Regulated Membrane Recruitment of Dynamin-2 Mediated by Sorting Nexin 9

The endocytic proteins sorting nexin 9 (SNX9) and dynamin-2 (Dyn2) assemble in the cytosol as a resting complex, together with a 41-kDa protein. We show here that the complex can be activated for membrane binding of SNX9 and of cytosol in the of SNX9 for the the that SNX9 as a of in The 41-kDa as the with a in SNX9 the membrane binding of of SNX9 the complex and SNX9 for membrane The that of the membrane SNX9 and The endocytic proteins sorting nexin 9 (SNX9) and dynamin-2 (Dyn2) assemble in the cytosol as a resting complex, together with a 41-kDa protein. We show here that the complex can be activated for membrane binding of SNX9 and of cytosol in the of SNX9 for the the that SNX9 as a of in The 41-kDa as the with a in SNX9 the membrane binding of of SNX9 the complex and SNX9 for membrane The that of the membrane SNX9 and The proteins in in the in as in and and The in the of the the of the the of the The for the of a the of a of proteins the and complex sorting nexin sorting nexin in the of the membrane and of a in a the of the and the the The be and in the a that in the of in in the of and the of the membrane for with in the the a the for a of proteins in the of the of that the of the be for the of the cytosol the membrane of the membrane of the in the in the dynamin-2 (Dyn2) in of the and that with as the in be for nexin 9 a of the sorting nexin of proteins a be in the endocytic as SNX9 binding for and in a and in the SNX9 membrane binding a a and a SNX9 with and the and in and of of SNX9 the of the cytosol of of SNX9 be in a complex together with and of The the of complex and the membrane of SNX9 and We show that SNX9 the of the and that the membrane binding of SNX9 a with the a with and the in and as in and for the in and for a as the cytosol with and cytosol in a of as The 41-kDa of the with and a the of as a of the complex and of binding of and with SNX9 with the of and of in the as proteins with the of the in SNX9 be for binding of in cytosol and with proteins with and and a of in and with in the of and of SNX9 as proteins as and The in the and as with proteins as and a for The and for in of The and in The as in a of of cytosol and and and and as in the with and cytosol the with the The in and the the a for the for with of and The in of in and the for as of in of of in cytosol with as in the for the for with of and for with cytosol with as in the of of in in and SNX9 with and the in the and the in in of SNX9 with the of cytosol for SNX9 with as in for in the of a of cytosol and in and The and the the cytosol with for in the of and and a and for the of SNX9 and for with cytosol in the of and with as the and proteins with cytosol and for the proteins a and with The for and cytosol together with and in the of and together with and The The for and SNX9 and of proteins binding for in of of and of in in and for with and as that and for the with and a the of and in in and of and of and cytosol and a a as and the of The and the a a of the The as a the membrane binding of SNX9 and with and of SNX9 and The in show that SNX9 in cytosol in the of for in and and for and the of of SNX9 the the binding of SNX9 in of with the that for of the with the of that in complex with SNX9 of SNX9 and and the cytosol for as the for SNX9 and and for binding of proteins as a with The the of of cytosol the for binding of with with and in the of with proteins as together with and SNX9 in cytosol as a complex, of SNX9 that and that the binding of the that for that the resting complex of SNX9 in the cytosol for membrane binding of the of that the with cytosol the of the membrane for binding of the membrane binding of SNX9 and cytosol of SNX9 and and in the of SNX9 in complex with in the in the of SNX9 in the cytosol of the that the complex and with for of the the protein. in the binding of SNX9 SNX9 for the binding of that SNX9 can in the cytosol of cytosol can be with SNX9 for the binding of that a that the can the binding of The show that the membrane with SNX9 and that the membrane binding in the SNX9 the binding of SNX9 a as for SNX9 the binding be with and and that in a of with cytosol together with and SNX9 binding and in with the binding of SNX9 and with of in the cytosol the the cytosol and with and and The that a of the the cytosol with and that the SNX9 complex in cytosol the binding for that in the cytosol can the for activated SNX9 with and of SNX9 and and with and for together with cytosol cytosol in of for the of and for proteins and and and and of of and and and with for proteins and The the of of cytosol the and of of the as in the binding as SNX9 for the of SNX9 for membrane binding of for in the with with the of and The show that SNX9 a for SNX9 as a that The and for the of the complex in cytosol in the with in the that SNX9 and for binding the of SNX9 for the of in in with the and in SNX9 the of SNX9 of and the of SNX9 cytosol and a membrane that together the of SNX9 and the of in the that the of in SNX9 and in the cytosol and of a in the and in the of the the in SNX9 with the a that with the of and that of that can for the of of SNX9 the of and in and in that SNX9 for a and membrane of a resting the for of the complex, of the of the 41-kDa in the complex of of cytosol with the 41-kDa as the the of the in the complex, and with and and the with and in and with SNX9 cytosol of SNX9 with that of the the proteins in the be the and with cytosol and proteins and of the of SNX9 in the that the binding of cytosol the of the in SNX9 a that binding for and The binding with a SNX9 and that of in the be for and binding the binding of in The binding of SNX9 be the of of the that the of in a for in the that in with a of the as a in cytosol and the in SNX9 the a of the and SNX9 be that the membrane binding of SNX9 the the of a of together with the and and with of the binding of the a of that in cytosol be The the the binding of the and The that SNX9 in the complex in membrane binding of a of the binding of SNX9 and the of as in and the for and the the in together with of The show the as binding of in the of of the binding of SNX9 be that a in the that SNX9 activated for with the of the complex SNX9 with the binding of that the of of proteins with cytosol in the of that in the that the the The the that a in the cytosol the in for the of SNX9 the for of and for the of the complex the the binding of SNX9 cytosol of in the the as the a a and the of the complex for binding of SNX9 the of the of a in the the of the in in the a SNX9 the and of a SNX9 the the of a in and the in and in and in the cytosol as a resting complex of SNX9 for membrane binding and with and The complex activated of the in SNX9 a that and SNX9 for binding membrane the SNX9 be SNX9 as a be the the of SNX9 that cytosol and a of together with a of SNX9 as SNX9 and the membrane as that SNX9 SNX9 and assemble and with the membrane for the of the complex and of SNX9 and the SNX9 with a resting complex in the together with as and as The and of SNX9 and The of the proteins in the complex with the of the SNX9 SNX9 a a in SNX9 as a the and that the of a of the in SNX9 as a SNX9 as a the membrane and the and of the membrane the The in SNX9 and for with and a the with SNX9 and the of SNX9 for of as a that be the membrane the the of the of and The the of in a of proteins and that can and membrane We and that the in SNX9 in a SNX9 that and the the can for SNX9 SNX9 as a in the complex the membrane as a of the the SNX9 and assemble the SNX9 that the membrane the proteins in the for proteins as the the the the and the and the We that binding of proteins can SNX9 the the of SNX9 in the membrane of SNX9 for binding of proteins the of the complex the of of a of and the of the in in that a the of a of SNX9 that the of a the of in the membrane the of SNX9 for that for of with the of be the of the in a for the and the cytosol be that SNX9 and the membrane membrane in a that SNX9 the of in the of that that activated in the cytosol the of the complex the binding of SNX9 the a in the that in in for that the as in of for that with in show in of binding that and proteins of the binding of the of as for SNX9 in the as that a the of in the The of of a of of in the binding of the of the and proteins the of the of the in SNX9 in of the in SNX9 of SNX9 in in of the membrane in the in in in the of that of proteins be the of SNX9 in the that for membrane of in in the membrane proteins and for a of the The of SNX9 in with SNX9 in the that for SNX9 and of and and SNX9 The of a complex SNX9 and in the cytosol for a of The that a of that be for The of for the that SNX9 and in the of the with The proteins in in the in as in and and The in the of the the of the the of the The for the of a the of a of proteins the and complex sorting nexin sorting nexin in the of the membrane and of a in a the of the and the the The be and in the a that in the of in in the of and the of the membrane for with in the the a the for a of proteins in the of the of that the of the be for the of the cytosol the membrane of the membrane of the in the in the dynamin-2 (Dyn2) in of the and that with as the in be for nexin 9 a of the sorting nexin of proteins a be in the endocytic as SNX9 binding for and in a and in the SNX9 membrane binding a a and a SNX9 with and the and in and of of SNX9 the of the cytosol of of SNX9 be in a complex together with and of The the of complex and the membrane of SNX9 and We show that SNX9 the of the and that the membrane binding of SNX9 a with the a with and the in and as in and for the in and for a as the cytosol with and cytosol in a of as The 41-kDa of the with and a the and of SNX9 as proteins as and The in the and as with proteins as and a for The and for in of The and in The as in a of of cytosol and and and and as in the with and cytosol the with the The in and the the a for the for with of and The in of in and the for as of in of of in cytosol with as in the for the for with of and for with cytosol with as in the of of in in and SNX9 with and the in the and the in in of SNX9 with the of cytosol for SNX9 with as in for in the of a of cytosol and in and The and the the cytosol with for in the of and and a and for the of SNX9 and for with cytosol in the of and with as the and proteins with cytosol and for the proteins a and with The for and cytosol together with and in the of and together with and The The for and SNX9 and of proteins binding for in of of and of in in and for with and as that and for the with and a the of and in in and of and of and cytosol and a a as and the of The and the a a of the The as a a with and the in and as in and for the in and for a as the cytosol with and cytosol in a of as The 41-kDa of the with and a the and of SNX9 as proteins as and The in the and as with proteins as and a for The and for in of The and in The as in a of of cytosol and and and and as in the with and cytosol the with the The in and the the a for the for with of and The in of in and the for as of in of of in cytosol with as in the for the for with of and for with cytosol with as in the of of in in and SNX9 with and the in the and the in in of proteins binding for in of of and of in in and for with and as that and for the with and a the of and in in and of and of and cytosol and a a as and the of The and the a a of the The as a the membrane binding of SNX9 and with and of SNX9 and The in show that SNX9 in cytosol in the of for in and and for and the of of SNX9 the the binding of SNX9 in of with the that for of the with the of that in complex with SNX9 SNX9 in cytosol as a complex, of SNX9 that and that the binding of the that for that the resting complex of SNX9 in the cytosol for membrane binding of the of that the with cytosol the of the membrane for binding of the membrane binding of SNX9 and cytosol of SNX9 and and in the of SNX9 in complex with in the in the of SNX9 in the cytosol of the that the complex and with for of the the protein. in the binding of SNX9 SNX9 for the binding of that SNX9 can in the cytosol of cytosol can be with SNX9 for the binding of that a that the can the binding of The show that the membrane with SNX9 and that the membrane binding in the SNX9 the binding of SNX9 a as for SNX9 the binding be with and and that in a of with cytosol together with and SNX9 binding and in with the binding of SNX9 and with of in the cytosol the the cytosol and with and and The that a of the the cytosol with and that the SNX9 complex in cytosol the binding for that in the cytosol can the for activated SNX9 with and of SNX9 and and with and for together with cytosol cytosol in of for the of and for proteins and and and and of of and and and with for proteins and The the of of cytosol the and of of the as in the binding as SNX9 for the of SNX9 for membrane binding of for in the with with the of and The show that SNX9 a for SNX9 as a that The and for the of the complex in cytosol in the with in the that SNX9 and for binding the of SNX9 for the of in in with the and in SNX9 the of SNX9 of and the of SNX9 cytosol and a membrane that together the of SNX9 and the of in the that the of in SNX9 and in the cytosol and of a in the and in the of the the in SNX9 with the a that with the of and that of that can for the of of SNX9 the of and in and in that SNX9 for a and membrane of a resting the for of the complex, of the of the 41-kDa in the complex of of cytosol with the 41-kDa as the the of the in the complex, and with and and the with and in and with SNX9 cytosol of SNX9 with that of the the proteins in the be the and with cytosol and proteins and of the of SNX9 in the that the binding of cytosol the of the in SNX9 a that binding for and The binding with a SNX9 and that of in the be for and binding the binding of in The binding of SNX9 be the of of the that the of in a for in the that in with a of the as a in cytosol and the in SNX9 the a of the and SNX9 be that the membrane binding of SNX9 the the of a of together with the and and with of the binding of the a of that in cytosol be The the the binding of the and The that SNX9 in the complex in membrane binding of a of the binding of SNX9 and the of as in and the for and the the in together with of The show the as binding of in the of of the binding of SNX9 be that a in the that SNX9 activated for with the of the complex SNX9 with the binding of that the of of proteins with cytosol in the of that in the that the the The the that a in the cytosol the in for the of SNX9 the for of and for the of the complex the the binding of SNX9 cytosol of in the the as the a a and the of the complex for binding of SNX9 the of the of a in the the of the We the membrane binding of SNX9 and with and of SNX9 and The in show that SNX9 in cytosol in the of for in and and for and the of of SNX9 the the binding of SNX9 in of with the that for of the with the of that in complex with SNX9 SNX9 in cytosol as a complex, of SNX9 that and that the binding of the that for that the resting complex of SNX9 in the cytosol for membrane binding of the of that the with cytosol the of the membrane for binding of the membrane binding of SNX9 and cytosol of SNX9 and and in the of SNX9 in complex with in the in the of SNX9 in the cytosol of the that the complex and with for of the the protein. in the binding of SNX9 SNX9 for the binding of that SNX9 can in the cytosol of cytosol can be with SNX9 for the binding of that a that the can the binding of The show that the membrane with SNX9 and that the membrane binding in the SNX9 the binding of SNX9 a as for SNX9 the binding be with and and that in a of with cytosol together with and SNX9 binding and in with the binding of SNX9 and with of in the cytosol the the cytosol and with and and The that a of the the cytosol with and that the SNX9 complex in cytosol the binding for that in the cytosol can the for activated SNX9 with and the binding as SNX9 for the of SNX9 for membrane binding of for in the with with the of and The show that SNX9 a for SNX9 as a that The and for the of the complex in cytosol in the with in the that SNX9 and for binding the The of SNX9 for the of in in with the and in SNX9 the of SNX9 of and the of SNX9 cytosol and a membrane that together the of SNX9 and the of in the that the of in SNX9 and in the cytosol and of a in the and in the of the the in SNX9 with the a that with the of and that of that can for the of of SNX9 the of and The in and in that SNX9 for a and membrane of a resting the for of the complex, of the of the 41-kDa in the complex of of cytosol with the 41-kDa as the the of the in the complex, and with and and the with and in and with SNX9 cytosol of SNX9 with that of the the proteins in the be the and with cytosol and proteins and of the of SNX9 in the that the binding of cytosol the of the in SNX9 a that binding for and The binding with a SNX9 and that of in the be for and binding the binding of in The binding of SNX9 be the of of the that the of in a for in the that in with a of the as a in cytosol and the in SNX9 the a of the and SNX9 be that the membrane binding of SNX9 the the of a of together with the and and with of the binding of the a of that in cytosol be The the the binding of the and The that SNX9 in the complex in membrane binding of a of the binding of SNX9 be that a in the that SNX9 activated for with the of the complex SNX9 with the binding of that the of of proteins with cytosol in the of that in the that the the The the that a in the cytosol the in for the of SNX9 the for of and for the of the complex the the binding of SNX9 cytosol of in the the as the a a and the of the complex for binding of SNX9 the of the of a in the the of the in in the a SNX9 the and of a SNX9 the the of a in and the in and in and in the cytosol as a resting complex of SNX9 for membrane binding and with and The complex activated of the in SNX9 a that and SNX9 for binding membrane the SNX9 be SNX9 as a be the the of SNX9 that cytosol and a of together with a of SNX9 as SNX9 and the membrane as that SNX9 SNX9 and assemble and with the membrane of as a that be the membrane the the of the of and The the of in a of proteins and that can and membrane We and that the in SNX9 in a SNX9 that and the the can for SNX9 SNX9 as a in the complex the membrane as a of the the SNX9 and assemble the SNX9 that the membrane the proteins in the for proteins as the the the the and the and the We that binding of proteins can SNX9 the the of SNX9 in the membrane of SNX9 for binding of proteins the of the complex the of of a of and the of the in in that a the of a of SNX9 that the of a the of in the membrane the of SNX9 for that for of with the of be the of the in a for the and the cytosol be that SNX9 and the membrane membrane in a that SNX9 the of in the of that that activated in the cytosol the of the complex the binding of SNX9 the a in the that in in for that the as in of for that with in show in of binding that and proteins of the binding of the of as for SNX9 in the as that a the of in the The of of a of of in the binding of the of the and proteins the of the of the in SNX9 in of the in SNX9 of SNX9 in in of the membrane in the in in in the of that of proteins be the of SNX9 in the that for membrane of in in the membrane proteins and for a of the The of SNX9 in with SNX9 in the that for SNX9 and of and and SNX9 The of a complex SNX9 and in the cytosol for a of The that a of that be for The of for the that SNX9 and in the of the with The in in the a SNX9 the and of a SNX9 the the of a in and the in and in and in the cytosol as a resting complex of SNX9 for membrane binding and with and The complex activated of the in SNX9 a that and SNX9 for binding membrane the SNX9 be SNX9 as a be the the of SNX9 that cytosol and a of together with a of SNX9 as SNX9 and the membrane as that SNX9 SNX9 and assemble and with the membrane The of as a that be the membrane the the of the of and The the of in a of proteins and that can and membrane We and that the in SNX9 in a SNX9 that and the the can for SNX9 SNX9 as a in the complex the membrane as a of the the SNX9 and assemble the SNX9 that the membrane the proteins in the for proteins as the the the the and the and the We that binding of proteins can SNX9 the the of SNX9 in the membrane of SNX9 for binding of proteins the The of the complex the of of a of and the of the in in that a the of a of SNX9 that the of a the of in the membrane the of SNX9 for that for of with the of be the of the in a for the and the cytosol be that SNX9 and the membrane membrane in a that SNX9 the of in the of that that activated in the cytosol the of the complex the binding of SNX9 the a in the that in in for that the as in of for that with in show in of binding that and proteins of the binding of the of as for SNX9 in the as that a the of in the The of of a of of in the binding of the of the and proteins the of the of the in SNX9 in of the in SNX9 of SNX9 in in of the membrane in the in in in the of that of proteins be the of SNX9 in the that for membrane of in in the membrane proteins and for a of the The of SNX9 in with SNX9 in the that for SNX9 and of and and SNX9 The of a complex SNX9 and in the cytosol for a of The that a of that be for The of for the that SNX9 and in the of the with We for