Antibodies capable of neutralizing HIV-1 often target variable regions 1 and 2 (V1V2) of the HIV-1 envelope, but the mechanism of their elicitation has been unclear. Here we define the developmental pathway by which such antibodies are generated and acquire the requisite molecular characteristics for neutralization. Twelve somatically related neutralizing antibodies (CAP256-VRC26.01–12) were isolated from donor CAP256 (from the Centre for the AIDS Programme of Research in South Africa (CAPRISA)); each antibody contained the protruding tyrosine-sulphated, anionic antigen-binding loop (complementarity-determining region (CDR) H3) characteristic of this category of antibodies. Their unmutated ancestor emerged between weeks 30–38 post-infection with a 35-residue CDR H3, and neutralized the virus that superinfected this individual 15 weeks after initial infection. Improved neutralization breadth and potency occurred by week 59 with modest affinity maturation, and was preceded by extensive diversification of the virus population. HIV-1 V1V2-directed neutralizing antibodies can thus develop relatively rapidly through initial selection of B cells with a long CDR H3, and limited subsequent somatic hypermutation. These data provide important insights relevant to HIV-1 vaccine development.

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Data deposits

Coordinates and structure factors for CAP256-VRC26 lineage Fabs have been deposited with the Protein Data Bank under accession codes 4ODH, 4OCR, 4OD1, 4ORG, 4OCW, 4OD3 and 4OCS. The EM reconstruction density for the CAP256-VRC26.09 complex with BG505 SOSIP.664 trimer has been deposited with the Electron Microscopy Data Bank under accession code EMD-5856. We have also deposited deep sequencing data used in this study to National Center for Biotechnology Information Short Reads Archives (SRA) under accession numbers SRP034555 and SRP017087. Information deposited with GenBank includes: the heavy- and light-chain variable region sequences of cloned antibodies CAP256-VRC26.01-12, UCA, I1 and I2 (accession numbers KJ134860KJ134889); bioinformatically identified VRC26-related sequences from B cell transcripts: 680 heavy chains and 472 light chains (accession numbers KJ133708KJ134387, KJ134388KJ134859); and CAP256 Env sequences (accession numbers KF996576KF996716).


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We thank the participants in the CAPRISA 002 study for their commitment. For technical assistance and advice, we thank: K. Mlisana, S. Sibeko, N. Naicker, the CAPRISA 002 clinical team, N. Samsunder, S. Heeralall, B. Lambson, M. Madzivhandila, T. Khoza, C. Mitchell Scheepers, E. Turk, C.-L. Lin, M. Roederer, J. Stuckey, B. Hartman, G. Loots, J. H. Lee, G. Ippolito, B. Briney, S. Hunicke-Smith and J. Wheeler, and members of the WCMC HIVRAD Core and the NIH Vaccine Research Center HIMS, HIMC, SBS and SBIS sections. We thank J. Baalwa, D. Ellenberger, F. Gao, B. Hahn, K. Hong, J. Kim, F. McCutchan, D. Montefiori, J. Overbaugh, E. Sanders-Buell, G. Shaw, R. Swanstrom, M. Thomson, S. Tovanabutra and L. Zhang for contributing the HIV-1 Envelope plasmids used in our neutralization panel. Funding was provided by the intramural research programs of the Vaccine Research Center and NIAID, the Fogarty International Center, NHGRI, and NIGMS of the National Institutes of Health, USA; the International AIDS Vaccine Initiative; the National Science Foundation; Scripps CHAV-ID; the South African Department of Science and Technology; and fellowships from the Wellcome Trust, Hertz Foundation, Donald D. Harrington Foundation, Poliomyelitis Research Foundation and the National Research Foundation of South Africa. Use of sector 22 (Southeast Region Collaborative Access team) at the Advanced Photon Source was supported by the US Department of Energy, Basic Energy Sciences, Office of Science, under contract number W-31-109-Eng-38.

Author information

Author notes

    • Nicole A. Doria-Rose
    • , Chaim A. Schramm
    • , Jason Gorman
    •  & Penny L. Moore

    These authors contributed equally to this work.


  1. Vaccine Research Center, National Institute of Allergy and Infectious Diseases, National Institutes of Health, Bethesda, Maryland 20892, USA

    • Nicole A. Doria-Rose
    • , Jason Gorman
    • , Michael J. Ernandes
    • , Ivelin S. Georgiev
    • , Marie Pancera
    • , Ryan P. Staupe
    • , Han R. Altae-Tran
    • , Robert T. Bailer
    • , Aliaksandr Druz
    • , Rui Kong
    • , Mark K. Louder
    • , Nancy S. Longo
    • , Krisha McKee
    • , Sijy O’Dell
    • , Ryan S. Roark
    • , Rebecca S. Rudicell
    • , Stephen D. Schmidt
    • , Cinque Soto
    • , Yongping Yang
    • , Peter D. Kwong
    • , Lawrence Shapiro
    •  & John R. Mascola
  2. Department of Biochemistry, Columbia University, New York, New York 10032, USA

    • Chaim A. Schramm
    • , Zhenhai Zhang
    •  & Lawrence Shapiro
  3. Center for HIV and STIs, National Institute for Communicable Diseases of the National Health Laboratory Service (NHLS), Johannesburg, 2131, South Africa

    • Penny L. Moore
    • , Jinal N. Bhiman
    • , Molati Nonyane
    • , Constantinos Kurt Wibmer
    •  & Lynn Morris
  4. Faculty of Health Sciences, University of the Witwatersrand, Johannesburg, 2050, South Africa

    • Penny L. Moore
    • , Jinal N. Bhiman
    • , Constantinos Kurt Wibmer
    •  & Lynn Morris
  5. Centre for the AIDS Programme of Research in South Africa (CAPRISA), University of KwaZulu-Natal, Congella, 4013, South Africa

    • Penny L. Moore
    • , Nigel J. Garrett
    • , Carolyn Williamson
    • , Salim S. Abdool Karim
    •  & Lynn Morris
  6. Department of Chemical Engineering, University of Texas at Austin, Austin, Texas 78712, USA

    • Brandon J. DeKosky
    •  & George Georgiou
  7. Department of Integrative Structural and Computational Biology, The Scripps Research Institute, La Jolla, California 92037, USA

    • Helen J. Kim
    • , Ian A. Wilson
    •  & Andrew B. Ward
  8. Center for HIV/AIDS Vaccine Immunology and Immunogen Discovery, The Scripps Research Institute, La Jolla, California 92037, USA

    • Helen J. Kim
    • , Ian A. Wilson
    •  & Andrew B. Ward
  9. IAVI Neutralizing Antibody Center, The Scripps Research Institute, La Jolla, California 92037, USA

    • Helen J. Kim
    • , Ian A. Wilson
    •  & Andrew B. Ward
  10. Torrey Pines Institute, San Diego, California 92037, USA

    • Ema T. Crooks
    •  & James M. Binley
  11. Weill Medical College of Cornell University, New York, New York 10065, USA

    • Albert Cupo
    •  & John P. Moore
  12. Department of Biomedical Engineering, University of Texas at Austin, Austin, Texas, USA

    • Kam H. Hoi
    •  & George Georgiou
  13. Institute of Infectious Diseases and Molecular Medicine, Division of Medical Virology, University of Cape Town and NHLS, Cape Town 7701, South Africa

    • Daniel J. Sheward
    •  & Carolyn Williamson
  14. NISC Comparative Sequencing program, National Institutes of Health, Bethesda, Maryland 20892, USA

    • James C. Mullikin
  15. NIH Intramural Sequencing Center, National Human Genome Research Institute, National Institutes of Health, Bethesda, Maryland 20892, USA

    • James C. Mullikin
  16. Department of Medical Microbiology, Academic Medical Center, Amsterdam 1105 AZ, Netherlands

    • Rogier W. Sanders
  17. Skaggs Institute for Chemical Biology, The Scripps Research Institute, La Jolla, California 92037, USA

    • Ian A. Wilson
  18. Department of Molecular Biosciences, University of Texas at Austin, Austin, Texas 78712, USA

    • George Georgiou
  19. Department of Epidemiology, Columbia University, New York, New York 10032, USA

    • Salim S. Abdool Karim


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N.A.D.-R., C.A.S., J.G. and P.L.M. contributed equally to this work. N.A.D.-R., C.A.S., J.G., P.L.M. and J.N.B., designed and performed experiments, analysed data and wrote the manuscript. L.M., P.D.K., L.S. and J.R.M. conceived and designed the experiments, analysed data, and wrote the manuscript. B.J.D., M.J.E., I.S.G, H.J.K., M.P. and R.P.S. conducted experiments and analysed data. H.R.A.-T., B.T.B., E.T.C., A.C., K.H.H., R.K., M.K.L., K.M., M.N., S.O., Ry.S.R., Re.S.R., S.D.S., C.K.W., Y.Y., J.C.M. and NISC conducted experiments. C.W. and A.D. contributed analysis tools and data analysis. S.S.A.K. and N.J.G conceived and managed the CAPRISA cohorts. J.M.B., R.W.S., I.A.W., J.P.M., A.B.W., G.G., N.S.L., D.J.S., C.S. and Z.Z. analysed data.

Competing interests

The authors declare no competing financial interests.

Corresponding authors

Correspondence to Lynn Morris or Peter D. Kwong or Lawrence Shapiro or John R. Mascola.

Extended data

Supplementary information

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  1. 1.

    Supplementary information

    This file contains Supplementary Notes, Supplementary Figures 1-11 and Supplementary Tables 1-5.

Text files

  1. 1.

    Supplementary Figure 12

    This file contains the NGS heavy chain reads in Nexus format.

  2. 2.

    Supplementary Figure 13

    This file contains the NGS light chain reads in Nexus format.

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