is to in the and of 1 The of the copper is the Cu(I) suggesting that Cox19 binds Cu(I) to Cox17. The as as as isolated per suggesting that in the Cox19 The of in a the is in the of the the copper mol and of the copper as The is to and the copper is to 1 mol of in in the and as in The is the Cu(I) of the protein in of the a as of in as 1 mol eq of Cu(I) per monomer The of Cu(I) in of 1 mol eq in in the Cu(I) 1 mol eq of Cu(I) and an of a 1 mol eq of Cu(I) in copper is in and the of Cu(I) is to the of to the of of the is 1 eq of the protein and of and as a dimeric The dimeric of and of and and of of a copper in The of Cox19 as a The copper the that copper is to the protein. of a The that of the The to the the of protein as a of also the protein a of is the and of the protein. and protein to the in The copper of a of a Cu(I) coordinate Cu(I) and the of Cu(I) in is The a Cu(I) and a Cu(I) also in The and of the in to the to in the suggesting Cu(I) in The in Cox19 to of of the copper of the and the and of and Cox17. the in copper and in and in the of The is of Cu(I) and Cu(I) to and to of the The is copper the of a in of in a is in the in of the and of and is in and The of the in Cox19 is The is copper and is that to a that of copper in a of copper is the of Cu(I) per copper in the a the to Cox19 be isolated as a protein when the protein is in The protein isolated as a dimeric a per monomer and the protein as the of Cox19 in to Cox19 the of copper mol mol the The of the to The of the also to be a in the of The protein suggesting that the protein in the within the Cox19 and to in the of is in Cox19 functional in The and Cox19 within the of that Cox19 of in of of Cox19 a of Cox19 and and tethered and of the of and and isolated the an and as mitochondrial and that Cox17 also functional when tethered to the mitochondrial the Cox19 protein a of its mitochondrial sequence and The Cu(I) of that the the Cox19 the Cox19 to the Cox19 of of The Cox19 also within the and the suggesting that the protein functional Cox19 Cox17 mitochondrial of and Cox19 and Cox17 also mitochondrial suggesting that the function in mitochondrial Cox19 in the twin CX9C motif The in and in the helical hairpin in The of the Cu(I) in Cox17 is function the in Cox19 to and and of Cox19 and and of of and of isolated the in a and a a The of Cox19 is the is the the is to the important is a sequence motif of is a Cu(I) a of Cox19 in function the Cox19 to a is of Cox19 in the and as as the that to in Cox19 the mitochondrial is the protein The of the and the the and in to the the protein of that of the Cox19 the in that to the of a protein an Cox19 when as an Cox19 The protein and Cox19 protein in of Cox19 and Cox19 of mitochondrial and and as mitochondrial and Cox19 copper The Cox19 as as the and in Cu(I) The the in Cu(I) Cu(I) in the copper also of as as Recombinant Cox19 in as a protein an to the of the protein in the of Cox19 mol eq of monomer and the protein as of Cox19 in the of the protein of The of copper to the is also the Cox17 and the of copper in to Cox19 is to in the and of 1 The of the copper is the Cu(I) suggesting that Cox19 binds Cu(I) to Cox17. The as as as isolated per suggesting that in the Cox19 The of in a the is in the of the the copper mol and of the copper as The is to and the copper is to 1 mol of in in the and as in The is the Cu(I) of the protein in of the a as of in as 1 mol eq of Cu(I) per monomer The of Cu(I) in of 1 mol eq in in the Cu(I) 1 mol eq of Cu(I) and an of a 1 mol eq of Cu(I) in copper is in and the of Cu(I) is to the of to the and as a dimeric The dimeric of and of and and to the in The copper of a of a Cu(I) coordinate Cu(I) and the of Cu(I) in is The a Cu(I) and a Cu(I) also in The and of the in to the to in the suggesting Cu(I) in The in Cox19 to The copper and in and in the of The is of Cu(I) and Cu(I) to and to of the The is copper the of a in is in the in of the and of and is in and The of the in Cox19 is The is copper and is that to a that of copper in a of copper is the of Cu(I) per copper in the a the to Recombinant Cox19 be isolated as a protein when the protein is in The protein isolated as a dimeric a per monomer and the protein as the of Cox19 in to Cox19 the of copper mol mol the The of the to The of the also to be a in the of The protein suggesting that the protein in the within the Cox19 and to in the of is in Cox19 functional in The and Cox19 within the of that Cox19 of in of of Cox19 a that Cox17 also functional when tethered to the mitochondrial the Cox19 protein a of its mitochondrial sequence and The Cu(I) of that the the Cox19 the Cox19 to the Cox19 of of The Cox19 also within the and the suggesting that the protein functional Cox19 Cox17 mitochondrial of and Cox19 and Cox17 also mitochondrial suggesting that the function in mitochondrial Cox19 in the twin CX9C motif The in and in the helical hairpin in The of the Cu(I) in Cox17 is function the in Cox19 to and and and a a The of Cox19 is the is the the is to the important is a sequence motif of is a Cu(I) a of Cox19 in function the Cox19 to a is The of Cox19 in the and as as the that to in Cox19 the mitochondrial is the protein The of the and the the and in to the the protein of that of the Cox19 the in The that to the of a protein an Cox19 when as an Cox19 The protein and The Cox19 copper The Cox19 as as the and in Cu(I) The the in Cu(I) Cu(I) in the copper also of as as and Cox19 a twin CX9C sequence Cox19 is functional when tethered to the mitochondrial that functional role is within the mitochondrial of and important function in the assembly of important in Cu(I) an ability to to dimeric and Cox19 exists in a Cox17 also as a of be of twin CX9C to be functional of Cox17 and Cox19 the is in a Cox19 to within the when the and Cox19 to be within the of Cox19 and Cox17 within the in the and Cox19 an ability to Cu(I) in Cox17 the in the twin CX9C Cox19 the motif Cu(I) in Cox19 motif the and important in a of the in a functional of the is important a Cox17 is a protein is The Cox19 to the Cox17 is the Cox19 that is in its in The Cox19 is also in Cu(I) the is in in function and Cu(I) a exists in function and Cu(I) within the motif appears to in function and Cu(I) be to be a Cu(I) of the protein that be to and be to to in the of the The of is a Cu(I) a role in the Cu(I) a of important to the the copper of The and of copper and in the and to the of the The the of a Cu(I) a copper in the The the of in the and is The Cu(I) and Cox19 is an the Cu(I) be its dimeric of the copper in Cox19 the Cu(I) in monomer is in to the Cu(I) in the a The in and Cox19 dimeric Cu(I) within a of to the of the a the of the copper the of the to the of the Cu(I) protein of that Cu(I) to a dimeric Cox19 is to Cox19 the in the protein of and The protein suggesting that the protein in the within the of Cox17 the protein of copper mol Cox19 a copper role to Cu(I) be the of the copper is in of Cox17 and Cox19 is that within a of that is the of the The that to of the and that an of the the Cox19 a to that of a of that also a twin CX9C The of is The of in the is and that Cox17 is in Cu(I) to and of and in and Cox17 Cu(I) to and the the role of is that Cox17 and Cox19 to the is a copper is that is of that is to Cu(I) and that the a a Cu(I) and to the that Cox17 Cu(I) to Cox19 is to the in Cox17 and Cox19 a twin CX9C sequence Cox19 is functional when tethered to the mitochondrial that functional role is within the mitochondrial of and important function in the assembly of important in Cu(I) an ability to to dimeric and Cox19 exists in a Cox17 also as a of be The of twin CX9C to be functional of Cox17 and Cox19 the is in a Cox19 to within the when the and Cox19 to be within the of Cox19 and Cox17 within the in the Cox17 and Cox19 an ability to Cu(I) in Cox17 the in the twin CX9C Cox19 the motif Cu(I) in Cox19 motif the and important in a of the in a functional of the is important a Cox17 is a protein is The Cox19 to the Cox17 is the Cox19 that is in its in The Cox19 is also in Cu(I) the is in in function and Cu(I) a exists in function and Cu(I) within the motif appears to in function and Cu(I) be to be a Cu(I) of the protein that be to and be to to in the of the The of is a Cu(I) a role in the Cu(I) a of important to the the copper of The and of copper and in the and to the of the The the of a Cu(I) a copper in the The the of in the and is The Cu(I) and Cox19 is an the Cu(I) be its dimeric The of the copper in Cox19 the Cu(I) in monomer is in to the Cu(I) in the a The in and Cox19 dimeric Cu(I) within a of to the of the a the of the copper the of the to the of the Cu(I) protein of that Cu(I) to a dimeric Cox19 is to Cox19 the in the protein of and The protein suggesting that the protein in the within the of Cox17 the protein of copper mol Cox19 a copper role to Cu(I) be the of the copper is in of Cox17 and Cox19 is that within a of that is the of the The that to of the and that an of the the Cox19 a to that of a of that also a twin CX9C The of is The of in the is and that Cox17 is in Cu(I) to and of and in and Cox17 Cu(I) to and the the role of is that Cox17 and Cox19 to the is a copper is that is of that is to Cu(I) and that the a a Cu(I) and to the that Cox17 Cu(I) to Cox19 is to the in the and of in

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