- Email: [email protected]
Assignment of the CD45-AP Gene to the Centromeric End of Mouse Chromosome 19 and Human Chromosome 11q13.1–q13.3
SHORT COMMUNICATION Assignment of the CD45-AP Gene to the Centromeric End of Mouse Chromosome 19 and Human Chromosome 11q13.1–q13.3 SETSUO TAKAI,* CHRISTINE A. KOZAK,† KOICHI KITAMURA,‡,1
AND
AKIKO TAKEDA‡,2
*Department of Genetics, Research Institute, International Medical Center of Japan, Tokyo 162, Japan; †Laboratory of Molecular Microbiology, National Institute of Allergy and Infectious Diseases, Bethesda, Maryland 20892; and ‡Department of Pathology, Roger Williams Medical Center–Brown University, Providence, Rhode Island 02908 Received May 13, 1996; accepted October 14, 1996
CD45-AP is a recently identified phosphorylated protein that specifically associates with the leukocytespecific transmembrane glycoprotein CD45. The gene for CD45-AP, Ptprcap (protein tyrosine phosphatase, receptor type c polypeptide associated protein), was mapped in mouse by typing the progeny of two multilocus crosses using the mouse CD45-AP cDNA as a Southern hybridization probe. The CD45-AP gene mapped to the centromeric region of Chr 19 proximal to the genes Fth, Cd5, and Pcna-rs. The gene for the human CD45-AP homologue, PTPRCAP, was localized to chromosome band 11q13.1–q13.3 by fluorescence in situ hybridization using human genomic CD45-AP DNA as a hybridization probe. The genetic mapping of the Ptprcap/PTPRCAP genes extends the previously defined synteny conservation of various genes that have been assigned to these regions of the mouse and the human chromosomes. q 1996 Academic Press, Inc.
CD45-AP is a recently identified protein in mouse (17, 18) that specifically associates with CD45, a leukocyte-specific transmembrane glycoprotein (19, 20). Subcellular localization analysis indicates that CD45-AP is a transmembrane protein, and its putative transmembrane domain interacts with the transmembrane segment of CD45 (5). In addition, the amino acid sequence and the pattern of protease susceptibility predict that only a short segment at the NH2-terminus of CD45-AP is located extracellularly and that the bulk of the protein is intracellular. The intracellular domain of CD45 exhibits protein tyrosine phosphatase activity (19, 20) and is thought to dephosphorylate key elements in leukocyte signal transduction, thus playing a critical role in signaling pathways essential for immune responses (4, 10, 22). It has been proposed that the cytoplasmic portion of CD45-AP, which constitutes 1
Present address: Department of Orthopedic Surgery, Goryokaku Hospital, Hakodate, Japan. 2 To whom correspondence should be addressed at Department of Pathology, Roger Williams Medical Center–Brown University, 825 Chalkstone Avenue, Providence, RI 02908. Telephone: (401) 4566557. Fax: (401) 456-6569. E-mail: [email protected].
the bulk of the protein, acts as an adapter that directs the interaction between CD45 and other molecules involved in CD45-mediated signal transduction (5). Mouse CD45-AP has been purified and its cDNA cloned by coimmunoprecipitation with CD45 and internal amino acid sequencing (18). The mouse CD45-AP cDNA consists of 997 basepairs with an open reading frame of 558 nucleotides encoding a predicted protein of 185 amino acids. CD45-AP shared no homology with previously known sequences available from the Basic Local Alignment Search Tool (BLAST) network service. The gene for CD45-AP was mapped in mouse by typing restriction fragment length polymorphisms in two sets of genetic crosses. Southern blotting using the EcoRI–BamHI CD45-AP cDNA fragment (nucleotides 2 to 865 of Ref. 18) as a hybridization probe identified cross-reactive ScaI fragments of 21 kb in Mus musculus musculus, 8.4 kb in Mus spretus, and 8.2 kb in NFS/N and C58/J. Inheritance of the polymorphic fragments was scored in two sets of multilocus crosses: (NFS/N or C58/J 1 M. m. musculus) F1 1 M. m. musculus (6) and (NFS/N 1 M. spretus) F1 1 M. spretus or C58/J (1). Progeny of these crosses have been typed for over 900 markers including the Chr 19 markers Cd5 (CD5), Fth (ferritin heavy chain), and Pcna-rs (proliferating cell nuclear antigen-related sequence). Cd5 was typed in both sets of crosses following HindIII digestion using an Ly1 probe obtained from Dr. B. Mock (National Cancer Institute, Bethesda, MD) (8). Fth was typed as a PstI polymorphism in both sets of crosses using the pMHFY20 clone (ATCC) as a probe (23). Pcna-rs was typed as a BamHI polymorphism in the M. spretus crosses and a BglII polymorphism in the M. m. musculus crosses using the Pcna-rs probe described previously (9). As shown in Fig. 1, the gene for CD45-AP, Ptprcap, mapped to the centromeric region of Chr 19 proximal to the genes Fth, Cd5, and Pcna-rs. The cDNA clone of a human CD45-AP homologue has been reported recently (2, 13) and the predicted amino acid sequence of the human homologue exhibits a high percentage of sequence identity with the mouse CD45-AP cDNA (18). A genomic clone of the human CD45-AP homologue was obtained from a human leuGENOMICS
429
38, 429–431 (1996) ARTICLE NO. 0648
0888-7543/96 $18.00 Copyright q 1996 by Academic Press, Inc. All rights of reproduction in any form reserved.
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SHORT COMMUNICATION
kocyte DNA library by screening with the mouse cDNA (18) as a hybridization probe.3 The genomic clone consisted of approximately 6.5 kb and contained a continuous sequence that encodes the entire coding region reported in the human cDNA (13) except for the methionine residue at the initiation site. This genomic DNA was labeled with biotin-16–dUTP (Boehringer Mannheim) as described previously (16) and fluorescence in situ hybridization (FISH) was performed on metaphase human chromosomes to localize the gene for the human CD45-AP homologue, PTPRCAP. R-banded chromosomes were prepared by standard methods (21) with some modifications (14). Hybridization was performed as described previously (7, 15, 16) with the signal amplification procedure (11, 16). A mixture of various human genomic sequences was added to the hybridization solution for competition to reduce the background binding that is probably caused by the presence of repetitive sequences such as the Alu repeats in the genomic clone. The chromosomes were stained with propidium iodide after hybridization and were observed under a Nikon Optiphot-2-EFD2 microscope with either a B-2A filter for R-bands or a UV-2A filter for G-bands. Forty-two of 50 cells exhibited symmetrical double spots on at least one copy of the long arm of chromosome 11 at the band q13.1–q13.3 when R-bands of human (pro)metaphase chromosomes were observed (Fig. 2). No symmetrical double spots were detected in other 3
K. Kitamura and A. Takeda, unpublished results.
FIG. 2. FISH of human metaphase chromosomes with the human CD45-AP genomic DNA as a probe. Original photographs were taken with Fujichrome films (Sensia, ASA 100). (A) R-banded chromosomes after propidium iodide staining were observed with a B-2A filter. Arrows indicate the symmetrical double spots of PTPRCAP, the gene for CD45-AP. (B) The same chromosomes with G-bands were observed with a UV-2A filter. Comparison of the patterns obtained under the two different filters revealed that the symmetrical double spots indicated by the arrows were located on chromosome 11 at the band q13.1–q13.3.
regions with any prominent coincidence. Thus, we conclude that the gene for the human CD45-AP homologue is located at human chromosome band 11q13.1–q13.3. This region of the human chromosome, i.e., 11q12–q13, shows conserved synteny with the centromeric region of mouse Chr 19 (12) where the mouse CD45-AP gene has been assigned as described above (Fig. 1). ACKNOWLEDGMENTS
FIG. 1. Genetic map location of Ptprcap on mouse Chr 19. To the right of the map between genes are recombination fractions for the M. m. musculus crosses (the first fraction) and the M. spretus crosses (the second fraction). Recombinational distances { the standard errors are given in parentheses. Data were stored and analyzed using the program LOCUS designed by C. E. Buckler (National Institute of Allergy and Infectious Diseases, Bethesda, MD). Recombinational distances were calculated according to Green (3), and genes were ordered by minimizing the number of recombinants. To the left of the map are corresponding map locations for the human homologues of the underlined genes.
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This work was supported in part by a grant from National Institutes of Health (GM 48188 to A.T.). We thank N. Yaseen for useful comments on the paper.
REFERENCES 1. Adamson, M. C., Silver, J., and Kozak, C. A. (1991). The mouse homolog of the gibbon ape leukemia virus receptor: Genetic mapping and a possible receptor function in rodents. Virology 183: 778–781. 2. Bruyns, E., Hendricks-Taylor, L. R., Meuer, S., Koretzky, G. A., and Schraven, B. (1995). Identification of the sites of interaction
gnmxa
AP: Genomics
431
SHORT COMMUNICATION between lymphocyte phosphatase-associated phosphoprotein (LPAP) and CD45. J. Biol. Chem. 270: 31372–31376. 3. Green, E. L. (1981). Linkage, recombination and mapping. In ‘‘Genetics and Probability in Animal Breeding Experiments,’’ pp. 77–113, Macmillan, New York. 4. Justement, L. B., Campbell, K. S., Chien, N. C., and Cambier, J. C. (1991). Regulation of B cell antigen receptor signal transduction and phosphorylation by CD45. Science 252: 1839–1842. 5. Kitamura, K., Maiti, A., Ng, D. H. W., Johnson, P., Maizel, A. L., and Takeda, A. (1995). Characterization of the interaction between CD45 and CD45-AP. J. Biol. Chem. 270: 21151–21157. 6. Kozak, C. A., Peyser, M., Krall, M., Mariano, T. M., Kumar, C. S., Pestka, S., and Mock, B. A. (1990). Molecular genetic markers spanning mouse chromosome 10. Genomics 8: 519– 524. 7. Lawrence, J. B., Villnave, C. A., and Singer, R. H. (1988). Sensitive, high resolution chromatin and chromosome mapping in situ: Presence and orientation of two closely integrated copies of EBV in a lymphoma line. Cell 52: 51–61. 8. Mock, B. A., Krall, M. M., and Dosik, J. K. (1993). Genetic mapping of tumor susceptibility genes involved in mouse plasmacytomagenesis. Proc. Natl. Acad. Sci. USA 90: 9499–9503. 9. Pilon, A. L., Kozak, C. A., Nebert, D. W., and Puga, A. (1993). Location of the murine Hmg1 gene, encoding an HMG-box protein, to mouse chromosome 2. Mamm. Genome 4: 612–614. 10. Pingel, J. T., and Thomas, M. L. (1989). Evidence that the leukocyte-common antigen is required for antigen-induced T lymphocyte proliferation. Cell 58: 1055–1065. 11. Pinkel, D., Strauma, T., and Gray, J. W. (1986). Cytogenetic analysis using quantitative, high sensitivity, fluorescence hybridization. Proc. Natl. Acad. Sci. USA 83: 2934–2938. 12. Poirier, C., and Guenet, J.-L. (1994). Mouse chromosome 19. Mammal. Genome 5: S266–S275. 13. Schraven, B., Schoenhaut, D., Bruyns, E., Koretzky, G., Eckerskorn, C., Wallich, R., Kirchgessner, H., Sakorafas, P., Labkovsky, B., Ratnofsky, S., and Meuer, S. (1994). LPAP, a novel 32kDa phosphoprotein that interacts with CD45 in human lymphocytes. J. Biol. Chem. 269: 29102–29111.
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6r24$$$322
11-21-96 09:21:57
14. Takahashi, E., Hori, T., O’Connell, P., Leppert, M., and White, R. (1990). R-banding and nonisotopic in situ hybridization: Precise localization of the human type II collagen gene (COL2A1). Hum. Genet. 86: 14–16. 15. Takahashi, E., Hori, T., O’Connell, P., Leppert, M., and White, R. (1991). Mapping of the MYC gene to band 8q24.12–q24.13 by R-banding and distal to fra(8)(q24.11), FRA8E, by fluorescence in situ hybridization. Cytogenet. Cell Genet. 57: 109–111. 16. Takai, S., Yamada, K., Hirayama, N., Miyajima, A., and Taniyama, T. (1994). Mapping of the human gene encoding the mutual signal-transducing subunit (b-chain) of the human granulocyte-macrophage colony-stimulating factor (GM-CSF), interleukin-3 (IL-3), and interleukin-5 (IL-5) receptor complexes to chromosome 22q13.1. Hum. Genet. 93: 198–200. 17. Takeda, A., Wu, J. J., and Maizel, A. L. (1992). Evidence for monomeric and dimeric forms of CD45 associated with a 30kDa phosphorylated protein. J. Biol. Chem. 267: 16651–16659. 18. Takeda, A., Maizel, A. L., Kitamura, K., Ohta, T., and Kimura, S. (1994). Molecular cloning of the CD45-associated 30-kDa protein. J. Biol. Chem. 269: 2357–2360. 19. Tonks, N. K., Charbonneau, H., Diltz, C. D., Fischer, E. H., and Walsh, K. A. (1988). Demonstration that the leukocyte common antigen CD45 is a protein tyrosine phosphatase. Biochemistry 27: 8695–8701. 20. Trowbridge, I. S., and Thomas, M. L. (1994). CD45: An emerging role as a protein tyrosine phosphatase required for lymphocyte activation and development. Annu. Rev. Immunol. 12: 85– 116. 21. Viegas-Pequignot, E., and Dutrillaux, B. (1978). Une methode simple pour obtenir desprophases et des prometaphases. Ann. Genet. 21: 122–125. 22. Volarevic, S., Niklinska, B. B., Burns, C. M., June, C. H., Weissman, A. M., and Ashwell, J. D. (1993). Regulation of TCR signaling by CD45 lacking transmembrane and extracellular domains. Science 260: 541–544. 23. Yachou, A.-K., Renaudie, F., Guenet, J.-L., Simon-Chazottes, D., Jones, R., Grandchamp, B., and Beaumont, C. (1991). Mouse ferritin H multigene family is polymorphic and contains a single multiallelic functional gene located on chromosome 19. Genomics 10: 531–538.
gnmxa
AP: Genomics
AND
AKIKO TAKEDA‡,2
*Department of Genetics, Research Institute, International Medical Center of Japan, Tokyo 162, Japan; †Laboratory of Molecular Microbiology, National Institute of Allergy and Infectious Diseases, Bethesda, Maryland 20892; and ‡Department of Pathology, Roger Williams Medical Center–Brown University, Providence, Rhode Island 02908 Received May 13, 1996; accepted October 14, 1996
CD45-AP is a recently identified phosphorylated protein that specifically associates with the leukocytespecific transmembrane glycoprotein CD45. The gene for CD45-AP, Ptprcap (protein tyrosine phosphatase, receptor type c polypeptide associated protein), was mapped in mouse by typing the progeny of two multilocus crosses using the mouse CD45-AP cDNA as a Southern hybridization probe. The CD45-AP gene mapped to the centromeric region of Chr 19 proximal to the genes Fth, Cd5, and Pcna-rs. The gene for the human CD45-AP homologue, PTPRCAP, was localized to chromosome band 11q13.1–q13.3 by fluorescence in situ hybridization using human genomic CD45-AP DNA as a hybridization probe. The genetic mapping of the Ptprcap/PTPRCAP genes extends the previously defined synteny conservation of various genes that have been assigned to these regions of the mouse and the human chromosomes. q 1996 Academic Press, Inc.
CD45-AP is a recently identified protein in mouse (17, 18) that specifically associates with CD45, a leukocyte-specific transmembrane glycoprotein (19, 20). Subcellular localization analysis indicates that CD45-AP is a transmembrane protein, and its putative transmembrane domain interacts with the transmembrane segment of CD45 (5). In addition, the amino acid sequence and the pattern of protease susceptibility predict that only a short segment at the NH2-terminus of CD45-AP is located extracellularly and that the bulk of the protein is intracellular. The intracellular domain of CD45 exhibits protein tyrosine phosphatase activity (19, 20) and is thought to dephosphorylate key elements in leukocyte signal transduction, thus playing a critical role in signaling pathways essential for immune responses (4, 10, 22). It has been proposed that the cytoplasmic portion of CD45-AP, which constitutes 1
Present address: Department of Orthopedic Surgery, Goryokaku Hospital, Hakodate, Japan. 2 To whom correspondence should be addressed at Department of Pathology, Roger Williams Medical Center–Brown University, 825 Chalkstone Avenue, Providence, RI 02908. Telephone: (401) 4566557. Fax: (401) 456-6569. E-mail: [email protected].
the bulk of the protein, acts as an adapter that directs the interaction between CD45 and other molecules involved in CD45-mediated signal transduction (5). Mouse CD45-AP has been purified and its cDNA cloned by coimmunoprecipitation with CD45 and internal amino acid sequencing (18). The mouse CD45-AP cDNA consists of 997 basepairs with an open reading frame of 558 nucleotides encoding a predicted protein of 185 amino acids. CD45-AP shared no homology with previously known sequences available from the Basic Local Alignment Search Tool (BLAST) network service. The gene for CD45-AP was mapped in mouse by typing restriction fragment length polymorphisms in two sets of genetic crosses. Southern blotting using the EcoRI–BamHI CD45-AP cDNA fragment (nucleotides 2 to 865 of Ref. 18) as a hybridization probe identified cross-reactive ScaI fragments of 21 kb in Mus musculus musculus, 8.4 kb in Mus spretus, and 8.2 kb in NFS/N and C58/J. Inheritance of the polymorphic fragments was scored in two sets of multilocus crosses: (NFS/N or C58/J 1 M. m. musculus) F1 1 M. m. musculus (6) and (NFS/N 1 M. spretus) F1 1 M. spretus or C58/J (1). Progeny of these crosses have been typed for over 900 markers including the Chr 19 markers Cd5 (CD5), Fth (ferritin heavy chain), and Pcna-rs (proliferating cell nuclear antigen-related sequence). Cd5 was typed in both sets of crosses following HindIII digestion using an Ly1 probe obtained from Dr. B. Mock (National Cancer Institute, Bethesda, MD) (8). Fth was typed as a PstI polymorphism in both sets of crosses using the pMHFY20 clone (ATCC) as a probe (23). Pcna-rs was typed as a BamHI polymorphism in the M. spretus crosses and a BglII polymorphism in the M. m. musculus crosses using the Pcna-rs probe described previously (9). As shown in Fig. 1, the gene for CD45-AP, Ptprcap, mapped to the centromeric region of Chr 19 proximal to the genes Fth, Cd5, and Pcna-rs. The cDNA clone of a human CD45-AP homologue has been reported recently (2, 13) and the predicted amino acid sequence of the human homologue exhibits a high percentage of sequence identity with the mouse CD45-AP cDNA (18). A genomic clone of the human CD45-AP homologue was obtained from a human leuGENOMICS
429
38, 429–431 (1996) ARTICLE NO. 0648
0888-7543/96 $18.00 Copyright q 1996 by Academic Press, Inc. All rights of reproduction in any form reserved.
AID
Genom 4454
/
6r24$$$321
11-21-96 09:21:57
gnmxa
AP: Genomics
430
SHORT COMMUNICATION
kocyte DNA library by screening with the mouse cDNA (18) as a hybridization probe.3 The genomic clone consisted of approximately 6.5 kb and contained a continuous sequence that encodes the entire coding region reported in the human cDNA (13) except for the methionine residue at the initiation site. This genomic DNA was labeled with biotin-16–dUTP (Boehringer Mannheim) as described previously (16) and fluorescence in situ hybridization (FISH) was performed on metaphase human chromosomes to localize the gene for the human CD45-AP homologue, PTPRCAP. R-banded chromosomes were prepared by standard methods (21) with some modifications (14). Hybridization was performed as described previously (7, 15, 16) with the signal amplification procedure (11, 16). A mixture of various human genomic sequences was added to the hybridization solution for competition to reduce the background binding that is probably caused by the presence of repetitive sequences such as the Alu repeats in the genomic clone. The chromosomes were stained with propidium iodide after hybridization and were observed under a Nikon Optiphot-2-EFD2 microscope with either a B-2A filter for R-bands or a UV-2A filter for G-bands. Forty-two of 50 cells exhibited symmetrical double spots on at least one copy of the long arm of chromosome 11 at the band q13.1–q13.3 when R-bands of human (pro)metaphase chromosomes were observed (Fig. 2). No symmetrical double spots were detected in other 3
K. Kitamura and A. Takeda, unpublished results.
FIG. 2. FISH of human metaphase chromosomes with the human CD45-AP genomic DNA as a probe. Original photographs were taken with Fujichrome films (Sensia, ASA 100). (A) R-banded chromosomes after propidium iodide staining were observed with a B-2A filter. Arrows indicate the symmetrical double spots of PTPRCAP, the gene for CD45-AP. (B) The same chromosomes with G-bands were observed with a UV-2A filter. Comparison of the patterns obtained under the two different filters revealed that the symmetrical double spots indicated by the arrows were located on chromosome 11 at the band q13.1–q13.3.
regions with any prominent coincidence. Thus, we conclude that the gene for the human CD45-AP homologue is located at human chromosome band 11q13.1–q13.3. This region of the human chromosome, i.e., 11q12–q13, shows conserved synteny with the centromeric region of mouse Chr 19 (12) where the mouse CD45-AP gene has been assigned as described above (Fig. 1). ACKNOWLEDGMENTS
FIG. 1. Genetic map location of Ptprcap on mouse Chr 19. To the right of the map between genes are recombination fractions for the M. m. musculus crosses (the first fraction) and the M. spretus crosses (the second fraction). Recombinational distances { the standard errors are given in parentheses. Data were stored and analyzed using the program LOCUS designed by C. E. Buckler (National Institute of Allergy and Infectious Diseases, Bethesda, MD). Recombinational distances were calculated according to Green (3), and genes were ordered by minimizing the number of recombinants. To the left of the map are corresponding map locations for the human homologues of the underlined genes.
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6r24$$$322
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This work was supported in part by a grant from National Institutes of Health (GM 48188 to A.T.). We thank N. Yaseen for useful comments on the paper.
REFERENCES 1. Adamson, M. C., Silver, J., and Kozak, C. A. (1991). The mouse homolog of the gibbon ape leukemia virus receptor: Genetic mapping and a possible receptor function in rodents. Virology 183: 778–781. 2. Bruyns, E., Hendricks-Taylor, L. R., Meuer, S., Koretzky, G. A., and Schraven, B. (1995). Identification of the sites of interaction
gnmxa
AP: Genomics
431
SHORT COMMUNICATION between lymphocyte phosphatase-associated phosphoprotein (LPAP) and CD45. J. Biol. Chem. 270: 31372–31376. 3. Green, E. L. (1981). Linkage, recombination and mapping. In ‘‘Genetics and Probability in Animal Breeding Experiments,’’ pp. 77–113, Macmillan, New York. 4. Justement, L. B., Campbell, K. S., Chien, N. C., and Cambier, J. C. (1991). Regulation of B cell antigen receptor signal transduction and phosphorylation by CD45. Science 252: 1839–1842. 5. Kitamura, K., Maiti, A., Ng, D. H. W., Johnson, P., Maizel, A. L., and Takeda, A. (1995). Characterization of the interaction between CD45 and CD45-AP. J. Biol. Chem. 270: 21151–21157. 6. Kozak, C. A., Peyser, M., Krall, M., Mariano, T. M., Kumar, C. S., Pestka, S., and Mock, B. A. (1990). Molecular genetic markers spanning mouse chromosome 10. Genomics 8: 519– 524. 7. Lawrence, J. B., Villnave, C. A., and Singer, R. H. (1988). Sensitive, high resolution chromatin and chromosome mapping in situ: Presence and orientation of two closely integrated copies of EBV in a lymphoma line. Cell 52: 51–61. 8. Mock, B. A., Krall, M. M., and Dosik, J. K. (1993). Genetic mapping of tumor susceptibility genes involved in mouse plasmacytomagenesis. Proc. Natl. Acad. Sci. USA 90: 9499–9503. 9. Pilon, A. L., Kozak, C. A., Nebert, D. W., and Puga, A. (1993). Location of the murine Hmg1 gene, encoding an HMG-box protein, to mouse chromosome 2. Mamm. Genome 4: 612–614. 10. Pingel, J. T., and Thomas, M. L. (1989). Evidence that the leukocyte-common antigen is required for antigen-induced T lymphocyte proliferation. Cell 58: 1055–1065. 11. Pinkel, D., Strauma, T., and Gray, J. W. (1986). Cytogenetic analysis using quantitative, high sensitivity, fluorescence hybridization. Proc. Natl. Acad. Sci. USA 83: 2934–2938. 12. Poirier, C., and Guenet, J.-L. (1994). Mouse chromosome 19. Mammal. Genome 5: S266–S275. 13. Schraven, B., Schoenhaut, D., Bruyns, E., Koretzky, G., Eckerskorn, C., Wallich, R., Kirchgessner, H., Sakorafas, P., Labkovsky, B., Ratnofsky, S., and Meuer, S. (1994). LPAP, a novel 32kDa phosphoprotein that interacts with CD45 in human lymphocytes. J. Biol. Chem. 269: 29102–29111.
AID
Genom 4454
/
6r24$$$322
11-21-96 09:21:57
14. Takahashi, E., Hori, T., O’Connell, P., Leppert, M., and White, R. (1990). R-banding and nonisotopic in situ hybridization: Precise localization of the human type II collagen gene (COL2A1). Hum. Genet. 86: 14–16. 15. Takahashi, E., Hori, T., O’Connell, P., Leppert, M., and White, R. (1991). Mapping of the MYC gene to band 8q24.12–q24.13 by R-banding and distal to fra(8)(q24.11), FRA8E, by fluorescence in situ hybridization. Cytogenet. Cell Genet. 57: 109–111. 16. Takai, S., Yamada, K., Hirayama, N., Miyajima, A., and Taniyama, T. (1994). Mapping of the human gene encoding the mutual signal-transducing subunit (b-chain) of the human granulocyte-macrophage colony-stimulating factor (GM-CSF), interleukin-3 (IL-3), and interleukin-5 (IL-5) receptor complexes to chromosome 22q13.1. Hum. Genet. 93: 198–200. 17. Takeda, A., Wu, J. J., and Maizel, A. L. (1992). Evidence for monomeric and dimeric forms of CD45 associated with a 30kDa phosphorylated protein. J. Biol. Chem. 267: 16651–16659. 18. Takeda, A., Maizel, A. L., Kitamura, K., Ohta, T., and Kimura, S. (1994). Molecular cloning of the CD45-associated 30-kDa protein. J. Biol. Chem. 269: 2357–2360. 19. Tonks, N. K., Charbonneau, H., Diltz, C. D., Fischer, E. H., and Walsh, K. A. (1988). Demonstration that the leukocyte common antigen CD45 is a protein tyrosine phosphatase. Biochemistry 27: 8695–8701. 20. Trowbridge, I. S., and Thomas, M. L. (1994). CD45: An emerging role as a protein tyrosine phosphatase required for lymphocyte activation and development. Annu. Rev. Immunol. 12: 85– 116. 21. Viegas-Pequignot, E., and Dutrillaux, B. (1978). Une methode simple pour obtenir desprophases et des prometaphases. Ann. Genet. 21: 122–125. 22. Volarevic, S., Niklinska, B. B., Burns, C. M., June, C. H., Weissman, A. M., and Ashwell, J. D. (1993). Regulation of TCR signaling by CD45 lacking transmembrane and extracellular domains. Science 260: 541–544. 23. Yachou, A.-K., Renaudie, F., Guenet, J.-L., Simon-Chazottes, D., Jones, R., Grandchamp, B., and Beaumont, C. (1991). Mouse ferritin H multigene family is polymorphic and contains a single multiallelic functional gene located on chromosome 19. Genomics 10: 531–538.
gnmxa
AP: Genomics