HIV Databases HIV Databases home HIV Databases home
HIV sequence database



Coreceptor Use by Primate Lentiviruses

Robert W. Doms, 1 Aimee L. Edinger,2 and John P. Moore3

1 Corresponding author: University of Pennsylvania, Department of Pathology and Laboratory Medicine, 807 Abramson, 34th and Civic Center Blvd., Philadelphia, PA 19104
phone: 215-898-0890, fax: 215-573-2883, email: doms@mail.med.upenn.edu

2 University of Pennsylvania, Department of Pathology and Laboratory Medicine, Philadelphia, PA 19104

3 The Aaron Diamond AIDS Research Center, The Rockefeller University, 455 First Avenue, New York, NY 10016

Since the original compilation of the coreceptors used by HIV-1 strains in 1998, considerable progress has been made in identifying coreceptors for SIV and HIV-2 strains. Hence, in addition to updating the record of coreceptor usage by HIV-1 strains, tables describing the coreceptors used by SIV and HIV-2 strains are now provided.

HIV-1

CCR5 and CXCR4 are the major coreceptors used by HIV-1 strains. All HIV-1 strains reported on to date use one or both of these receptors. For the vast majority of strains, CCR5 and CXCR4 support virus entry more efficiently than any of the growing number of alternative coreceptors, including CCR2, CCR3, CCR8, CX3CR1 (formerly V28), STRL33 (Bonzo), GPR1, GPR15 (BOB), APJ, and ChemR23. Each of these alternative coreceptors can function for virus entry or Env-mediated cell-cell fusion, to a greater or lesser extent, when they are transfected into CD4+ cell lines in vitro . However, none of them appears to be efficiently utilized by large numbers of virus strains. The question arises as to whether use of alternative coreceptors in vivo impacts viral tropism and pathogenesis. Potentially, use of receptors other than CCR5 or CXCR4 could enable virus strains to infect new types of cells, for example in the brain, thymus or mucosa. However, to the best of our knowledge infection of primary cells by HIV-1 is always mediated by CCR5 or CXCR4. It is particularly striking that infection of primary cells (CD4+ T-cells and macrophages) obtained from CCR5-negative individuals is always mediated by CXCR4. Use of receptors other than CXCR4 to infect these CD4-negative cells has not yet been demonstrated. Development of antibodies to the alternative coreceptors is needed, however, to determine the cell types in which they are expressed. Specific antibodies, or other inhibitors, targeted at the alternative coreceptors would also be helpful to gauge their importance in primary cells. Our experience is, however, that inhibitors targeted at CCR5 and CXCR4 are sufficient to inhibit the replication of all tested HIV-1 strains in primary cells.

The HIV-1 coreceptor use Table is updated from last year. The major coreceptor(s) used by each virus strain is listed, as are alternative coreceptors that support virus infection in vitro to an extent that is >10% of the efficiency of the major coreceptor used by each virus strain.

Proposed             Primary  Other 
Strain     Accessiona   Clade   Tropismb    Designationc     Receptord    Receptorse     References

DJ258 L22939 A NSI R5 CCR5 [Trkola (1998)] 92RW026 NA A NSI R5 CCR5 [Trkola (1998)] 93KE101 NA A NSI R5 CCR5 [Zhang (1996)] 93IN103 NA A NSI R5 CCR5 [Zhang (1996)] 92UG037-8 U51190 A NSI R5 CCR5 CCR8* [Bjorndal (1997), Rucker (1997)] 92RW020-5 U08794 A NSI R5 CCR5 [Rucker (1997)] 92UG31 L34667 A NSI R5 CCR5 [Dittmar (1997)] 92RW20 U08794 A NSI R5 CCR5 [Dittmar (1997)] 92UG029 NA A SI X4 CXCR4 [Trkola (1998)] 92RW009 U88823 A SI R5X4 CXCR4,CCR5 [Zhang (1996)] JR-FL U63632 B NSI R5 CCR5 CCR3 [Deng (1997), Farzan (1997),Rucker (1997)] JR-CSF M38429 B NSI R5 CCR5 [Simmons (1996), Trkola (1998),Zhang (1996)] SF162 M65024 B NSI R5 CCR5 STRL33* [Liao (1997),Rucker (1997)] YU2 M93258 B NSI R5 CCR5 CCR3,GPR15 [Choe (1996),Farzan (1997)] ADA AF004394 B NSI R5 CCR5 CCR3,GPR15, [Choe (1996), STRL33*,CCR8 Farzan (1997),Rucker (1997)] Ba-L M68893 B NSI R5 CCR5 CCR3,STRL33* [Deng (1997),Dragic (1996), Liao (1997),Rucker (1997)] 92US657 U04908 B NSI R5 CCR5 [Trkola (1998)] 92US715.6 U08451 B NSI R5 CCR5 [Bjorndal (1997)] 92Br20-4 U08797 B NSI R5 CCR5 [Choe96,Rucker (1997)] 91US005.11 U27434 B NSI R5 CCR5 [Bjorndal (1997), Rucker (1997)] SL-2 NA B NSI R5 CCR5 [Simmons (1996)] 92TH014.12 U08801 B NSI R5 CCR5 [Bjorndal (1997)] CM243 NA B NSI R5 CCR5 GPR15,STRL33 [Rucker (1997)] M23 NA B NSI R5 CCR5 [Dittmar (1997)] E80 NA B NSI R5 CCR5 [Dittmar (1997)] BR92 NA B NSI R5 CCR5 [Dittmar (1997)] BR49 NA B NSI R5 CCR5 [Dittmar (1997)] BR53 NA B NSI R5 CCR5 [Dittmar (1997)] BR90 NA B NSI R5 CCR5 [Dittmar (1997)] 92HA593g U08444 B SI R5X4 CXCR4,CCR5 [Zhang (1996)] 92HT593.1g U08444 B NSI R5X4 CXCR4,CCR5 [Bjorndal (1997)] 2028 NA B SI R5X4 CXCR4,CCR5 CCR3 [Dittmar (1997), Simmons (1996)] 2076 NA B SI R5X4 CXCR4,CCR5 [Dittmar (1997), Simmons (1996),Trkola (1998)] 89.6 U39362 B SI R5X4 CXCR4,CCR5 CCR3,CCR2b, [Farzan (1997),Rucker (1997)] CCR8,V28 DH123 NA B SI R5X4 CXCR4,CCR5 [Trkola (1998)] Isolate NA B SI R5X4 CXCR4,CCR5 [Trkola (1998)] C 7/86 92HA594 U08445 B SI R5X4 CXCR4,CCR5 [Zhang (1996)] 92HA596 U08446 B SI R5X4 CXCR4,CCR5 [Zhang (1996)] M13 NA B SI R5X4 CXCR4,CCR5 [Simmons (1996)] 2006 NA B SI R5X4 CXCR4,CCR5 [Simmons (1996)] 2044 NA B SI R5X4 CXCR4,CCR5 [Simmons (1996)] 2036 NA B SI R5X4 CXCR4,CCR5 [Simmons (1996)] 2005 NA B SI R5X4 CXCR4,CCR5 [Simmons (1996)] 92HT599.24 U08447 B SI X4 CXCR4 [Bjorndal (1997)] BK132 L03697 B SI X4 CXCR4 CCR3*,CCR8* [Rucker(1997)] BR65 NA B SI X4 CXCR4 [Dittmar (1997)] HC4 NA B SI X4 CXCR4 [Trkola (1998)] SF2 K02007 B SI/TCLA R5X4 CXCR4,CCR5 [Trkola (1998)] RF M17451 B SI/TCLA R5X4 CXCR4,CCR5 [Alkhatib (1996), Deng (1997),Rucker (1997)] NL 4-3 M19921 B SI/TCLA X4 CXCR4 [Trkola (1998),Zhang (1996)] LAI X01762 B SI/TCLA X4 CXCR4 [Trkola (1998)] HXBc2 K03455 B SI/TCLA X4 CXCR4 [Choe (1996)] GUN-1 D34590 B SI/TCLA R5X4 CXCR4,CCR5 [Simmons (1996)] BH8 K02011 B SI/TCLA X4 CXCR4 CCR3*,STRL33* [Rucker (1997)] P1 22-211-V4 B NSI R5 CCR5 [Zhang (1998)] P2 22-204-V2 B NSI R5 CCR5 [Zhang (1998)] P3 19-245-V6 B NSI R5 CCR5 [Zhang (1998)] P4 03-212-V8 B NSI R5 CCR5 [Zhang (1998)] P5 11-213-V4 B NSI R5 CCR5 [Zhang (1998)] P6 02-217-V3 B NSI R5 CCR5 BONZO [Zhang (1998)] P7 22-216-V5 B NSI R5 CCR5 [Zhang (1998)] P8 02-236-V6 B NSI R5 CCR5 [Zhang (1998)] P9 22-236-V6 B NSI R5 CCR5 [Zhang (1998)] P10 03-237-V6 B NSI R5 CCR5 [Zhang (1998)] P11 22-202-V3 B NSI R5 CCR5 [Zhang (1998)] P13 19-242-V3 B NSI R5 CCR5 [Zhang (1998)] P14 22-207-V6 B NSI R5 CCR5 [Zhang (1998)] P15 22-237-V4 B NSI R5 CCR5 [Zhang (1998)] 301657 B NSI R5 CCR5 [Zhang (1998)] 301714 B NSI R5 CCR5 [Zhang (1998)] 301073 B NSI R5 CCR5 [Zhang (1998)] 301056 B NSI R5 CCR5 [Zhang (1998)] 301660 B NSI R5 CCR5 [Zhang (1998)] 301727 B NSI R5 CCR5 [Zhang (1998)] MWB B NSI R5 CCR5 [Zhang (1998)] 301593 B SI R5X4 CXCR4,CCR5 V28 [Zhang (1998)] AD73 B SI R5X4 CXCR4,CCR5 V28 [Zhang (1998)] DH123 B SI R5X4 CXCR4,CCR5 CCR8 [Zhang (1998)] NL4-3 B SI R4 CXCR4 [Zhang (1998)] P6 02-217-V3-m1 B NSI R5 CCR5 BONZO [Zhang (1998)] P6 02-217-V4 m2 B NSI R5 CCR5 BONZO [Zhang (1998)] P6-02-217-V6-m6 B NSI R5 CCR5 BONZO [Zhang (1998)] P6 02-217-V8-m12 B NSI R5 CCR5 BONZO [Zhang (1998)] P6 02-217-V9-m18 B NSI R5 CCR5 [Zhang (1998)] M6-V2-m4b B SI R5X4 CXCR4,CCR5 BONZO,V28, [Zhang (1998)] APJ M6-V3-m1b B SI R5X4 CXCR4,CCR5 BONZO,V28, [Zhang (1998)] APJ M6-V6-m6a B SI R5X4 CXCR4,CCR5 BONZO,V28, [Zhang (1998)] APJ EL1 B SI X4 CXCR4 APJ,CCR3 [Choe (1998)] 2005 B SI X4 CXCR4 [Simmons (1998)] 2044 B SI X4 CXCR4 [Simmons (1998)] 2028 B SI R5X4 CXCR4,CCR5 [Simmons (1998)] 2076 B SI R5X4 CXCR4,CCR5 CCR3,CCR8, [Simmons (1998)] STRL33 SL-2 B NSI R5 CCR5 GPR15 [Simmons (1998)] 92ZW101 NA C NSI R5 CCR5 [Zhang (1996)] 92BR025.9 U52953 C NSI R5 CCR5 [Bjorndal (1997), Dittmar (1997)] BR28 U16217 C NSI R5 CCR5 [Dittmar (1997)] 93MW965.26 U08455 C NSI R5 CCR5 [Bjorndal (1997)] BR70 NA C NSI R5 CCR5 [Dittmar (1997)] JW1 NA C NSI R5 CCR5 [Dittmar (1997)] JW4 NA C NSI R5 CCR5 [Dittmar (1997)] 92ZW102 NA C NSI R5 CCR5 [Zhang (1996)] DJ259 L22940 C NSI R5 CCR5 [Trkola (1998)] 94ZW103 NA C NSI R5 CCR5 [Trkola (1998)] 94ZW109 NA C NSI R5 CCR5 [Trkola (1998)] 92ZW106 NA C SI X4 CXCR4 [Zhang (1996)] ZAM20 L22956 C SI X4 CXCR4 [Trkola (1998)] 94ZW106 NA C SI X4 CXCR4 [Trkola (1998)] 94KE102 NA D NSI R5 CCR5 [Trkola (1998),Zhang (1996)] 94KE103 NA D NSI R5 CCR5 [Trkola (1998),Zhang (1996)] 92UG046 U08737 D SI X4 CXCR4 [Trkola (1998)] UG270 NA D SI X4 CXCR4 [Trkola (1998)] 92UG024.2 U08726 D SI X4 CXCR4 CCR8*,V28*, [Bjorndal (1997), Rucker (1997),Trkola (19980] CCR3,f 92UG021.16 U27399 D SI X4 CXCR4 [Bjorndal (1997)] JW5 NA D SI X4 CXCR4 [Dittmar (1997)] NDK M27323 D SI X4 CXCR4 GPR15 [Deng (1997),Pleskoff (1997)] 93ZR001.3 U27419 D NA X4 CXCR4 V28* [Rucker (1997)] 91ZR001.3 D SI R5X4 CXCR4,CCR5 [Bazan (1998)] SE7076 D SI X4 CXCR4 [Vallejo (1998)] SE8384 D SI X4 CXCR4 [Vallejo (1998)] SE8646 D SI R5X4 CCR5,CXCR4 [Vallejo (1998)] CM235 NA E NSI R5 CCR5 [Trkola (1998)] 92TH001 NA E NSI R5 CCR5 [Trkola (1998)] M53 NA E NSI R5 CCR5 [Dittmar (1997)] 92TH22 U09131 E NSI R5 CCR5 [Dittmar (1997)] 92TH23 NA E NSI R5 CCR5 [Dittmar (1997)] C2 NA E NSI R5 CCR5 [Dittmar (1997)] 93TH305 NA E NSI R5 CCR5 [Zhang (1996)] 93TH307 NA E NSI R5 CCR5 [Zhang (1996)] 93TH966.8 U08456 E NSI R5 CCR5 [Bjorndal (1997)] 93TH976.17 U08458 E NA R5 CCR5 [Bjorndal (1997)] 93TH304 NA E SI R5X4 CXCR4,CCR5 [Zhang (1996)] SL6 NA E SI X4 CXCR4 [Dittmar (1997)] SL7 NA E SI X4 CXCR4 [Dittmar (1997)] SL8 NA E SI X4 CXCR4 [Dittmar (1997)] 94TH304 NA E SI X4 CXCR4 [Trkola (1998)] BR58 NA F SI R5X4 CXCR4,CCR5 CCR3 [Dittmar (1997)] BZ162 L22084 F NSI R5 CCR5 [Trkola (1998)] R1 NA F NSI R5 CCR5 [Trkola (1998)] 93BR029.2 U27413 F NA R5 CCR5 [Rucker (1997)] RU23 F NSI R5 CCR5 [Vallejo (1998)] RU29 F NSI R5 CCR5 [Vallejo (1998)] RU30 F NSI R5 CCR5 [Vallejo (1998)] 92UG975.10 U27426 G NSI R5 CCR5 [Bjorndal (1997)] CA9 NA O NSI R5 CCR5 [Zhang (1996)] MVP5180 L20571 O SI R5X4 CXCR4,CCR5 [Zhang (1996)] ES1158.1 O NSI R5 CCR5 [Vallejo (1998)] ES1159.1 O NSI R5 CCR5 [Vallejo (1998)] MD.1 O NSI R5 CCR5 [Vallejo (1998)] MVP-5180 O SI R5X4 CXCR4,CCR5 [Vallejo (1998)]

Footnotes:

a Accession numbers refer to sequences from the described isolate that contain the longest available sequence including Env (full length genome when available, down to a minimum of the V3 region), but may not pertain directly to the sample used to determine the isolate's phenotype. Accession numbers have been deemed "NA" if there are no related locus names in the HIV Database, or if there is insufficient information to conclude that a similarly named locus in the database is referring to the same isolate.

b The tropism for each virus strain is indicated. SI = syncytium inducing; NSI = non syncytium inducing; SI/TCLA = syncytium inducing T-cell line adapted; NA = not available.

c As described in the text, we have proposed that viruses which use CCR5 be termed R5 viruses, viruses that use CXCR4 be termed X4 viruses, and viruses that use both be termed R5X4 viruses. For a virus to be termed an R5X4 virus, CXCR4 must be used at >10% of the efficiency of CCR5 for viruses that use CCR5 as their primary receptor, or CCR5 must be used at >10% of the efficiency of CXCR4 for viruses that use CXCR4 as their primary receptor.

d All virus strains in the Table have been tested for the ability to use CCR5 and CXCR4. The receptors used by each strain are indicated. For both receptors to be listed, the least efficiently used receptor must support virus entry by >10% of the levels supported by the most efficiently used receptor. For viruses that use both receptors, CXCR4 is listed first; this does not mean that CXCR4 is used more efficiently than CCR5.

e Receptors other than CCR5 or CXCR4 that are used by the indicated strains are listed. Since most viruses have not yet been tested for the ability to use receptors other than CCR5 and CXCR4, the absence of other receptors used in the Table should not be taken to mean that a virus uses only CCR5 or CXCR4. For example, if CCR3 is not listed as being used by a given virus strain, it may be because it has not yet been tested. Receptors that have been shown by experiments not to be used by a given virus strain are not indicated. Occasionally there are discrepancies in the literature concerning the use of one or more receptors. In these cases, we opted for `majority rules'. If two papers report a positive result and one a negative result, the receptor is shown as being used.

f The virus isolate used CCR3 whereas the cloned env from this isolate did not [Bjorndal (1997)].

*Indicates that coreceptor use was determined only by fusion assays rather than by virus infection.

g 92HT593.1 (cloned env gene) and 92HA593 (whole virus) come from the same isolate and gave slightly different results in references [Dittmar (1997)] and [Zhang (1996)] .

HIV-2

HIV-2 strains appear to be remarkably promiscuous in their use of coreceptors, with receptors other than CCR5 or CXCR4 often supporting very efficient virus infection in transfected cells. This is obvious from the table, where coreceptors that are used efficiently are listed separately from those that support infection inefficiently. In addition, whenever possible, information is provided about the virus (whether it is a molecular clone, biological clone, or virus swarm, and whether it can infect PBMCs from CCR5-negative individuals) and the clinical stage of the patient from which each virus was isolated. See the key at the end of the table for an explanation of abbreviations.

                               (delta)
                    Primary   32 PBMCs    Patient                        Weak 
Isolate     Status  Isolate  Replication   Status   Coreceptors       Coreceptors        References 


ST MC NO + ASYMP CCR5/GPR15- CCR1/- [Edinger(1997b), STRL33(GPR1/APj) CCR2b/- Edinger (1998a), CCR3/- Edinger(1998b), CXCR4 Rucker(1997), Hill (1997), McKnight (1998), Deng (1997)] ST/24.1C#2 MC CCR5/GPR15- [Rucker (1997), STRL33 Hill (1997), Hill (1998),Deng (1997)] SBL6669 MC SYMP (CCR5/CXCR4/- [Rucker (1997), CCR8/V28 Chakrabarti (1990), Albert (1987)] ROD/A MC AIDS (CCR5/CXCR4/- [Hill (1997), CCR3/US28/V28 Sol (1998), Sol (1997), Clavel (1986), Bron (1997),Reeves (1997)] ROD/B MC NO CXCR4 CCR3/V28 [Clapham (1992), Reeves (1997)] prCBL-20 MC YES + AIDS CXCR4 CCR1/- [McKnight (1998), CCR2b/- Schulz (1990)] CCR3/CCR5 CBL-20 MC NO + CXCR4 [McKnight (1998)] prCBL-23 MC YES SYMP CXCR4 CCR1/- [McKnight (1998), CCR2b/- Schulz (1990)] CCR3/CCR5 UC1 MC SYMP CCR5 [Hill (1997), Evans (1988), Deng (1997)] UC2 MC NO AIDS CCR5/CXCR4/- CCR2b/CCR3 [Hill (1997), STRL33/GPR15 Evans (1988), Barnett (1996), Deng (1997)] MIR MC + AIDS CXCR4 CCR1/- [Sol (1997), CCR2b/- McKnight (1998), CCR3/CCR5 Clavel (1986)] V9 AIDS CCR1/CCR2b/- [McKnight (1998)] CCR3/CCR5/CXCR4 A-ND + SYMP CXCR4 CCR1/- [McKnight (1998)] CCR2b/CCR3 VCP MC NO CXCR4 [Endres (1996)] F0784 MC CCR5/GPR15 [Deng (1997),Gao (1992)] A195811 YES - AIDS CCR5/GPR15/- [Chen (1998a),Owen (1998)] STRL33 A227011 YES - ASYMP CCR5/GPR15/- [Chen (1998a)] STRL33 G0415k YES CCR5/GPR15/- [Chen (1998a)] STRL33 7924A YES + AIDS CXCR4/CCR2b/- [Chen (1998a)] CCR3/CCR5/- STRL33/GPR15 JK7312A MC + SYMP CCR5/GPR15/- [Hill (1997), CCR4/CXCR4 Owen (1998),Gao (1992), Deng (1997)] 310342 S YES CCR5/CXCR4/- [Owen (1998)] STRL33/CCR1 GB87 S YES + SYMP CCR5/GPR15/- [Owen (1998)] CXCR4/STRL33/- CCR4/CCR2b/CCR1 310248 S YES + CCR1/CCR4/CCR5/- [Owen (1998)] CXCR4 310319 S YES + CCR2b/CCR3/- [Owen (1998)] CCR4/CXCR4/- STRL33/GPR15/- CCR5 77618 S YES + AIDS CCR2b/CCR3/- [Owen (1998)] CXCR4/GPR15/- STRL33/CCR5 GB122 S YES + CCR2b/CCR3/- CCR5/- [Owen (1998)] GPR15 CXCR4 A2267 S YES - ASYMP CCR5 [Owen (1998)] SLRHC S YES - ASYMP CCR5 [Owen (1998)] 310072 S YES - DONOR CCR5 [Owen (1998)] 310340 S YES - DONOR CCR5 [Owen (1998)] 60415k S YES - ASYMP CCR5 [Owen (1998)] BAJE (B) S YES - ASYMP CCR5/CCR3 [Sol (1998)] BATI (A) S YES - ASYMP CCR5 CCR3 [Sol (1997)] BAPA (C) S YES + ASYMP CCR5 CCR3 [Sol (1997)] DESY (E) S YES ASYMP CCR5 CCR3 [Sol (1997)] BAYO (G) S YES + AIDS CXCR4 [Sol (1997)] B3 S YES CCR5 [Heredia (1997)] B4 S YES CCR5 [Heredia (1997)] B7 S YES CCR5 [Heredia (1997)] B8 S YES CCR5 [Heredia (1997)] B9 S YES CCR5 [Heredia (1997)] B2 S YES CCR1/CCR2b/- [Heredia (1997)] CCR5/CXCR4 B5 S YES CCR1/CCR2b/- [Heredia (1997)] CCR5/CXCR4 B10 S YES CCR1/CCR2b/- [Heredia (1997)] CCR5/CXCR4 RH-2-1 S YES EARLY CCR5 [Guillon (1998)] PH-2-1 C1 BC YES INTERMED CCR5 [Guillon (1998)] PH-2-1 C12 BC YES INTERMED CCR5/CCR1/CCR3 [Guillon (1998)] PH-2-1 E6 BC YES INTERMED CCR5/CXCR4/- [Guillon (1998)] CCR3/CCR1 PH-2-1 H8 BC YES INTERMED CCR5/CCR3/CCR1 [Guillon (1998)] PH-2-1 D5 BC YES INTERMED CCR5 CCR3/- [Guillon (1998)] CXCR4/- CCR1 PH-2-1 H12 BC YES INTERMED CCR5/CCR3/CCR1 CXCR4 [Guillon (1998)] RH-2-5 A10 BC YES INTERMED CCR5 CCR3 [Guillon (1998)] RH-2-5 E4 BC YES INTERMED CCR5/CCR3/CCR1 CXCR4 [Guillon (1998)] RH-2-5 E11 BC YES INTERMED CCR5/CCR3/CCR1 CXCR4 [Guillon (1998)] RH-2-5 F7 BC YES INTERMED CCR5/CCR3/CCR1/- [Guillon (1998)] CXCR4 RH-2-5 G7 BC YES INTERMED CCR5 CXCR4/- [Guillon (1998)] CCR1/CCR3 RH-2-7 A5 BC YES INTERMED CCR5/CCR3 CCR1/- [Guillon (1998)] CXCR4 RH-2-7 C9 BC YES INTERMED CCR5/CCR1/CCR3/- [Guillon (1998)] CCR2b/CXCR4 RH-2-7 C12 BC YES INTERMED CCR5 CCR3/- [Guillon (1998)] CXCR4/- CCR1 RH-2-7 D7 BC YES INTERMED CCR5/CCR3/CCR1 CXCR4/- [Guillon (1998)] CCR2b RH-2-7 G12 BC YES INTERMED CCR5/CCR3/CCR1 CXCR4/- [Guillon (1998)] CCR2b RH-2-2 S YES LATE CCR3/CCR1/- CCR2b [Guillon (1998)] CXCR4/CCR5 RH-2-6 S YES LATE CXCR4 [Guillon (1998)]
KEY FOR HIV-2 CORECEPTOR USE TABLE MC = molecular clone BC = biologic clone S = virus swarm ASYMP = asymptomatic SYMP = symptomatic DONOR = blood donor EARLY = isolated at early timepoints INTERMED = isolated at an intermediate point in disease progression LATE = isolated late in disease V28 has been redesignated CX3CR1 ( ) indicates results are from cell-cell fusion assays, all other data are from infection assays

SIV

SIV strains are also rather promiscuous in their use of coreceptors. SIV strains use CCR5 with few exceptions, while CXCR4 is only rarely used, even by laboratory-adapted viruses, a feature that distinguishes them from HIV-1 strains. The Table indicates the tropism of the virus (when known) as well as whether the data were derived from a molecular clone or virus swarm.

Strain              Status        Tropism       Coreceptors             References 

SIVmac251 S M CCR5/GPR15/STRL33 [Chen (1997),Chen (1998a), Kuhmann (1997),Luciw (1992)] SIVmac251(BK28 clone) MC CCR5/GPR15/STRL33/- [Edinger (1997b),Edinger (1999)] GPR1 SIVmac251(v194 clone) MC (CCR5/STRL33/GPR1/- [Rucker (1997),Edinger (1997a), GPR15) Edinger (1997b),Koenig (1989)] SIVmac 239 MC T CCR5/DTRL33/GPR15/- [Edinger (1997a),Edinger (1997b), GPR1(ChemR23) Edinger (1999),Choe (1998), Rucker (1997),Chen (1997),Chen (1998a), Hill (1997),Hill (1998),Kirchhoff (1997), Marcon (1997),Samson (1998), Farzan (1997),Deng (1997)] SIVmac316 MC M CCR5/STRL33/GPR15/- Edinger (1997a),Edinger (1997b), APJ/CCR8/GPR1/- Choe (1998),Rucker (1997), (ChemR23) Kirchhoff (1997),Samson (1998), Farzan (1997)] SIVmac316mut MC Md CCR5/(STRL33/GPR1/- [Edinger (1997a),Edinger (1997b), GPR15/CHEMR23/APJ) Edinger (1998b),Rucker (1997), Kirchhoff (1997),Kirchhoff (1994), Samson (1998)] SIVmac316EM MC M CCR5/GPR15/STRL33 [Chen (1997),Chen (1998a)] SIVmac1A11 MC M CCR5/STRL33/GPR15 Edinger (1997b),Edinger (1999), Chen (1997),Chen (1998a), Hill (1997),Banapour (1991a), Banapour (1991b),Deng (1997), Chen (1998a),Chen (1998b)] SIVmac/17E-Fr MC M CCR5/STRL33/GPR13/- [Edinger (1997a),Edinger (1997b), APJ/(ChemR23/CCR8) Edinger (1998b),Edinger(1999), Rucker (1997),Flaherty (1997)] SIVmac/17E-Cl MC M (CCR5/GPR1/STRL33/- [Edinger (1998a),Flaherty (1997)] GPR15) SIVmacCP-MAC S CXCR4 [Endres (1996)] SIVmacCP-MAC MC CCR5/STRL33/GPR15/APJ [Edinger (1997b),Edinger (1998b), Edinger (1999)] SIVmac(mne) CCR5/GPR15 [Chen (1998a)] SIVsm(delta)B670 clone 3 MC CCR5/GPR15/STRL33 [Edinger (1997a),Edinger (1997b), Rucker (1997)] SIVsm(delta)B670 clone 12 MC (M) CCR5/GPR15 [Edinger (1997b),Edinger (1999), Amedee (1995)] SIVsm62A MC T (CCR5/GPR1/GPR15/STRL33/ChemR23) [Edinger (1997b),Hirsch (1994)] SIVsm62B MC (CCR5) [Edinger (1997b),Hirsch (1994)] SIVsm62D MC M CCR5 [Edinger (1997b),Hirsch (1994)] SIVsmE543-3 MC M CCR5/GPR15 [Edinger (1997b),Hirsch (1997)] SIVsmE543/B10 MC (CCR5/GPR15/GPR1/STRL33) [Edinger (1997b),Hirsch (1997)] SIVsmPBj6.6 MC (STRL33/CCR5/GPR1/GPR15) [Edinger (1998a),Novembre (1993)] SIVsmSL92a S CCR5/GPR15/STRL33 [Chen (1997),Chen(1998a)] SIVsmSL92b S CCR5/GPR15/STRL33 [Chen (1998a)] SIVsmLib-1 S CCR5/GPR15/STRL33 [Chen (1997),Chen (1998a)] SIVsmFNS S CCR5/GPR15/STRL33 [Chen (1998a)] SIVagmSab1.4 MC (CCR5/GPR15/GPR1/STRL33) [Edinger (1997b)] SIVagm9063-2 MC M (CCR5) [Edinger (1997b),Hirsch (1995)] SIVagmTYO MC CCR5/GPR15/STRL33 [Deng (1997),Chen (1998a),Chen (1998b)] SIVmneCl8 MC CCR5 [Chackerian (1997)] SIVrcm CCR2b/STRL33 [Marx (1998)] SIVcpzGAB S CCR5 [Chen (1997),Marx (1998)] SIVmnd(GB-1) S CXCR4 [Schols (1998)]
KEY FOR SIV CORECEPTOR TABLE S = uncloned virus swarm MC = molecular clone M = macrophage tropic, Md = delayed infection kinetics in macrophages, (M) = probably M-tropic, T = T-tropic ( ) indicate fusion results, all other data from infection assays

References

[Albert (1987)] J. Albert, U. Bredberg, F. Chiodi, B. Bottiger, E. M. Fenyo, E. Norrby, G. Biberfeld.
   A new human retrovirus isolate of West African origin (SBL-6669) and its relationship to HTLV-IV,
   LAV-II, and HTLV-IIIB. AIDS Res Hum Retroviruses 3:3-10, 1987.
   
   NOTE:   (Medline: 87299193).

[Alkhatib (1996)] G. Alkhatib, C. Combadiere, C. C. Broder, Y. Feng, P. E. Kennedy, P. M. Murphy, 
   E. A. Berger. CC CKR5: a RANTES, MIP-1alpha, MIP-1beta receptor as a fusion cofactor for     
   macrophage-tropic  HIV-1. Science 272:1955-8, 1996.
    
   NOTE:   (Medline: 96275693).

[Amedee (1995)] A. M. Amedee, N. Lacour, J. L. Gierman, L. N. Martin, J. E. Clements, R. Bohm, 
   R. M. Harrison, M. Murphey-Corb. Genotypic selection of simian immunodeficiency virus in macaque
   infants infected transplacentally. J Virol 69:7982-90, 1995.
    
   NOTE:   (Medline: 96079048).

[Banapour (1991a)] B. Banapour, M. L. Marthas, R. J. Munn, P. A. Luciw.  In vitro macrophage tropism of
   pathogenic and nonpathogenic molecular clones of simian immunodeficiency virus (SIVmac).    
   Virology 183:12-9, 1991a.
    
   NOTE:   (Medline: 91272471).

[Banapour (1991b)] B. Banapour, M. L. Marthas, R. A. Ramos, B. L. Lohman, R. E. Unger, M. B. Gardner, 
   N. C. Pedersen, P. A. Luciw. Identification of viral determinants of macrophage tropism for simian   
   immunodeficiency virus SIVmac. J Virol 65:5798-805, 1991b.

   NOTE:   (Medline: 92015471).

[Barnett  (1996)] S. W. Barnett, H. S. Legg, Y. Sun, J. Klinger, D.J. Blackbourn, C. P. Locher, J. A. Levy.
   Molecular cloning of the human immunodeficiency virus subtype 2 strain HIV-2UC2. 
   Virology 222:257-61, 1996.

   NOTE:   (Medline: 96400130).

[Bazan (1998)] H. A. Bazan, G. Alkhatib, C. C. Broder, E. A. Berger. Patterns of CCR5, CXCR4, and CCR3   
   usage by envelope glycoproteins from human immunodeficiency virus type 1 primary isolates. 
   J Virol 72:4485-91, 1998.
    
   NOTE:   (Medline: 98216824).

[Bjorndal (1997)] A. Bjorndal, H. Deng, M. Jansson, J. R. Fiore, C. Colognesi, A. Karlsson, J. Albert,   
   G. Scarlatti, D. R. Littman, E. M. Fenyo. Coreceptor usage of primary human immunodeficiency virus type
   1 isolates varies according to biological phenotype.  J Virol 71:7478-87, 1997.

   NOTE:   (Medline: 97456513).

[Bron (1997)] R. Bron, P. J. Klasse, D. Wilkinson, P. R. Clapham, A. Pelchen-Matthews, C. Power, T. N.    
   Wells, J. Kim, S. C. Peiper, J. A. Hoxie, M. Marsh. Promiscuous use of CC and CXC chemokine receptors in
   cell-to-cell fusion mediated by a human immunodeficiency virus type 2 envelope protein. 
   J Virol 71:8405-15, 1997.
    
   NOTE:   (Medline: 98001362).

[Chackerian  (1997)] B. Chackerian, E. M. Long, P. A. Luciw, J. Overbaugh. Human immunodeficiency virus
   type 1 coreceptors participate in postentry stages in the virus replication cycle and function in simian
   immunodeficiency virus infection. J Virol 71:3932-9, 1997.
    
   NOTE:   (Medline:  97248442).

[Chakrabarti  (1990)] S. Chakrabarti, T. Mizukami, G. Franchini, B. Moss. Synthesis, oligomerization, and
   biological activity of the human immunodeficiency virus type 2 envelope glycoprotein expressed by a
   recombinant vaccinia virus. Virology 178:134-42, 1990.
    
   NOTE:   (Medline: 90357756).

[Chen (1998a)] Z. Chen, A. Gettie, D. D. Ho, P. A. Marx. Primary SIVsm isolates use the CCR5 coreceptor
   from sooty mangabeys naturally infected in west Africa: a comparison of coreceptor usage of primary 
   SIVsm, HIV-2, and SIVmac. Virology 246:113-24, 1998a.
    
   NOTE:   (Medline: 98321155).

[Chen (1998b)] Z. Chen, D. Kwon, Z. Jin, S. Monard, P. Telfer, M. S. Jones, C.Y. Lu, R. F. Aguilar, 
   D. D. Ho, P. A. Marx. Natural infection of a homozygous delta24 CCR5 red-capped mangabey with an 
   R2b-tropic simian immunodeficiency virus. J Exp Med 188:2057-65, 1998b.
    
   NOTE:   (Medline: 99059829).

[Chen (1997)] Z. Chen, P. Zhou, D. D. Ho, N. R. Landau, P. A. Marx. Genetically divergent strains of simian
   immunodeficiency virus use CCR5 as a coreceptor for entry. J Virol 71:2705-14,
   1997.
    
   NOTE:   (Medline: 97213934).

[Choe (1998)] H. Choe, M. Farzan, M. Konkel, K. Martin, Y. Sun, L. Marcon, M. Cayabyab, M. Berman, 
   M. E. Dorf, N. Gerard, C. Gerard, J. Sodroski. The orphan seven-transmembrane receptor apj supports the
   entry of primary T-cell-line-tropic and dualtropic human immunodeficiency virus type 1. 
   J Virol 72:6113-8, 1998.
   
   NOTE:   (Medline: 98285773).

[Choe (1996)] H. Choe, M. Farzan, Y. Sun, N. Sullivan, B. Rollins, P. D. Ponath, L. Wu, C. R. Mackay, 
   G. LaRosa, W. Newman, N. Gerard, C. Gerard, J. Sodroski. The beta-chemokine receptors CCR3 and CCR5 
   facilitate infection by primary  HIV-1  isolates. Cell 85:1135-48, 1996.
    
   NOTE:   (Medline: 96270515).

[Clapham (1992)] P. R. Clapham, A. McKnight, R. A. Weiss. Human immunodeficiency virus type 2 infection and
   fusion of CD4- negative human cell lines: induction and enhancement by soluble CD4. 
    J Virol 66:3531-7, 1992.
    
   NOTE:   (Medline: 92260623).

[Clavel (1986)] F. Clavel, D. Guetard, F. Brun-Vezinet, S. Chamaret, M. A. Rey, M. O. Santos-Ferreira,   
   A. G. Laurent, C. Dauguet, C. Katlama, C. Rouzioux. Isolation of a new human retrovirus from West
   African patients with AIDS. Science 233:343-6, 1986.
    
   NOTE:   (Medline:  86261773).

[Deng (1997)] H. K. Deng, D. Unutmaz, V. N. KewalRamani, D. R. Littman. Expression cloning of new receptors
   used by simian and human immunodeficiency viruses (see comments). 
   Nature 388:296-300, 1997.
    
   NOTE:   (Medline: 97373958).

[Dittmar (1997)] M. T. Dittmar, G. Simmons, S. Hibbitts, M. O'Hare, S. Louisirirotchanakul, S. Beddows,    
   J. Weber, P. R. Clapham, R. A. Weiss. Langerhans cell tropism of human immunodeficiency virus type 1
   subtype A through F isolates derived from different transmission groups.
   J Virol 71:8008-13, 1997.

   NOTE:   (Medline: 97456582).

[Dragic (1996)] T. Dragic, V. Litwin, G. P. Allaway, S. R. Martin, Y. Huang, K. A. Nagashima, C. Cayanan, 
   P. J. Maddon, R. A. Koup, J. P. Moore, W. A. Paxton. HIV-1  entry into  CD4+  cells is mediated by the 
   chemokine receptor CC- CKR-5 (see comments). Nature 381:667-73, 1996.

   NOTE:   (Medline:  96260018).

[Edinger (1997a)] A. L. Edinger, A. Amedee, K. Miller, B. J. Doranz, M. Endres, M. Sharron, M. Samson, 
   Z. H. Lu, J. E. Clements, M. Murphey-Corb, S. C. Peiper, M. Parmentier, C. C. Broder, R. W. Doms.
   Differential utilization of CCR5 by macrophage and T cell tropic simian immunodeficiency virus strains. 
   Proc Natl Acad Sci U S A 94:4005-10, 1997a.

   NOTE:   (Medline: 97268687).

[Edinger (1999)] A. L. Edinger, C. Blanpain, K. Kunstman, S. M. Wolinsky, M. Endres, M. Parmentier,  
   R.~W. Doms. Functional dissection of CCR5 coreceptor function through the use of CD4-independent SIV 
   strains. 1999.
    
   NOTE:   Submitted.

[Edinger (1998a)] A. L. Edinger, T. L. Hoffman, M. Sharron, B. Lee, B. O'Dowd, R. W. Doms. Use of GPR1, 
   GPR15, and STRL33 as coreceptors by diverse human immunodeficiency virus type 1 and simian 
   immunodeficiency virus envelope proteins. Virology 249:367-78, 1998a.

   NOTE:   (Medline:  99009390).

[Edinger (1998b)] A. L. Edinger, T. L. Hoffman, M. Sharron, B. Lee, Y. Yi, W. Choe, D. L. Kolson, 
   B. Mitrovic, Y. Zhou, D. Faulds, R. G. Collman, J. Hesselgesser, R. Horuk, R. W. Doms. An orphan 
   seven-transmembrane domain receptor expressed widely in the brain functions as a coreceptor for human 
   immunodeficiency virus type 1 and simian immunodeficiency virus. 
   J Virol 72:7934-40, 1998b.

   NOTE:   (Medline: 98406195).

[Edinger (1997b)] A. L. Edinger, J. L. Mankowski, B. J. Doranz, B. J. Margulies, B. Lee, J. Rucker, 
   M. Sharron , T. L. Hoffman, J. F. Berson, M. C. Zink, V. M. Hirsch, J. E. Clements, R. W. Doms.
   CD4-independent, CCR5-dependent infection of brain capillary endothelial cells by a neurovirulent simian
   immunodeficiency virus strain. Proc Natl Acad Sci U S A 94:14742-7, 1997b.
    
   NOTE:   (Medline:  98070822).

[Endres (1996)] M. J. Endres, P. R. Clapham, M. Marsh, M. Ahuja, J. D. Turner, A. McKnight, J. F. Thomas, 
    B. Stoebenau-Haggarty, S. Choe, P. J. Vance, T. N. Wells, C. A. Power, S. S. Sutterwala, R. W. Doms, 
    N. R. Landau, J. A. Hoxie. CD4-independent  infection by  HIV-2  is mediated by  fusin/CXCR4. 
    Cell 87:745-56, 1996.
    
    NOTE:   (Medline:  97083584).

[Evans  (1988)] L. A. Evans, J. Moreau, K. Odehouri, D. Seto, G. Thomson-Honnebier, H. Legg, A. Barboza, 
    C. Cheng-Mayer, J. A. Levy. Simultaneous isolation of HIV-1 and HIV-2 from an AIDS patient. 
    Lancet 2:1389-91, 1988.

    NOTE:   (Medline:  89069975).

[Farzan (1997)] M. Farzan, H. Choe, K. Martin, L. Marcon, W. Hofmann, G. Karlsson, Y. Sun, P. Barrett, 
    N. Marchand, N. Sullivan, N. Gerard, C. Gerard, J. Sodroski. Two orphan seven-transmembrane segment 
    receptors which are expressed in CD4-positive cells support simian immunodeficiency virus infection.
    J Exp Med. 186:405-11, 1997.

    NOTE:   (Medline:  98105758).

[Flaherty (1997)] M. T. Flaherty, D. A. Hauer, J. L. Mankowski, M. C. Zink, J. E. Clements. Molecular and 
    biological characterization of a neurovirulent molecular clone of simian immunodeficiency virus. 
    J Virol 71:5790-8, 1997.
    
    NOTE:   (Medline: 97366637).

[Gao (1992)] F. Gao, L. Yue, A. T. White, P. G. Pappas, J. Barchue, A. P. Hanson, B. M. Greene, 
    P. M. Sharp, G. M. Shaw, B. H. Hahn. Human infection by genetically diverse SIVSM-related HIV-2 in west
    Africa. Nature 358:495-9, 1992.
    
    NOTE:   (Medline:  92350299).

[Guillon (1998)] C. Guillon, M. E. van der Ende, P. H. Boers, R. A. Gruters, M. Schutten, A. D. Osterhaus.
    Coreceptor usage of human immunodeficiency virus type 2 primary isolates and biological clones is broad
    and does not correlate with their syncytium-inducing capacities. 
    J Virol 72:6260-3, 1998.
    
    NOTE:   (Medline:  98285800).

[Heredia (1997)] A. Heredia, A. Vallejo, V. Soriano, J. S. Epstein, I. K. Hewlett. Chemokine receptors and 
    HIV-2 [letter]. AIDS 11:1198-9, 1997.

    NOTE:   (Medline:  97377877).

[Hill (1997)] C. M. Hill, H. Deng, D. Unutmaz, V. N. Kewalramani, L. Bastiani, M. K. Gorny, S.  
    Zolla-Pazner, D. R. Littman. Envelope glycoproteins from human immunodeficiency virus types 1 and
    2 and simian immunodeficiency virus can use human CCR5 as a coreceptor for viral entry and make direct 
    CD4-dependent interactions with this chemokine receptor.
    J Virol 71:6296-304, 1997.
    
    NOTE:   (Medline:  97404634).

[Hill (1998)] C. M. Hill, D. Kwon, M. Jones, C. B. Davis, S. Marmon, B. L. Daugherty, J. A. DeMartino, 
    M. S. Springer, D. Unutmaz, D.R. Littman. The amino terminus of human CCR5 is required for its function
    as a receptor for diverse human and simian immunodeficiency virus envelope
    glycoproteins. Virology 248:357-71, 1998.

    NOTE:   (Medline:  98389855).

[Hirsch (1997)] V. Hirsch, D. Adger-Johnson, B. Campbell, S. Goldstein, C. Brown, W. R. Elkins, 
    D. C. Montefiori. A molecularly cloned, pathogenic, neutralization-resistant simian immunodeficiency 
    virus, SIVsmE543-3. J Virol 71:1608-20, 1997.

    NOTE:   (Medline:  97151152).

[Hirsch (1995)] V. M. Hirsch, G. Dapolito, P. R. Johnson, W. R. Elkins, W. T. London, R. J. Montali, 
    S. Goldstein, C. Brown. Induction of AIDS by simian immunodeficiency virus from an African
    green monkey: species-specific variation in pathogenicity correlates with the extent of in vivo 
    replication. J Virol 69:955-67, 1995.

    NOTE:   (Medline:  95115151).

[Hirsch (1994)] V. M. Hirsch, J. E. Martin, G. Dapolito, W. R. Elkins, W. T. London, S. Goldstein,
    P. R. Johnson. Spontaneous substitutions in the vicinity of the V3 analog affect cell tropism and
    pathogenicity of simian immunodeficiency virus. J Virol 68:2649-61, 1994.

    NOTE:   (Medline:  94187106).

[Kirchhoff (1994)] F. Kirchhoff, K. Mori, R. C. Desrosiers. The "V3" domain is a determinant of simian 
    immunodeficiency virus cell tropism. J Virol 68:3682-92, 1994.
 
    NOTE:   (Medline:  94246713).

[Kirchhoff (1997)] F. Kirchhoff, S. Pohlmann, M. Hamacher, R. E. Means, T. Kraus, K. Uberla, P. Di Marzio.
    Simian immunodeficiency virus variants with differential T-cell and macrophage tropism use CCR5 and an 
    unidentified cofactor expressed in CEMx174 cells for efficient entry. 
    J Virol 71:6509-16, 1997.

    NOTE:   (Medline:  97404658).

[Koenig (1989)] S. Koenig, V. M. Hirsch, R. A. Olmsted, D. Powell, W. Maury, A. Rabson, A. S. Fauci, 
   R. H. Purcell, P. R. Johnson. Selective infection of human CD4+ cells by simian immunodeficiency
   virus: productive infection associated with envelope glycoprotein- induced fusion. 
   Proc Natl Acad Sci U S A 86:2443-7, 1989.

   NOTE:   (Medline:  89184643).

[Kuhmann (1997)] S. E. Kuhmann, E. J. Platt, S. L. Kozak, D. Kabat. Polymorphisms in the CCR5 genes of 
   African green monkeys and mice implicate specific amino acids in infections by simian and human
   immunodeficiency viruses. J Virol 71:8642-56, 1997.
    
   NOTE:   (Medline:  98001387).

[Liao (1997)] F. Liao, G. Alkhatib, K. W. Peden, G. Sharma, E. A. Berger, J. M. Farber. STRL33, A novel 
   chemokine receptor-like protein, functions as a fusion cofactor for both macrophage-tropic and T cell
   line-tropic HIV-1. J Exp Med 185:2015-23, 1997.

   NOTE:   (Medline:  97311099).

[Luciw (1992)] P. A. Luciw, K. E. Shaw, R. E. Unger, V. Planelles, M. W. Stout, J. E. Lackner, 
   E. Pratt-Lowe, N. J. Leung, B. Banapour, M. L. Marthas. Genetic and biological comparisons of pathogenic
   and nonpathogenic molecular clones of simian immunodeficiency virus (SIVmac). 
   AIDS Res Hum Retroviruses 8:395-402, 1992.

   NOTE:   (Medline:  92239216).

[Marcon (1997)] L. Marcon, H. Choe, K. A. Martin, M. Farzan, P. D. Ponath, L. Wu, W. Newman, N. Gerard, 
   C. Gerard, J. Sodroski. Utilization of C-C chemokine receptor 5 by the envelope glycoproteins of a 
   pathogenic simian immunodeficiency virus, SIVmac239. 
   J Virol 71:2522-7, 1997.

   NOTE:   (Medline:  9784592).

[Marx & Chen (1998)] P. A. Marx & Z. Chen. The function of simian chemokine receptors in the replication of
   SIV. Semin Immunol 10:215-23, 1998.

   NOTE:   (Medline:  98317008).

[McKnight (1998)] A. McKnight, M. T. Dittmar, J. Moniz-Periera, K. Ariyoshi, J. D. Reeves, S. Hibbitts, 
   D. Whitby, E. Aarons, A. E. Proudfoot, H. Whittle, P. R. Clapham. A broad range of chemokine receptors 
   are used by primary isolates of human immunodeficiency virus type 2 as coreceptors with CD4. 
   J Virol 72:4065-71, 1998.

   NOTE:   (Medline:  98216773).

[Novembre (1993)] F. J. Novembre, P. R. Johnson, M. G. Lewis, D. C. Anderson, S. Klumpp, H. M. McClure, 
   V. M. Hirsch. Multiple viral determinants contribute to pathogenicity of the acutely lethal simian 
   immunodeficiency virus SIVsmmPBj variant. J Virol 67:2466-74, 1993.

   NOTE:   (Medline:  93233209).

[Owen (1998)] S. M. Owen, D. Ellenberger, M. Rayfield, S. Wiktor, P. Michel, M. H. Grieco, F. Gao, 
   B. H. Hahn, R. B. Lal. Genetically divergent strains of human immunodeficiency virus type 2 use multiple
   coreceptors for viral entry. J Virol 72:5425-32, 1998.

   NOTE:   (Medline:  98285695).

[Pleskoff (1997)] O. Pleskoff, N. Sol, B. Labrosse, M. Alizon. Human immunodeficiency virus strains differ 
   in their ability to infect CD4+ cells expressing the rat homolog of CXCR-4 (fusin).
   J Virol 71:3259-62, 1997.
 
   NOTE:   (Medline: 97214002).


[Reeves (1997)] J. D. Reeves, A. McKnight, S. Potempa, G. Simmons, P. W. Gray, C. A. Power, T. Wells, 
   R. A. Weiss, S. J. Talbot. CD4-independent  infection by  HIV-2 (ROD/B): use of the 7-transmembrane
   receptors CXCR-4, CCR-3, and V28 for entry. Virology 231:130-4, 1997.

   NOTE:   (Medline: 97288333).

[Rucker (1997)] J. Rucker, A. L. Edinger, M. Sharron, M. Samson, B. Lee, J. F. Berson, Y. Yi, B. Margulies,
   R. G. Collman, B. J. Doranz, M. Parmentier, R. W. Doms. Utilization of chemokine receptors, orphan
   receptors, and herpesvirus encoded receptors by diverse human and simian immunodeficiency viruses.
   J Virol 71:8999-9007, 1997.
 
   NOTE:   (Medline: 98037607).


[Samson (1998)] M. Samson, A. L. Edinger, P. Stordeur, J. Rucker, V. Verhasselt, M. Sharron, C. Govaerts,
   C. Mollereau, G. Vassart, R. W. Doms, M. Parmentier. ChemR23, a putative chemoattractant receptor, is 
   expressed in monocyte- derived dendritic cells and macrophages and is a coreceptor for SIV and some 
   primary HIV-1 strains. Eur J Immunol 28:1689-700, 1998.
    
   NOTE:   (Medline: 98264639).


[Schols & De Clercq (1998)] D. Schols & E. De Clercq. The simian immunodeficiency virus mnd(GB-1) strain
   uses CXCR4, not CCR5, as coreceptor for entry in human cells. 
   J Gen Virol 79( Pt 9):2203-5, 1998.
    
   NOTE:   (Medline: 98418501).


[Schulz (1990)] T. F. Schulz, D. Whitby, J. G. Hoad, T. Corrah, H. Whittle, R. A. Weiss.
   Biological and molecular variability of human immunodeficiency virus type 2 isolates from The Gambia. 
   J Virol 64:5177-82, 1990.
    
   NOTE:   (Medline: 90376478).


[Simmons (1998)] G. Simmons, J. D. Reeves, A. McKnight, N. Dejucq, S. Hibbitts, C. A. Power, E. Aarons, 
   D. Schols, E. De Clercq, A. E. Proudfoot, P. R. Clapham. CXCR4 as a functional coreceptor for human   
   immunodeficiency virus type 1 infection of primary macrophages. 
   J Virol 72:8453-7, 1998.
    
   NOTE:   (Medline: 98406265).


[Simmons (1996)] G. Simmons, D. Wilkinson, J. D. Reeves, M. T. Dittmar, S. Beddows, J. Weber,
   G. Carnegie, U. Desselberger, P. W. Gray, R. A. Weiss, P. R. Clapham. Primary, syncytium-inducing human
   immunodeficiency virus type 1 isolates are dual-tropic and most can use either Lestr or CCR5 coreceptors
   for virus entry. J Virol 70:8355-60, 1996.
 
   NOTE:   (Medline: 97126031).


[Sol (1997)] N. Sol, F. Ferchal, J. Braun, O. Pleskoff, C. Treboute, I. Ansart, M. Alizon. Usage of the
   coreceptors CCR-5, CCR-3, and CXCR-4 by primary and cell line-adapted human immunodeficiency virus type
   2. J Virol 71:8237-44, 1997.

   NOTE:   (Medline: 98001340).


[Sol (1998)] N. Sol, C. Treboute, E. Gomas, F. Ferchal, B. Shacklett, M. Alizon. The rhesus macaque CCR3
   chemokine receptor is a cell entry cofactor for HIV-2, but not for HIV-1. 
   Virology 240:213-20, 1998.
    
   NOTE:   (Medline: 98118446).


[Trkola (1998)] A. Trkola, W. A. Paxton, S. P. Monard, J. A. Hoxie, M. A. Siani, D. A. Thompson, L. Wu,
   C. R. Mackay, R. Horuk, J. P. Moore. Genetic subtype-independent inhibition of human immunodeficiency
   virus type 1 replication by CC and CXC chemokines. 
   J Virol 72:396-404, 1998.

   NOTE:   (Medline: 98080427).


[Vallejo (1998)] A. Vallejo, A. Heredia, A. Mas, S. F. Lee, J. S. Epstein, V. Soriano, I. K. Hewlett.
   Tropism, coreceptor use, and phylogenetic analysis of both the V3 loop and the protease gene of three
   novel HIV-1 group O isolates. 
   J Acquir Immune Defic Syndr Hum Retrovirol 18:417-25, 1998.
    
   NOTE:   (Medline: 98379694).


[Zhang (1996)] L. Zhang, Y. Huang, T. He, Y. Cao, D. D. Ho. HIV-1  subtype and second-receptor use
   (letter). Nature 383:768, 1996.
    
   NOTE:   (Medline: 97048157).

[Zhang (1998)] Y. J. Zhang, T. Dragic, Y. Cao, L. Kostrikis, D. S. Kwon, D. R. Littman, V. N. KewalRamani,
   J. P. Moore. Use of coreceptors other than CCR5 by non-syncytium-inducing adult and pediatric isolates
   of human immunodeficiency virus type 1 is rare in vitro. 
   J Virol 72:9337-44, 1998.
    
   NOTE:   (Medline: 98440604).

   
last modified: Wed Sep 16 14:06 2009


Questions or comments? Contact us at seq-info@lanl.gov.

 
Operated by Los Alamos National Security, LLC, for the U.S. Department of Energy's National Nuclear Security Administration
Copyright © 2005-2012 LANS LLC All rights reserved | Disclaimer/Privacy

Dept of Health & Human Services Los Alamos National Institutes of Health