23
Views
12
CrossRef citations to date
0
Altmetric
Original Article

B-Cell Superantigens: Molecular and Cellular Implications

, &
Pages 259-290 | Received 02 Sep 1996, Published online: 10 Jul 2009

References

  • White J., Herman A., Pullen A. M., Kubo R., Kappler J. W., Marrack P. The V beta-specific super antigen staphylococcal enterotoxin B: Stimulation of mature T cells and atonal deletion in neonatal mice. Cell 1989; 56: 27–35
  • Herman A., Kappler J. W., Marrack P., Pullen A. M. Superantigens: Mechanism of T-cell stimulation and role in immune responses. Annu. Rev. Immunol. 1991; 9: 745–772
  • Irwin M. J., Gascoigne N. R. Interplay between superantigens and the immune system. J Leukoc. Biol. 1993; 54: 495–503
  • Held W., Acha-Orbea H., MacDonald H. R., Waanders G. A. Superantigens and retroviral infection: Insights from mouse mammary tumor virus. Immunology Today 1994; 15: 184–190
  • Blackman M. A., Woodland D. L. In vivo effects of superantigens. Life Sciences 1995; 57: 1717–1735
  • McCormack J. E., Kappler J., Marrack P. Stimulation with specific antigen can block superantigen-mediated deletion of T cells in vivo. Proc. Natl. Acad. Sci. USA 1994; 91: 2086–2090
  • He X. W., Goronzy J., Weyand C, Paliard X., West S. G., Lafferty J. A., Clements J. R., Kappler J. W., Marrack P., Kotzin B. L. Selective induction of rheumatoid factors by superantigens and human helper T cells. Evidence for the effects of a superantigen in rheumatoid arthritis. Science 1991; 253: 325–329
  • Bohach G. A., Fast D. J., Nelson R. D., Schlievert P. M. Staphylococcal and streptococcal pyrogenic toxins involved in toxic shock syndrome and related illnesses. Crit. Rev. Microbiol. 1990; 17: 251–272
  • Abe J., Kotzin B. L., Jujo K., Melish M. E., Glode M. P., Kohsaka T., Leung D. Y. Selective expansion of T cells expressing T-cell receptor variable regions V beta 2 and V beta 8 in Kawasaki disease. Proc. Natl. Acad. Sci. USA 1992; 89: 4066–4070
  • Paliard X., West S. G., Lafferty J. A., Clements J. R., Kappler J. W., Marrack P., Kotzin B. L. Evidence for the effects of a superantigen in rheumatoid arthritis. Science 1991; 253: 325–329
  • Inganas M. Comparison of mechanisms of interaction between protein A from Staphylococcus aureus and human monoclonal IgG, IgA and IgM in relation to the classical Fc gamma and the alternative F(ab)2 epsilon protein A interactions. Scand. J. Immunol. 1981; 13: 343–352
  • Endressen C. The binding of protein A of immunoglobulin G and of Fab and Fc fragments. Acta pathol. Microbiol. Immunol. Scand. (C) 1979; 87: 185–189
  • Romagnani S., Giudizi M. G., Del Prete G., Maggi E., Biagiotti R., Almerigogna F., Ricci M. Demonstration on protein A of two distinct immunoglo-bulin-binding sites and their role in the mitogenic activity of Staphylococcal aureus Cowan I on human B cells. J. Immunol. 1982; 129: 596–602
  • Schrezenmeier H., Fleischer B. Mitogenic activity of staphylococcal protein A is due to contaminating staphylococcal enterotoxins. J. Immunol. Meth. 1987; 105: 133–137
  • Goodglick L., Zevit N., Neshat M. S., Braun J. Mapping the Ig superantigen-binding site of HIV-1 gp120. J. Immunol. 1995; 155: 515–5159
  • Kristiansen S. V., Pascual V., Lipsky P. E. Staphylococcal protein A induces biased production of Ig by VH3-expressing B lymphocytes. J. Immunol. 1994; 153: 2974–2984
  • Kozlowski L. M., Kunning S. R., Zheng Y., Wheatley L. M., Levinson A. I. Staphylococcus aureus Cowan I-induced human immunoglobin responses: Preferential IgM rheumatoid factor production and VH3 mRNA expression by protein A-binding B cells. J. Clin. Immunol. 1995; 15: 145–151
  • Berberian L., Valles-Ayoub Y., Sun N., Martinez-Maza O., Braun J. A VH clonal deficit in human immunodeficiency virus-positive individuals reflects a B-cell maturational arrest. Blood 1991; 78: 175–179
  • Berberian L., Goodglick L., Kipps T. J., Braun J. Immunoglobulin VH3 gene products: Natural ligands for HIV gp120. Science 1993; 261: 1588–1591
  • Muller S., Wang H., Silverman G. J., Bramlet G., Haigwood N., Kohler H. B-cell abnormalities in AIDS: Stable and clonally-restricted antibody response in HIV-1 infection. Scand. J. Immunol. 1993; 38: 327–334
  • Silverman G. J., Roben P., Bouvet J.-P., Sasano M. Superantigen properties of a human sialoprotein involved in gut-associated immunity. J. Clin. Invest. 1995; 96: 417–426
  • Patella V., Bouvet J. P., Marone G. Protein Fv produced during viral hepatitis is a novel activator of human basophils and mast cells. J. Immunol. 1993; 151: 5685–5698
  • Lenert P., Kroon D., Spiegelberg H., Goiub E. S., Zanetti M. Human CD4 binds immunoglobulin. Science 1990; 248: 1639–1642
  • Langone J. J. Protein A of Staphylococcus aureus and related immunoglobulin receptors produced by streptococci and pneumonococci. Adv. Immunol. 1982; 32: 157–252
  • Young W. W., Tamura Y., Wolock D. M., Fox J. W. Staphylococcal protein A binding to the Fab of mouse monoclonal antibodies. J. Immunol. 1984; 133: 3163–3166
  • Sasano M., Burton D. R., Silverman G. J. Molecular selection of human antibodies with an unconventional bacterial B cell superantigen. J. Immunol. 1993; 151: 5822–5839
  • Silverman G. J., Goni F., Fernandez J., Chen P. P., Frangione B., Carson D. A. Distinct patterns of heavy chain variable region subgroup use by human monoclonal autoantibodies of different specificity. J. Exp. Med. 1988; 168: 2361–2366
  • Silverman G. J., Goldfien R., Mageed R. A., Jefferis R., Goni F., Frangione B., Fong S., Carson D. A. Idiotypic and subgroup analysis of human monoclonal rheumatoid factors. Implications for structural and genetic basis of autoantibodies in humans. J. Clin. Invest. 1988; 82: 469–475
  • Sasso E. H., Silverman G. J., Mannik M. Human IgM molecules that bind staphylococcal protein A contain VHIIIH chains. J. Immunol. 1989; 142: 2778–2783
  • Sasso E. H., Silverman G. J., Mannik M. Human IgA and IgG F(ab)2 that bind to staphylococcal protein A belong to the VHIII subgroup. J. Immunol. 1991; 147: 1877–1883
  • Silverman G. J. Human antibody responses to bacterial antigens: Studies of a model conventional antigen and a proposed model B cell superantigen. Int. Rev. Immunol. 1992; 9: 57–78
  • Hakoda M., Hayashimoto S., Yamanaka H., Terai C., Kamatani N., Kashiwazaki S. Molecular basis for the interaction between human IgM and staphylococcal protein A. Clin. Immunol. Immunopathol. 1994; 72: 394–401
  • Ibrahim S., Kaartinen M., Seppala I., Matoso-Ferreira A., Makela O. The alternative binding site for protein A in the Fab fragment of immunoglobulins. Scand. J. Immunol. 1993; 37: 257–264
  • Hillson J. L., Karr N. S., Oppliger I. R., Mannik M., Sasso E. H. The structural basis of germline-encoded VH3 immunoglobulin binding to staphylococcal protein A. J. Exp. Med. 1993; 178: 331–336
  • Ibrahim S., Seppala I., Makela O. V-region-mediated binding of human Ig by protein A. J. Immunol. 1993; 151: 3597–3603
  • Randen I., Potter K. N., Li Y., Thompson K. M., Pascual V., Forre O., Natvig J. B., Capra J. D. Complementarity-determining region 2 is implicated in the binding staphylococcal protein A to human immunoglobulin VHIII variable regions. Eur. J. Immunol. 1993; 23: 2682–2686
  • Silverman G. J. Unconventional antigen-antibody binding interactions and B cell super antigens. The Immunologist 1994; 2: 51–57
  • Riechmann L., Davies J. Backbone assignment, secondary structure and Protein A binding of an isolated human antibody VH domain. J. Biomolec. NMR 1995; 6: 141–152
  • Roben P., Salem A., Silverman G. J. VH3 antibodies bind domain D of staphylococcal protein A. J. Immunol. 1995; 154: 6437–6446
  • Schroeder H. W. J., Hillson J. L., Perlmutter R. M. Structure and evolution of mammalian VH families. Int. Immunol. 1990; 2: 41–50
  • Schroeder H. W., Jr., Mortari F., Shiokawa S., Kirkham P. M., Elgavish R. A., Bertrand F. E. III Developmental regulation of the human antibody repertoire. Ann. N. Y. Acad. Sci. 1995; 764: 242–260
  • Myhre E. B. Interaction of bacterial immunoglobulin receptors with sites in the Fab region. Bacterial Immunoglobulin-Binding Proteins, M. D.P. Boyle. Academic Press, Inc., San Diego, CA 1990; vol. I: 243–256
  • Kirkham P. M., Mortari F., Newton J. A., Schroeder H. W.J. Immunoglobulin VH clan and family identity predicts variable domain structure and may influence antigen binding. EM BO J. 1992; 11: 603–609
  • Malchiodi E. L., Eisenstein E., Fields B. A., Ohlendorf D. H., Schlievert P. M., Karialainen K., Mariuzza R. A. Superantigen binding to a T cell receptor beta chain of known three-dimensional structure. J. Exp. Med. 1995; 182: 1833–1845
  • Seth A., Stem L. J., Ottenhoff T. H., Engel I., Owen M. J., Lamb J. R., Klausner R. D., Wiley D. C. Binary and ternary complexes between T-cell receptor, class II MHC and superantigen in vitro. Nature 1994; 369: 324–327
  • Padlan E. A. Anatomy of the antibody molecule. Mol. Immunol. 1994; 31: 169–217
  • Hakoda M., Kamatani N., Kurumanda S., Kashiwazaki S., Silverman G. J. Involvement of framework region 3 of VH3 IgM in the binding of the B cell superantigen, staphylococcal protein A. Arthritis Rheum. 1995; 38: S190
  • Silverman G. J., Lucas A. H. Variable region diversity in human circulating antibodies specific for the capsular polysaccharide of Haemophilus influenzae type b. Preferential usage of two types of VH3 heavy chains. J. Clin. Invest. 1991; 88: 911–920
  • Hakoda M., Kamatani N., Hayashimoto-Kurumada S., Silverman G. J., Yamanaka H., Terai C., Kashiwazaki S. Differential binding avidities of human IgM for staphylococcal protein A derive from specific germline VH3 gene usage. J. Immunol 1996, in press
  • Radic M. Z., Mackle J., Erikson J., Mol C, Anderson W. F., Weigert M. Residues that mediate DNA binding of autoimmune antibodies. J. Immunol. 1993; 150: 4966–4977
  • Hong S. C., Waterbury G., Janeway C. A., Jr. Different superantigens interact with distinct sites in the Vβ domain of a single T cell receptor. J. Exp. Med. 1996; 183: 1437–1446
  • Choi Y., Herman A., DiGustro D., Wade T., Marrack P., Kappler J. Residues of the variable region of the T-cell-receptor β chain that interact with S. aureus toxin superantigens. Nature 1996; 346: 471–473
  • Pullen A. M., Wade T., Marrack P., Kappler J. W. Identification of the region of T cell receptor beta chain that interacts with the self-superantigen MIs-la. Cell 1990; 61: 1365–1374
  • Pullen A. M., Bill J., Kubo R. T., Marrack P., Kappler J. W. Analysis of the interaction site for the self superantigen Mis-la on T cell receptor V beta. J. Exp. Med. 1991; 173: 1183–1192
  • Cazenave P. A., Marche P. N., Jouvin-Marche E., Voegtle D., Bonhomme F., Bandeira A., Coutinho A. V beta 17 gene polymorphism in wild-derived mouse strains: Two amino acid substitutions in the V beta 17 region greatly alter T cell receptor specificity. Cell 1990; 63: 717–728
  • Pontzer C. H., Irwin M. J., Gascoigne N. R., Johnson H. M. T-cell antigen receptor binding sites for the microbial superantigen staphylococcal enterotoxin A. Proc. Natl. Acad. Sci. USA 1992; 89: 7727–7731
  • White J., Pullen A., Choi K., Marrack P., Kappler J. W. Antigen recognition properties of mutant V beta 3+ T cell receptors are consistent with an immunoglobulin-like structure for the receptor. J. Exp. Med 1993; 177: 119–125
  • Bentley G. A., Boulot G., Karjalianen K., Mariuzza R. A. Crystal structure of the b chain of a T cell antigen receptor. Science 1995; 267: 1984–1987
  • Alam S. M., Sim B. C., Gascoigne N. R. Selection of TCR V alpha by MHC class II predicts superantigen reactivity. Int. Immunol. 1995; 7: 1311–1318
  • Woodland D. L., Smith H. P., Surman S., Le P., Wen R., Blackman M. A. Major histocompatibility complex-specific recognition of Mls-1 is mediated by multiple elements of the T cell receptor. J. Exp. Med. 1993; 177: 433–442
  • Waanders G. A., Lussow A. R., MacDonald H. R. Skewed T cell receptor V alpha repertoire among superantigen reactive murine T cells. Int. Immunol. 1993; 5: 55–61
  • Vacchio M. S., Kanagawa O., Tomonari K., Hodes R. J. Influence of T cell receptor V alpha expression on Mlsa superantigen-specific T cell responses. J. Exp. Med. 1992; 175: 1405–1408
  • Blackman M. A., Smith H. P., Le P., Woodland D. L. Influence of the T cell receptor alpha-chain on T cell reactivity and tolerance to Mls-1 in T cell receptor beta-chain transgenic mice. J. Immunol. 1993; 151: 556–565
  • Labrecque N., Thibodeau J., Mourad W., Sekaly R. P. T cell receptor-major histocompatibility complex class II interaction is required for the T cell response to bacterial superantigens. J. Exp. Med. 1994; 180: 1921–1929
  • Deckhut A. M., Chien Y., Blackman M. A., Woodland D. L. Evidence for a functional interaction between the beta chain of major histocompatibility complex class II and the T cell receptor alpha chain during recognition of a bacterial superantigen. J. Exp. Med. 1994; 180: 1931–1935
  • Silverman G. J., Sasano M., Wormsley S. B. Age-associated changes in binding of human B lymphocytes to a VH3-restricted unconventional bacterial antigen. J. Immunol. 1993; 151: 5840–5855
  • Brezinschek H. P., Brezinschek R. I., Lipsky P. E. Analysis of the heavy chain repertoire of human peripheral B cells using single cell polymerase chain reaction. J. Immunol. 1995; 155: 190
  • Klein U., Kuppers R., Rajewsky K. Variable region gene analysis of B cell subsets derived from a 4-year-old child: Somatically mutated memory B cells accumulate throughout life. J. Exp. Med. 1994; 180: 1383–1393
  • Klein U., Kuppers R., Rajewsky K. Human IgM+IgD+ B cells, the major B cell subset in the peripheral blood, express V kappa genes with no or little somatic mutation throughout life. Eur. J. Immunol. 1993; 23: 3272–3277
  • Pascual V., Liu Y. J., Magalski A., De Bouteiller O., Banchereau J., Capra J. D. Analysis of somatic mutation in five B cell subsets of human tonsil. J. Exp. Med. 1994; 180: 329–339
  • Sjodahl J. Structural studies on the four repetitive Fc-binding regions in protein A from Staphylococcus aureus. Eur. J. Biochem. 1977; 78: 471–479
  • Lofdahl S., Guss B., Uhlen M., Philipson L., Lindberg M. Gene for staphylococcal protein A. Proc. Natl. Acad. Sci. USA 1983; 80: 697–701
  • Uhlen M., Guss B., Nilsson B., Gatenbeck S., Philipson L., Lindberg M. Complete sequence of the staphylococcal gene encoding protein A. A gene evolved through multiple duplications. J. Biol. Chetn. 1984; 259: 1695–1702
  • Moks T., Abrahmsen L., Nilsson B., Hellman U., Sjoquist J., Uhlen M. Staphylococcal protein A consists of five IgG-binding domains. Eur. J. Biochem. 1986; 156: 637–643
  • Deisenhofer J., Jones T. A., Huber R., Sjodahl J., Sjoquist J. Crystallographic refinement and atomic models of a human Fc fragment and its complex with fragment B of protein A from Staphylococcus aureus at 2.9- and 2.8-resolution. Hoppe. Seylers. Z. Physiol. Chem. 1978; 359: 975–985
  • Gouda H., Torigoe H., Saito A., Sato M., Arata Y., Shimada I. Three-dimensional solution structure of the B domain of staphylococcal protein A: Comparisons of the solution and crystal structures. Biochemistry 1992; 31: 9665–9672
  • Inganas M., Johansson S. G., Sjoquist J. Further characterization of the alternative protein-A interaction of immunoglobulins: Demonstration of an Fc-binding fragment of protein A expressing the alternative reactivity. Scand. J. Immunol. 1981; 14: 379–388
  • Erntell M., Myhre E. B., Kronvall G. Non-immune F(ab)2- and Fc-mediated interactions of mammalian immunoglobulins with S. aureus and group C and G streptococci. Acta Pathol. Microbiol. Immunol. Scand. [B]. 1986; 94: 377–385
  • Ibrahim S. Immunoglobulin binding specificities of the homology regions (domains) of protein A. Scand. J. Immunol. 1993; 38: 368–374
  • Ljungberg U. K., Jansson B., Niss U., Nilsson R., Sandberg B. E., Nilsson B. The interaction between different domains of staphylococcal protein A and human polyclonal IgG, IgA, IgM and F(ab)2: Separation of affinity from specificity. Mol. Immunol. 1993; 30: 1279–1285
  • Lando P. A., Dohlstein M., Hedlund G., Akerblom E., Kalland T. T cell killing of human colon carcinomas by monoclonal antibody-targeted super-antigens. Cancer Immunol. Immunother. 1993; 36: 223–228
  • Avery A. C., Markowitz J. S., Grusby M. J., Glimscher L. H., Cantor H. Activation of T cells by superantigen in class II-negative mice. J. Immunol. 1994; 153: 483–491
  • Bouvet J. P., Pirès R., Lunel-Fabiani F., Crescenzo-Chaigne B., Maillard P., Valla D., Opolon P., Pillot J. Protein F. A novel F(ab)-binding factor, present in normal liver, and largely released in the digestive tract during hepatitis. J. Immunol. 1990; 145: 1176–1180
  • Bouvet J. P., Pirès R., Charlemagne J., Pillot J., Iscaki S. Non-immune binding of human protein Fv to immunoglobulins from various mammalian and non-mammalian species. Scand. J. Immunol. 1991; 34: 491–496
  • Ruffet E., Pirès R., Pillot J., Bouvet J. P. Activation of the classical pathway of complement by non-immune complexes of immunoglobulins with human protein Fv (Fv fragment-binding protein). Scand. J. Immunol. 1994; 40: 359–362
  • Bouvet J. P., Pirés R., Iscaki S., Pillot J. Nonimmune macromolecular complexes of Ig in human gut lumen. Probable enhancement of antibody functions. J. Immunol. 1993; 151: 2562–2571
  • Silverman G. J., Pirés R., Bouvet J. P. An endogenous sialoprotein and a bacterial B-cell superantigen compete in their VH family-specific binding interactions with human immunoglobulins. J. Immunol 1996, in press
  • Yamada M., Wasserman R., Reichard B. A., Shane S., Caton A. J., Rovera G. Preferential utilization of specific immunoglobulin heavy chain diversity. J. Exp. Med. 1991; 173: 395–407
  • Sanz I. Multiple mechanisms participate in the generation of diversity of human H chain CDR3 regions. J. Immunol. 1991; 147: 1720–1729
  • Luther S., Shakhov A. N., Xenarios I., Haga S., Imai S., Acha-Orbea H. New infectious mammary tumor virus superantigen with V beta-specificity identical to staphylococcal enterotoxin B (SEB). J. Immunol. 1994; 24: 1757–1764
  • Sasso E. H., Johnson T., Kipps T. J. Expression of the immunoglobulin VH gene 51 P1 is proportional to its germline gene copy number. J. Clin. Invest. 1996; 97: 2074–2080
  • Pascual V., Capra J. D. B-cell superantigens. Curr. Opin. in Immunol. 1991; 1: 315–317
  • Childs R., Feizi T. Cross idiotypic specificity among heavy chains of macroglobulins with blood group I and i specificities. Nature 1975; 255: 562–564
  • Silverman G. J., Carson D. A. Structural characterization of human monoclonal cold agglutinins: Evidence for a distinct primary sequence-defined VH4 idiotype. Eur. J. Immunol. 1990; 20: 351–356
  • Pascual V., Victor K., Lelsz D., Spellerberg M. B., Hamblin T. J., Thompson K. M., Randen I., Natvig J., Capra J. D., Stevenson F. K. Nucleotide sequence analysis of the V regions of two IBM cold agglutinins. Evidence that the VH4–21 gene segment is responsible for the major cross-reactive idiotype. J. Immunol. 1991; 146: 4385–4391
  • Leoni J., Ghiso J., Goni F., Frangione B. The primary structure of the Fab fragment of protein KAU, a monoclonal immunoglobulin M cold agglutinin. J. Biol. Chem. 1991; 266: 2836–2842
  • Silberstein L. E., Jefferies L. C., Goldman J., Friedman D., Moore J. S., Nowell P. C., Roelcke D., Pruzanski W., Roudier J., Silverman G. J. Variable region gene analysis of pathologic human autoantibodies to the related i and I red blood cell antigens. Blood 1991; 78: 2372–2386
  • Pascual V., Victor K., Spellerberg M., Hamblin T. J., Stevenson F. K., Capra J. D. VH restriction among human cold agglutinins. The VH4–21 gene segment is required to encode anti-I and anti-i specificities. J. Immunol. 1992; 149: 2337–2344
  • Stevenson F. K., Smith G. J., North J., Hamblin T. J., Glennie M. J. Identification of normal B-cell counterparts of neoplastic cells which secrete cold agglutinins of anti-I and anti-i specificity. Br. J. Haematol. 1989; 72: 9–15
  • Li Y., Spellerberg M. B., Stevenson F. K., Capra J. D., Potter K. N. The I binding specificity of human VH4–34 (VH4–21) encoded antibodies is determined by both VH framework region 1 and complementarity determining region 3. J. Mol. Biol. 1996; 256: 577–589
  • Feeney A. J., Victor K. D., Vu K., Nadel B., Chukwuocha R. U. Influence of the V(D)J recombination mechanism on the formation of the primary T and B cell repertoires. Semin. Immunol. 1994; 6: 155–163
  • Gu H., Tarlington D., Muller W., Rajewsky K., Forster L. Most peripheral B cells in mice are ligand selected. J. Exp. Med. 1991; 173: 1357–1371
  • Pennell C. A., Sheehan K. M., Brodeur P. H., Clarke S. H. Organization and expression of VH gene families preferentially expressed by Ly-1+ (CD5) B cells. Eur. J. Immunol. 1989; 19: 2115–2121
  • Ye J., McCray S. K., Clarke S. H. The transition of pre-BI to pre-BII cells is dependent on the VH structure of the mu/surrogate L chain receptor. EMBOJ. 1996; 15: 1524–1533
  • Kozlowski L., Lambris J. D., Silverman G. J., Levinson A. Complement activation by a B cell superantigen. J. Immunol. 1996; 157: 1200–1206
  • Silverman G. J. Unconventional B-cell antigens and human immune repertoires. Ann. N. Y. Acad. Sci. 1995; 764: 566–585
  • Silverman G. J. B-cell superantigens and the somatic diversification of immune repertoires: Molecular and cellular implications. Human B-cell superantigens, M. Zouali. Landes Co., Austin, TX 1996, in press
  • Fearon D. T., Carter R. H. The CD19/CR2/TAPA-1 complex of B lymphocytes: Linking natural and acquired immunity. Annu. Rev. Immunol. 1995; 13: 127–149

Reprints and Corporate Permissions

Please note: Selecting permissions does not provide access to the full text of the article, please see our help page How do I view content?

To request a reprint or corporate permissions for this article, please click on the relevant link below:

Academic Permissions

Please note: Selecting permissions does not provide access to the full text of the article, please see our help page How do I view content?

Obtain permissions instantly via Rightslink by clicking on the button below:

If you are unable to obtain permissions via Rightslink, please complete and submit this Permissions form. For more information, please visit our Permissions help page.