Association of RNA Helicase A with Human Immunodeficiency Virus Type 1 Particles

RNA helicase A (RHA) belongs to the DEAH family of proteins that are capable of unwinding double-stranded RNA structure. In addition to its involvement in the metabolism of cellular RNA, RHA has been shown to stimulate RNA transcription from the long terminal repeat promoter of human immunodeficiency virus type 1 (HIV-1) as well as to enhance Rev/Rev response element-mediated gene expression. In this study, we provide evidence that RHA associates with HIV-1 in in of RHA HIV-1 of RHA in virus to of HIV-1 that are in as of RHA the of cellular RNA that in HIV-1 RNA helicase A (RHA) belongs to the DEAH family of proteins that are capable of unwinding double-stranded RNA structure. In addition to its involvement in the metabolism of cellular RNA, RHA has been shown to stimulate RNA transcription from the long terminal repeat promoter of human immunodeficiency virus type 1 (HIV-1) as well as to enhance Rev/Rev response element-mediated gene expression. In this study, we provide evidence that RHA associates with HIV-1 in in of RHA HIV-1 of RHA in virus to of HIV-1 that are in as of RHA the of cellular RNA that in HIV-1 RNA helicase A (RHA) RNA helicase human immunodeficiency type response response of RNA helicase human immunodeficiency type response response of of the family of proteins with the the family are to as RNA that are to the of RNA RHA helicase of that are RNA the of RHA are of type A double-stranded RNA with the double-stranded RNA RNA as well as helicase of RHA RHA the the with the of of the of the RHA to of RHA to double-stranded RNA with the from of of its RHA to in cellular from RNA transcription to RNA to RNA the of RHA in the of the with RHA of RHA in RNA transcription its the RNA RHA has been shown to the RNA transcription as RHA with the of gene of RHA in transcription the of its in the that gene from the RHA has been as of of RHA In to RNA RHA to the RNA of type as with cellular RHA of RNA of its in cellular RNA that RHA the of human immunodeficiency virus type 1 (HIV-1) RHA transcription of HIV-1 RNA of RHA to the response that the of HIV-1 the double-stranded RNA of of that to RHA the of the transcription of RNA RHA Rev/Rev response gene RHA with in the of HIV-1 RNA, RHA the of HIV-1 RNA from In addition to its involvement in HIV-1 gene we that RHA associates with the virus of RHA in HIV-1 the with the in the of that the with to of the from transcription the of the with as with to the the from with the with the the with the the with the in as in of with with to in as in of with with to to the of with the from the with to with in the with with 1 the with in proteins with in 1 1 with 1 1 the proteins with 1 1 proteins with to of with HIV-1 in the in the virus the of the to In to the of cellular proteins with virus with in the with RNA from virus in in In RNA with of RNA to with RNA to RNA with HIV-1 RNA to of RNA that been with RNA the cellular RNA with in with the with with the with of with with 1 the from of in of RHA with to RHA from to of RHA in with to of as RNA from virus the of RNA in from the of RNA HIV-1 in of to the the in to of virus from with to the virus to 1 of in 1 with with of in with with the 1 in the of of to of to of in the the that are with HIV-1 long terminal with to of the of to the of the of has been in to in cellular that with HIV-1 we the of A from the to the of HIV-1 of from to the the in to from the with that the that the of the proteins shown in as of with in are of cellular RNA RNA helicase of that RHA RNA helicase with RHA to with virus in to the of In this study, we the of RHA in of virus with HIV-1 of the cellular proteins that with HIV-1 of the shown the A are to the of in the of HIV-1 of with to the proteins with of proteins shown the of the with the the A with HIV-1 in we the from in that RHA with in the the from the of RHA in the in of the to in the of that from the with the the with the from transcription the the with RHA the of in we that of from the the that RNA the as the from HIV-1 the in of RHA to with the the that the of the in to of RHA that In the of that the with RHA with RHA in RHA associates with HIV-1 the with the from the transcription proteins from the in the from the as in the of in as the with the with the in the to are with the the of of RHA with proteins in of RHA to with the in cellular RNA as shown the of the of RHA in of RHA with the that the gene to the of in the of the HIV-1 RHA with of RHA of this A to of A the of of that A the of RHA that with that of RHA the the HIV-1 to the to to RHA that with this with the of are shown in that RHA the with the of RHA the with the of RHA as of the the RHA we that RHA the to the of RHA the we to the RHA this HIV-1 of the of RHA virus we HIV-1 virus RHA to with virus RHA virus to the virus the of proteins as well as the of of that the of the proteins with of with RNA, with the the virus the of the that with virus virus as well the RHA in the the of RHA with virus of RHA HIV-1 of RHA with virus to of with with in to RHA as well as proteins as of RNA with the with to the RNA with RNA that been with of in to the of the of with to of RHA with virus of RHA with HIV-1 virus the that of RHA with virus of that with we HIV-1 that the in the of the as to to the of the this with RHA RHA are to the virus of RHA as with the of RHA that with the of RHA of RHA with the the virus of that the RHA with virus to the evidence of RHA with virus HIV-1 the RNA from the cellular RNA virus of that RHA RNA as the to with virus this the with the that with the to to that RNA with that RHA to to HIV-1 RNA of HIV-1 RHA with HIV-1 RNA with the that the to RNA that RNA of RNA from in to the of RNA in of in to the of the of RHA the virus to of in to of RHA to of in to the of the virus of RNA with the of virus in that the RNA of are of the RHA HIV-1 RNA the RHA in virus that we HIV-1 that RNA with the virus of RNA of of RHA that the as with the virus with HIV-1 RNA RHA to virus of RHA to of with RHA in of virus this we with that RHA of of RHA in RHA RHA as with with of HIV-1 with of the that the of that the HIV-1 RNA virus of virus as of RHA the as with that that RHA in as with from the in virus the of of virus in that been with RHA RHA of virus virus of RHA with RHA with the HIV-1 in the of virus in in the to virus of virus in that RHA to with RHA in the with that of the virus in as from RHA to RNA RHA with in HIV-1 RNA that RNA are virus in are the transcription that of RHA the of virus RNA we to of RNA with virus that RHA the of RNA of RNA in that RHA to in of the of RNA as with with A of RHA HIV-1 RNA RNA in RNA the RNA are RNA from virus to of from to of RNA RNA in RNA from virus to of in HIV-1 as of RNA that are RHA HIV-1 RNA we to RNA that from virus that RNA of been with RHA RHA RNA the that RHA RNA in that the of RHA has in HIV-1 we to the of the that to RNA virus A in the of virus that RHA that RHA to the transcription of this RNA transcription of in that the RHA virus of that of that the virus in as with the RHA virus with the RHA in the to of virus with transcription of the to of RNA in the of RNA in the from human to in HIV-1 RNA to the the as transcription HIV-1 are with from this RNA that to the the the as to the from in to the of are shown in the as well as in the with from virus as this study, we provide evidence that RHA associates with HIV-1 the of HIV-1 the of the that RHA of the HIV-1 virus the of RNA of the RNA that been to in HIV-1 the of HIV-1 RNA shown to gene with the RNA helicase of HIV-1 RNA of of as in HIV-1 with the of proteins in HIV-1 been the RNA RHA the that shown to with HIV-1 In of the with RNA to that of proteins to HIV-1 RNA the of of RNA that with that RNA are to HIV-1 RNA HIV-1 RNA that to with the RNA the that of virus that the virus RNA as to in of RNA of cellular to as of RHA are that to A this we are to the that RHA RHA in to the of to RNA RHA to we that RHA to HIV-1 RNA in the of of RHA to HIV-1 RNA the of of RHA RNA with the that RHA with the of HIV-1 RNA HIV-1 gene the that RHA with HIV-1 RNA the that this the that HIV-1 RNA virus to HIV-1 RNA, RHA to the virus as well as the transcription virus the of of its helicase the cellular that has been shown to with virus to the of HIV-1 of the in the as the of that with this shown that RNA RNA virus as as virus to cellular of RHA with HIV-1 virus the of that from RHA that RHA in the of virus the virus the virus the of to in RNA that with RHA to the of in of RNA of that the transcription of virus RHA RHA in the of we provide evidence the involvement of RHA in the of to transcription in to RNA unwinding RNA RHA virus RNA as well as the of RNA with as to transcription that RHA the cellular the of this the of RHA with HIV-1 virus that to virus RHA to the of RNA as well as to RNA helicase A (RHA) RNA helicase human immunodeficiency type response response of RNA helicase human immunodeficiency type response response of of the family of proteins with the the family are to as RNA that are to the of RNA RHA helicase of that are RNA the of RHA are of type A double-stranded RNA with the double-stranded RNA RNA as well as helicase of RHA RHA the the with the of of the of the RHA to of RHA to double-stranded RNA with the from of of its RHA to in cellular from RNA transcription to RNA to RNA the of RHA in the of the with RHA of RHA in RNA transcription its the RNA RHA has been shown to the RNA transcription as RHA with the of gene of RHA in transcription the of its in the that gene from the RHA has been as of of RHA In to RNA RHA to the RNA of type as with cellular RHA of RNA In of its in cellular RNA that RHA the of human immunodeficiency virus type 1 (HIV-1) RHA transcription of HIV-1 RNA of RHA to the response that the of HIV-1 the double-stranded RNA of of that to RHA the of the transcription of RNA RHA Rev/Rev response gene RHA with in the of HIV-1 RNA, RHA the of HIV-1 RNA from In addition to its involvement in HIV-1 gene we that RHA associates with the virus of RHA in HIV-1 the with the in the of that the with to of the from transcription the of the with as with to the the from with the with the the with the the with the in as in of with with to in as in of with with to to the of with the from the with to with in the with with 1 the with in proteins with in 1 1 with 1 1 the proteins with 1 1 proteins with to of with HIV-1 in the in the virus the of the to In to the of cellular proteins with virus with in the with RNA from virus in in In RNA with of RNA to with RNA to RNA with HIV-1 RNA to of RNA that been with RNA the cellular RNA with in with the with with the with of with with 1 the from of in of RHA with to RHA from to of RHA in with to of as RNA from virus the of RNA in from the of RNA HIV-1 in of to the the in to of virus from with to the virus to 1 of in 1 with with of in with with the 1 in the of of to of to of in the the that are with HIV-1 long terminal with to of the of to the of the with the in the of that the with to of the from transcription the of the with as with to the the from with the with the the with the the with the in as in of with with to in as in of with with to to the of with the from the with to with in the with with 1 the with in proteins with in 1 1 with 1 1 the proteins with 1 1 proteins with to of with HIV-1 in the in the virus the of the to In to the of cellular proteins with virus with in the with RNA from virus in in In RNA with of RNA to with RNA to RNA with HIV-1 RNA to of RNA that been with RNA the cellular RNA with in with the with with the with of with with 1 the from of in of RHA with to RHA from to of RHA in with to of as RNA from virus the of RNA in from the of RNA HIV-1 in of to the the in to of virus from with to the virus to 1 of in 1 with with of in with with the 1 in the of of to of to of in the the that are with HIV-1 long terminal with to of the of to the of the of has been in to in cellular that with HIV-1 we the of A from the to the of HIV-1 of from to the the in to from the with that the that the of the proteins shown in as of with in are of cellular RNA RNA helicase of that RHA RNA helicase with RHA to with virus in to the of In this study, we the of RHA in of virus with HIV-1 in we the from in that RHA with in the the from the of RHA in the in of the to in the of that from the with the the with the from transcription the the with RHA the of in we that of from the the that RNA the as the from HIV-1 the in of RHA to with the the that the of the in to of RHA that In the of that the with RHA with RHA in RHA associates with HIV-1 the with the from the transcription proteins from the in the from the as in the of in as the with the with the in the to are with the the of of RHA with proteins in of RHA to with the in cellular RNA as shown the of the of RHA in of RHA with the that the gene to the of in the of the HIV-1 RHA with of RHA of this A to of A the of of that A the of RHA that with that of RHA the the HIV-1 to the to to RHA that with this with the of are shown in that RHA the with the of RHA the with the of RHA as of the the RHA we that RHA the to the of RHA the we to the RHA this HIV-1 of the of RHA virus we HIV-1 virus RHA to with virus RHA virus to the virus the of proteins as well as the of of that the of the proteins with of with RNA, with the the virus the of the that with virus virus as well the RHA in the the of RHA with virus of RHA HIV-1 of RHA with virus to of with with in to RHA as well as proteins as of RNA with the with to the RNA with RNA that been with of in to the of the of with to of RHA with virus of RHA with HIV-1 virus the that of RHA with virus of that with we HIV-1 that the in the of the as to to the of the this with RHA RHA are to the virus of RHA as with the of RHA that with the of RHA of RHA with the the virus of that the RHA with virus to the evidence of RHA with virus HIV-1 the RNA from the cellular RNA virus of that RHA RNA as the to with virus this the with the that with the to to that RNA with that RHA to to HIV-1 RNA of HIV-1 RHA with HIV-1 RNA with the that the to RNA that RNA of RNA from in to the of RNA in of in to the of the of RHA the virus to of in to of RHA to of in to the of the virus of RNA with the of virus in that the RNA of are of the RHA HIV-1 RNA the RHA in virus that we HIV-1 that RNA with the virus of RNA of of RHA that the as with the virus with HIV-1 RNA RHA to virus of RHA to of with RHA in of virus this we with that RHA of of RHA in RHA RHA as with with of HIV-1 with of the that the of that the HIV-1 RNA virus of virus as of RHA the as with that that RHA in as with from the in virus the of of virus in that been with RHA RHA of virus virus of RHA with RHA with the HIV-1 in the of virus in in the to virus of virus in that RHA to with RHA in the with that of the virus in as from RHA to RNA RHA with in HIV-1 RNA that RNA are virus in are the transcription that of RHA the of virus RNA we to of RNA with virus that RHA the of RNA of RNA in that RHA to in of the of RNA as with with A of RHA HIV-1 RNA RNA in RNA the RNA are RNA from virus to of from to of RNA RNA in RNA from virus to of in HIV-1 as of RNA that are RHA HIV-1 RNA we to RNA that from virus that RNA of been with RHA RHA RNA the that RHA RNA in that the of RHA has in HIV-1 we to the of the that to RNA virus A in the of virus that RHA that RHA to the transcription of this RNA transcription of in that the RHA virus of that of that the virus in as with the RHA virus with the RHA in the to of virus with transcription of the to of RNA in the of RNA in the from human to in HIV-1 RNA to the the as transcription HIV-1 are with from this RNA that to the the the as to the from in to the of are shown in the as well as in the with from virus as HIV-1 the of has been in to in cellular that with HIV-1 we the of A from the to the of HIV-1 of from to the the in to from the with that the that the of the proteins shown in as of with in are of cellular RNA RNA helicase of that RHA RNA helicase with RHA to with virus in to the of In this study, we the of RHA in of virus RHA with HIV-1 in we the from in that RHA with in the the from the of RHA in the in of the to in the of that from the with the the with the from transcription the the with RHA the of in we that of from the the that RNA the as the from HIV-1 the in of RHA to with the the that the of the in to of RHA that In the of that the with RHA of RHA of this A to of A the of of that A the of RHA that with that of RHA the the HIV-1 to the to to RHA that with this with the of are shown in that RHA the with the of RHA the with the of RHA as of the the RHA we that RHA the to the of RHA the we to the RHA this RHA HIV-1 of the of RHA virus we HIV-1 virus RHA to with virus RHA virus to the virus the of proteins as well as the of of that the of the proteins with of with RNA, with the the virus the of the that with virus virus as well the RHA in the the of RHA with virus the that of RHA with virus of that with we HIV-1 that the in the of the as to to the of the this with RHA RHA are to the virus of RHA as with the of RHA that with the of RHA of RHA with the the virus of that the RHA with virus to the evidence of RHA with virus RHA HIV-1 the RNA from the cellular RNA virus of that RHA RNA as the to with virus this the with the that with the to to that RNA with that RHA to to HIV-1 RNA of of the RHA HIV-1 RNA the RHA in virus that we HIV-1 that RNA with the virus of RNA of of RHA that the as with the virus with HIV-1 RNA RHA to virus of RHA to of with RHA in of virus this we with that RHA of of RHA in RHA RHA as with with of HIV-1 with of the that the of that the HIV-1 RNA virus of virus as of RHA the as with that that RHA in as with from the in virus the of of virus in that been with RHA RHA of virus from RHA to RNA RHA with in HIV-1 RNA that RNA are virus in are the transcription that of RHA the of virus RNA we to of RNA with virus that RHA the of RNA of RNA in that RHA to in of the of RNA as with with A HIV-1 of RNA that are RHA HIV-1 RNA we to RNA that from virus that RNA of been with RHA RHA RNA the that RHA RNA in that the of RHA has in HIV-1 we to the of the that to RNA virus A in the of virus that RHA that RHA to the transcription of this RNA transcription of in that the RHA virus of that of that the virus in as with the RHA virus with the RHA in the this study, we provide evidence that RHA associates with HIV-1 the of HIV-1 the of the that RHA of the HIV-1 virus the of RNA of the RNA that been to in HIV-1 the of HIV-1 RNA shown to gene with the RNA helicase of HIV-1 RNA of of as in HIV-1 with the of proteins in HIV-1 been the RNA RHA the that shown to with HIV-1 In of the with RNA to that of proteins to HIV-1 RNA the of of RNA that with that RNA are to HIV-1 RNA HIV-1 RNA that to with the RNA the that of virus that the virus RNA as to in of RNA of cellular to as of RHA are that to A this we are to the that RHA RHA in to the of to RNA RHA to we that RHA to HIV-1 RNA in the of of RHA to HIV-1 RNA the of of RHA RNA with the that RHA with the of HIV-1 RNA HIV-1 gene the that RHA with HIV-1 RNA the that this the that HIV-1 RNA virus to HIV-1 RNA, RHA to the virus as well as the transcription virus the of of its helicase the cellular that has been shown to with virus to the of HIV-1 of the in the as the of that with this shown that RNA RNA virus as as virus to cellular of RHA with HIV-1 virus the of that from RHA that RHA in the of virus the virus the virus the of to in RNA that with RHA to the of in of RNA of that the transcription of virus RHA RHA in the of we provide evidence the involvement of RHA in the of to transcription in to RNA unwinding RNA RHA virus RNA as well as the of RNA with as to transcription that RHA the cellular the of this the of RHA with HIV-1 virus that to virus RHA to the of RNA as well as to In this study, we provide evidence that RHA associates with HIV-1 the of HIV-1 the of the that RHA of the HIV-1 virus the of RNA RHA of the RNA that been to in HIV-1 the of HIV-1 RNA shown to gene with the RNA helicase of HIV-1 RNA of of as in HIV-1 with the of proteins in HIV-1 been the RNA RHA the that shown to with HIV-1 In of the with RNA to that of proteins to HIV-1 RNA the of of RNA that with that RNA are to HIV-1 RNA HIV-1 RNA that to with the RNA the that of virus that the virus RNA as to in of RNA of cellular to as of RHA are that to A this we are to the that RHA RHA in to the of to RNA RHA to we that RHA to HIV-1 RNA in the of of RHA to HIV-1 RNA the of of RHA RNA with the that RHA with the of HIV-1 RNA HIV-1 gene the that RHA with HIV-1 RNA the that this the that HIV-1 RNA virus to HIV-1 RNA, RHA to the virus as well as the transcription virus the of of its helicase RHA the cellular that has been shown to with virus to the of HIV-1 of the in the as the of that with this shown that RNA RNA virus as as virus to cellular of RHA with HIV-1 virus the of that from RHA that RHA in the of virus the virus the virus the of to in RNA that with RHA to the of in of RNA of that the transcription of virus RHA RHA in the of we provide evidence the involvement of RHA in the of RHA to transcription in to RNA unwinding RNA RHA virus RNA as well as the of RNA with as to transcription that RHA the cellular In the of this the of RHA with HIV-1 virus that to virus RHA to the of RNA as well as to RHA the

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