Age- and disease-related decline in immune function: An opportunity for "Thymus-Boosting" therapies

Tissue Engineering

Volumn 5, Issue 6, 1999, Pages 499-514

  Berthiaume, François ^a   Aparicio, Carlos L ^a   Eungdamrong, John ^a   Yarmush, Martin L ^a  

^a MASSACHUSETTS GENERAL HOSPITAL   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

CYTOKINE;

AGING; CELLULAR IMMUNITY; CYTOKINE PRODUCTION; HUMAN; INJURY; PRIORITY JOURNAL; REVIEW; STRESS; T LYMPHOCYTE; THYMUS;

EID: 0032765086     PISSN: 10763279     EISSN: None     Source Type: Journal    
DOI: 10.1089/ten.1999.5.499     Document Type: Article

References (155)

1. Miller, J.F.A.P. Immunological function of the thymus. Lancet 2, 748, 1961.
2. Martinez, C., Kersey, J., Papermaster, B., and Bood, R. Skin homograft survival in thymectomized mice. Proc. Soc. Exp. Biol. Med. 109, 193, 1962.
3. Scollay, R., Butcher, E., and Weissman, I. Thymus migration: quantitative studies on the rate of migration on cells from the thymus to the periphery in mice. Eur. J. Immunol. 10, 210, 1980.
4. Blackwell, B.N., Bucci, T.J., Hart, R.W., and Turturro, A. Longevity, body weight, and neoplasia in ad libitumfed and diet-restricted C57BL6 mice fed NIH-313 open formula diet. Toxicol. Pathol. 23, 570, 1995.
5. Hirokawa, K. Understanding the mechanism of the age-related decline in immune function. Nutr. Rev. 50, 361, 1992.
6. + T cell clones in elderly humans. J. Immunol. 158, 4493, 1997.
7. Aspinall, R. Age-associated thymic atrophy in the mouse is due to deficiency affecting rearrangement of the TCR during intrathymic development. J. Immunol. 158, 3037, 1997.
8. Hadden, J.W., Malec, P.H., Coto, J., and Hadden, E.M. Thymic involution in aging. Prospects for correction. Ann. N.Y. Acad. Sci. 673, 231, 1992.
9. Hadden, J.W. Immunostimulants. Immunol. Today 14, 275, 1993.
10. Chatamra, K., Daniel, P.M., Kendall, M.D., and Lam, D.K. Atrophy of the thymus in rats rendered diabetic by streptozotocin. Horm. Metab. Res. 17, 630, 1985.
11. Gelfand, D.W., Goldman, A.S., Law, E.J., et al. Thymic hyperplasia in children recovering from theraml burns. J. Trauma 12, 813, 1972.
12. van Baarlen, J., Schuurman, H.-J., and Huber, J. Acute thymus involution in infancy and childhood: a reliable marker for duration of acute illness. Hum. Pathol. 19, 1155, 1988.
13. Harris, V.J., Ramilo, J., and White, H. The thymic mass as a mediastinal dilemma. Clin. Radiol. 31, 263, 1980.
14. Flurkey, K., Miller, R.A., and Harrison, D.E. Cellular determinants of age-related decrements in the T-cell mitrogen response of B6CBAF1 mice. J. Gerontol. 47, B115, 1992.
15. Kirschmann, D.A., and Murasko, D.M. Splenic and inguinal lymph node T cells of aged mice respond differently to polyclonal and antigen-specific stimuli. Cell. Immunol. 139, 426, 1992.
16. Bloom, E.T., and Horvath, J.A. Cellular and molecular mechanisms of the IL-12-induced increase in allospecific murine cytolytic T cell activity: implications for the age-related decline in CTL. J. Immunol. 152, 4242, 1994.
17. Kelley, K.W., Davila, D.R., Brief, S., Simon, J., and Arkins, S. A pituitary-thymus connection during aging. Ann. N.Y. Acad. Sci. 521, 88, 1988.
18. Palacios, R., and Imhof, B.A. At day 8-8.5 of mouse development the yolk sac, not the embryo proper, has lymphoid precursor potential in vivo and in vitro. Proc. Natl. Acad. Sci. U.S.A. 90, 6581, 1993.
19. Savagner, P., Imhof, B.A., Yamada, K.M., and Thiery, J.-P. Homing of haematopoietic precursor cells to the embryonic thymus: characterization of an invasive mechanism induced by chemotactic peptides. J. Cell Biol. 103, 2715, 1986.
20. Shores, E.W., van Ewijk, W., and Singer, A. Disorganization and restoration of thymic medullary epithelial cells in T cell receptor-negative seid mice: evidence that receptor-bearing lymphocytes influence maturation of the thymic microenvironment. Eur. J. Immunol. 21, 1657, 1991.
21. Surh, C.D., Ernst, B., and Sprent, J. Growth of epithelial cells in the thymic medulla is under the control of mature T cells. J. Exp. Med. 176, 611, 1992.
22. Shores, E.W., Nakayama, T., Wiest, D.L., Takahama, Y., Sharrow, S., and Singer, A. Structurally distinct T cell receptor complexes on developmentally distinct T cell populations in severe combined immunodeficiency mice expressing a TCR beta transgene. J. Immunol. 150, 1263, 1992.
23. Nezelof, C. Pathology of the thymus in immunodeficiency states. Curr. Top. Pathol. 75, 151, 1986.
24. Nezelof, C. Thymic pathology in primary and secondary immunodeficiencies. Histopathology 21, 499, 1992.
25. Raviola, E., and Karnovsky, J. Evidence for a blood-thymus barrier using electron-opaque tracers. J. Exp. Med. 136, 466, 1972.
26. van Ewijk, W. Cell surface topography of thymic microenvironments. Lab. Invest. 59, 579, 1988.
27. Steinmann, G.G. Changes in the human thymus during aging. In: Müller-Hermelink, H.-K., ed. The Human Thymus. Histopathology and Pathology. Berlin: Springer-Verlag, 1986, pp. 43-88.
28. Palacios, R., Kiefer, M., Brockhaus, M., et al. Molecular, cellular, and functional properties of bone marrow T lymphocytes progenitor clones. J. Exp. Med. 166, 33, 1987.
29. Antica, M., Wu, L., Shortman, K., and Scollay, R. Thymic stem cells in mouse bone marrow. Blood 84, 111, 1994.
30. Galy, A., Travis, M., Cen, D., and Chen, B. Human T, B, natural killer, and dendritic cells arise from a common marrow progenitor cell subset. Immunity 3, 459, 1995.
31. Wu, L., Antica, M., Johnson, G.R., Scollay, R., and Shortman, K. Developmental potential of the earliest precursor cells from the adult mouse thymus. J. Exp. Med. 174, 1617, 1991.
32. Matsuzaki, Y., Gyotoku, J., Ogawa, M., et al. Characterization of c-kit positive intrathymic stem cells that are restricted to lymphoid differentiation. J. Exp. Med. 178, 1283, 1993.
33. Wu, L., Scollay, R., Egerton, M., Pearse, M., Spangrude, G.J., and Shortman, K. CD4 expressed on earliest T-lineage precursor cells in the adult murine thymus. Nature 349, 71, 1991.
34. - triple-negative adult mouse thymocytes defined by CD44 and CD25 expression. J. Immunol. 150, 4244, 1993.
35. - thymocyte differentiation. J. Immunol. 152, 4783, 1994.
36. +) thymocytes in normal and genetically modified mice. J. Immunol. 154, 5103, 1995.
37. Zúniga-Pflücker, J.C., and Lenardo, M.J. Regulation of thymocyte development from immature progenitors. Curr. Opin. Immunol. 8, 215, 1996.
38. Anderson, G., Moore, N.C., Owen, J.J.T., and Jenkinson, E.J. Cellular interactions in thymocyte development. Annu. Rev. Immunol. 14, 73, 1996.
39. Mallick, C.A., Dudley, E.C., Viney, J.L., Owen, M.J., and Hayday, A.C. Rearrangement and diversity of T cell receptor β chain genes in thymocytes: a critical role for the β chain in development. Cell 73, 513, 1993.
40. Rothenberg, E.V., Chen, D., and Diamond, R.A. Functional and phenotypic analysis of thymocytes in SCID mice. J. Immunol. 151, 3530, 1993.
41. Fehling H.J., Krotkova, A., Saint-Ruf, C., and von Boehmer, H. Crucial role of the pre-T-cell receptor α gene in development of αβ but not γδ T cells. Nature 375, 795, 1995.
42. Fehling, H.J., and von Boehmer, H. Early αβ T cell development in the thymus of normal and genetically altered mice. Curr. Opin. Immunol. 9, 263, 1997.
43. Hoffman, E.S., Passoni, L., Crompton, T., et al. Productive T-cell receptor β-chain gene rearrangement: coincident regulation of cell cycle and clonality during development in vivo. Genes Dev. 10, 948, 1996.
44. Petrie, H.T., Livak, F., Schatz, D.G., Strasser, A., Crispe, N., and Shortman, K. Multiple rearrangements in T cell receptor a chains maximize the production of useful thymocytes. J. Exp. Med. 178, 615, 1993.
45. Kundig, T.M., Shahinian, A., Kawai, K., et al. Duration of TCR stimulation determines costimulatory requirement of T cells. Immunity 5, 41, 1996.
46. Wilkinson, R.W., Anderson, G., Owen, J.J., and Jenkinson, E.J. Positive selection of thymocytes involves sustained interaction with thymic microenvironment. J. Immunol. 155, 5234, 1995.
47. Merkenschlager, M., Graf, D., Lovatt, M., Bommhardt, U., Zamoyska, R., and Fisher, A.G. How many thymocytes audition for selection? J. Exp. Med. 186, 1149, 1997.
48. Tough, D.F., and Sprent, J. Turnover of naive- and memory-phenotype T cells. J. Exp. Med. 179, 1127, 1994.
49. Scollay, R., and Godfrey, D.I. Thymic emigration: conveyor belts or lucky dips. Immunol. Today 16, 268, 1995.
50. Gabor, M.J., Godfrey, D.I., and Scollay, R. Recent thymic emigrants are distinct from most medullary thymocytes. Eur. J. Immunol. 27, 2010, 1997.
51. Pénit, C., and Vasseur, F. Expansion of mature thymocyte subsets before emigration to the periphery. J. Immunol. 159, 4848, 1997.
52. Pereia, P., and Rocha, B. Post-thymic in vivo expansion of mature αβ T cells. Int. Immunol. 3, 1077, 1991.
53. Berzins, S.P., Boyd, R.L., and Miller, J.F.A.P. The role of the thymus and recent thymic migrants in the maintenance of the adult peripheral lymphocyte pool. J. Exp. Med. 187, 1839, 1998.
54. Tanchot, C., and Rocha, B. Peripheral selection of T cell repertoires: the role of continuous thymus output. J. Exp. Med. 186, 1099, 1997.
55. Coltey, M., Jotereau, F.V., and Le Douarin, N.M. Evidence for a cyclic renewal of lymphocyte precursor cells in the embryonic chick thymus. Cell Differ. 22, 71, 1987.
56. Coltey, M., Bucy, R.P., Chen, C.H., et al. Analysis of the first two waves of thymus homing stem cells and their T cell progeny in chick-quail chimeras. J. Exp. Med. 170, 543, 1989.
57. Dunon, D., Courtois, D., Vainio, O., et al. Ontogeny of the immune system: γ/δ and α/β T cells migrate from the thymus to the periphery in alternating waves. J. Exp. Med. 186, 977, 1997.
58. Le Douarin, N.M., and Jotereau, F.V. Tracing of cells of the avian thymus through embryonic life in interspecific chimeras. J. Exp. Med. 142, 17, 1975.
59. 2-microglobulin is involved in thymus colonization of chicken embryos. E.M.B.O. J. 9, 3315, 1990.
60. Dunon, D., Allioli, N., Vainio, O., Ody, C., and Imhof, B.A. Quantification of T-cell progenitors during ontogeny: thymus colonization depends on blood delivery of progenitors. Blood 93, 2234, 1999.
61. Jotereau, F., Heuze, F., Salomon-Vie, V., and Gascan, H. Cell kinetics in the fetal mouse thymus: precursor cell input, proliferation, and emigration. J. Immunol 138, 1026, 1987.
62. Morin, C., Jotereau, F., and Augustin, A. Patterns of responsiveness of T cell lines and thymocytes reveal waves of specific activity in the post-natal thymus. Int. Immunol. 4, 1091, 1992.
63. Dunon, D., Cooper, M.D., and Imhof, B.A. Thymic origin of embryonic intestinal γ/δ T cells. J. Exp. Med. 177, 257, 1993.
64. Dunon, D., Cooper, M.D., and Imhof, B.A. Migration patterns of thymus derived γ/δ T cells during chicken development. Eur. J. Immunol. 23, 2545, 1993.
65. Dunon, D., Schwager, J., Dangy, J.P., Cooper, M.D., and Imhof, B.A. T cell migration during development: homing is not related to TCR Vβ1 repertoire selection. E.M.B.O. J. 13, 808, 1994.
66. Goldschneider, I., Komschlies, K.L., and Greiner, D.L. Studies of thymocytopoiesis in rats and mice. I. Kinetics of appearance of thymocytes using a direct intrathymic adoptive transfer assay for thymocyte precursors. J. Exp. Med. 163, 1, 1986.
67. Scollay, R., Smith, J., and Stauffer, V. Dynamics of early T cells: prothymocyte migration and proliferation in the adult mouse thymus. Immunol. Rev. 91, 129, 1986.
68. + T cells. J. Exp. Med. 175, 1067, 1992.
69. Shortman, K., Egerton, M., Spangrude, G.J., and Scollay, R. The generation and fate of thymocytes. Semin. Immunol. 2, 3, 1990.
70. Spangrude, G.J., and Scollay, R. Differentiation of hematopoietic stem cells in irradiated mouse thymic lobes. Kinetics and phenotype of progeny. J. Immunol. 145, 3661, 1990.
71. - thymocytes. Proliferation, renewal, and differentiation cell subsets studied by DNA biosynthetic labeling and surface antigen detection. Eur. J. Immunol. 18, 1343, 1988.
72. Egerton, M., Shortman, K., and Scollay, R. The kinetics of immature murine thymocytes development in vivo. Int. Immunol. 2, 501, 1990.
73. Egerton, M., Scollary, R., and Shortman, K. Kinetics of mature T-cell development in the thymus. Proc. Natl. Acad. Sci. U.S.A. 87, 2579, 1990.
74. Aparicio, C.L. Migration of progenitor T cells to the thymus: effect of tumor necrosis factor-alpha on the adhesion of progenitor T cells to endothelial cells [Ph.D. dissertation]. Rutgers University, New Brunswick, 1996.
75. Butcher, E.C. Leukocyte-endothelial cell recognition: Three (or more steps) to specificity and diversity. Cell 67, 1033, 1991.
76. Berg, E.L., Robinson, M.K., Warnock, R.A., and Butcher, E.C. The human peripheral lymph node vascular addressin is a ligand for LECAM-1, the peripheral lymph node homing receptor. J. Cell Biol. 114, 343, 1991.
77. Imhof, B.A., Ruiz, P., Hesse, B., Palacios, R., and Dunon, D. EA-1, a novel adhesion molecule involved in the homing of progenitor T lymphocytes to the thymus. J. Cell Biol. 114, 1069, 1991.
78. 6 integrins participate in pro-T cell homing to the thymus. Eur. J. Immunol. 25, 2034, 1995.
79. de la Hera, A., Acevedo, A., Martson, W., and Sanchez-Madrid, F. Function of CD44 (Pgp-1) homing receptor in human T cell precursors. Int. Immunol. 1, 598, 1989.
80. O'Neill, H.C. Antibody which defines a subset of bone marrow cells that can migrate to thymus. Immunology 68, 59, 1989.
81. O'Neill, H.C. Prothymocyte seeding in the thymus. Immunol. Lett. 27, 1, 1991.
82. Horst, E., Meijer, C.J.L.M., Duijvestjin, A.M., Hartwig, N., Van der Harten, H.J., and Pals, S. The ontogeny of human lymphocyte recirculation: high endothelial cell antigen (HECA-452) and CD44 homing receptor expression in the development of the immune system. Eur. J. Immunol. 20, 1483, 1990.
83. Wu, L., Kincade, P.W., and Shortman, K. The CD44 expressed on the earliest intrathymic precursor population functions as a thymus homing molecule but does not bind to hyaluronate. Immunol. Lett. 38, 69, 1993.
84. Nakano, N., Hardy, R.R., and Kishimoto, T. Identification of intrathymic T progenitor cells by expression of Thy-1, IL2 receptor and CD3. Eur. J. Immunol. 17, 1567, 1987.
85. Hyman, R., Lesley, J., Schulte, R., and Trotter, J. Progenitor cells in the thymus: most thymus-homing progenitor cells in the adult mouse thymus bear Pgp-1 glycoprotein but not interleukin-2 receptor on their cell surface. Cell. Immunol. 101, 320, 1986.
86. Aparicio, C.L., Berthiaume, F., Chang, C.C., and Yarmush, M.L. Tumor necrosis factor-alpha (TNF-α) induces a reversible, time- and dose-dependent adhesion of progenitor T cells to endothelial cells. Mol. Immunol. 33, 671, 1996.
87. Bereta, J., Bereta, M., Cohen, S., and Cohen, M.C. Regulation of VCAM-1 expression and involvement in cell adhesion to murine microvascular endothelium. Cell. Immunol. 147, 313, 1993.
88. Shimizu, Y., Newman, W., Gopal, T.V., et al. Four molecular pathways of T cell adhesion to endothelial cells: roles of LFA-1, VCAM-1, and ELAM-1 and changes in pathway hierarchy under different activation conditions. J. Cell Biol. 113, 1203, 1991.
89. Deman, J., Martin, M.-T., Delvenne, P., Humblet, C., Boniver, J., and Defresne, M.-P. Analysis by in situ hybridization of cells expressing mRNA for tumor necrosis factor in the developing thymus of mice. Dev. Immunol. 2, 103, 1992.
90. Agus, D.B., Surh, C.D., and Sprent, J. Re-entry of T cells to the adult thymus is restricted to activated T cells. J. Exp. Med. 173, 1039, 1991.
91. Pfeffer, K., Matsuyama, T., Kundig, T.M., et al. Mice deficient for the 55 kD tumor necrosis factor receptor are resistant to endotoxic shock, yet succumb to L. monocytogenes infection. Cell 73, 457, 1993.
92. Zlotnik, A., and Moore, T.A. Cytokine production and requirements during T cell development. Curr. Opin. Immunol. 7, 206, 1995.
93. + hemopoietic progenitors. J. Cell Biol. 135, 1655, 1996.
94. Mackall, C.L., Punt, J.A., Morgan, P., Farr, A.G., and Gress, R.E. Thymic function in young/old chimeras: substantial thymic T cell regenerative capacity despite irreversible age-associated thymic involution. Eur. J. Immunol. 28, 1886, 1998.
95. Thoman, M.L. The pattern of T lymphocyte differentiation is altered during thymic involution. Mech. Ageing Dev. 82, 155, 1995.
96. + thymocytes in BALB/c mice. Mech. Ageing Dev. 62, 307, 1992.
97. Thoman, M.L. Early steps in T cell development are affected by aging. Cell. Immunol. 178, 117, 1997.
98. Perez-Mera, M.L., Carnero, D.G., and Rey-Mendez, M. Effect of age on the proportion of mouse bone marrow cells migrating in response to newborn thymus supernatant. Cell migration and thymus evolution in mice. Thymus 18, 237, 1991.
99. Donskoy, E., and Goldschneider, I. Thymocytopoiesis is maintained by blood-borne precursors throughout postnatal life. J. Immunol. 148, 1604, 1992.
100. Eren, R., Globerson, A., Abel, L., and Zharhary, D. Quantitative analysis of bone marrow thymic progenitors in young and aged mice. Cell. Immunol. 127, 238, 1990b.
101. Fridkis-Hareli, M., Mehr, R., Abel, L., and Globerson, A. Developmental interactions of CD4 T cells and thymocytes: age-related differential effects. Mech. Ageing Dev. 73, 169, 1994.
102. Sharp, A., Fridkis-Hareli, M., Eren, R., Kukulansky, T., Abel, L., and Globerson, A. MHC-linked colonization of the thymus and thymocyte development: effects of mature T lymphocytes. Int. Arch. Allergy Immunol. 106, 13, 1995.
103. Lum, L.G. The kinetics of immune reconstitution after human marrow transplantation. Blood 69, 369, 1987.
104. + T-lymphocyte regeneration after intensive chemotherapy. N. Engl. J. Med. 332, 143, 1995.
105. Peylan-Ramu, N., Haddy, T.B., Jones, E., Horvath, K., Adde, M.A., and Magrath, I.T. High frequency of benign mediastinal uptake of gallium-67 after completion of chemotherapy in children with high-grade non-Hodgkin's lymphoma. J. Clin. Oncol. 7, 1800, 1989.
106. Dumont-Girard, F., Roux, E., van Lier, R.A., et al. Reconstitution of the T-cell compartment after bone marrow transplantation: restoration of the repertoire by thymic emigrants. Blood 92, 4464, 1998.
107. Mackall, C.L., Bare, V.C., Granger, L.A., Sharrow, S.O., Titus, J.A., and Gress, R.E. Thymic-independent T cell regeneration occurs via antigen-driven expansion of peripheral T cells resulting in a repertoire that is limited in diversity and prone to skewing. J. Immunol. 156, 4609, 1996.
108. +) T-helper cells after human allogeneic bone marrow transplantation. Blood 90, 850, 1997.
109. + T lymphocytes in children on highly active antiretroviral therapy. Dutch study group for children with HIV infections. AIDS 12, 2155, 1998.
110. Hendrickx, P., and Döhring, W. Thymic atrophy and rebound enlargement following chemotherapy for testicular cancer. Acta Radiol. 30, 263, 1989.
111. Willich, E. Acute and stress-induced involution of the thymus. In: Walter, E., Willich, E., and Webb, W.R., eds. The Thymus. Diagnostic Imaging, Functions, and Pathologic Anatomy. Berlin: Springer-Verlag, 1992, pp. 57-61.
112. Barlow, Y. T lymphocytes and immunosuppression in the burned patient: a review. Burns 20, 487, 1994.
113. Douek, D.C., McFarland, R.D., Keiser, P.H., et al. Changes in thymid function with age and during the treatment of HIV infection. Nature 396, 690, 1998.
114. Willich, E., Walter, E., Otto, H.F., and Hofman, W.J. Developmental abnormalities of the thymus. In: Walter, E., Willich, E., and Webb, W.R., eds. The Thymus. Diagnostic Imaging, Functions, and Pathologic Anatomy. Berlin: Springer-Verlag, 1992, pp. 63-108.
115. Hirokawa, K. Reversing and restoring immune functions. Mech. Ageing Dev. 93, 119, 1997.
116. Binz, K., Joller, P., Froesch, P., Binz, H., Zapf, J., and Froesch, E.R. Repopulation of the atrophied thymus in diabetic rats by insulin-like growth factor I. Proc. Natl. Acad. Sci. U.S.A. 87, 3690, 1990.
117. Ernst, E. Thymus therapy for cancer? A criteria-based, systematic review. Eur. J. Cancer 33, 517, 1997.
118. Hirokawa, K., and Utsuyama, M. The effect of sequential multiple grafting of syngeneic newborn thymus on the immune functions and life expectancy of aging mice. Mech. Ageing Dev. 28, 111, 1984.
119. Metcalf, D. Multiple thymus grafts in aged mice. Nature 208, 87, 1966b.
120. Fabris, N., Mocchegiani, E., and Provinciali, M. Plasticity of neuro-endocrine thymus interactions during aging. A minireview. Cell. Mol. Biol. 43, 529, 1997.
121. Kendall, M.D., Fitzpatrick, F.T.A., Greenstein, B.D., Khoylou, F., Safieh, B., and Hamblin, A. Reversal of ageing changes in the thymus of rats by chemical or surgical castration. Cell Tissue Res. 261, 555, 1990.
122. Greenstein, B.D., Fitzpatrick, F.T.A., Kendall, M.D., and Wheeler, M.J. Regeneration of the thymus in old male rats treated with a stable analog of LHRH. J. Endocrinol. 112, 345, 1987.
123. Prasad, A.S., Fitzgerald, J.T., Hess, J.W., Kaplan, J., Pelen, F., and Dardenne, M. Zinc deficiency in elderly patients. Nutrition 9, 218, 1993.
124. Dardenne, M., Boukaiba, N., Gagnereault, M.C., et al. Restoration of the thymus in aging mice by in vivo zinc supplementation. Clin. Immunol. Immunopathol. 66, 127, 1993.
125. Mocchegiani, E., Santarelli, L., Muzzioli, M., and Fabris, N. Reversibility of the thymic involution and of age-related peripheral immune dysfunctions by zinc supplementation in old mice. Int. J. Pharmacol. 17, 703, 1995.
126. Jenkinson, E.J., Kingston, R., and Owen, J.J. Stromal cell populations in the developing thymus of normal and nude mice. Adv. Exp. Med. Biol. 186, 245, 1985.
127. Cordier, A.C., and Haumont, S.M. Development of thymus, parathyroids, and ultimobranchial bodies in NMRI and nude mouse. Am. J. Anat. 157, 227, 1980.
128. Bulloch, K., and Pomerantz, W. Autonomie nervous system innervation of thymic-related lymphoid tissue in the wild type and nude mice. J. Comp. Neurol. 228, 57, 1984.
129. Bulloch, K., Cullen, M.R., Schwartz, R.H., and Longo, D.L. Development of innervation within syngeneic thymus tissue transplanted under the kidney capsule of the nude mouse: a light and ultrastructural microscope study. J. Neurosci. Res. 18, 16, 1987.
130. Okazaki, H., Ito, M., Sudo, T., et al. IL-7 promotes thymocyte proliferation and maintains immunocompetent thymocytes bearing αβ or γδ T-cell receptors in vitro: synergism with IL-2. J. Immunol. 143, 2917, 1989.
131. - thymocytes. J. Immunol. 146, 3068, 1991.
132. Godfrey, D.I., Zlotnik, A., and Suda, T. Phenotypic and functional characterization of c-kit expression during intrathymic T cell development. J. Immunol. 149, 2281, 1992.
133. - thymocytes: synergy with IL-7. Cell. Immunol. 157, 118, 1994.
134. Rodewald, H.-R., Kretschmar, K., Swat, W., and Takeda, S. Intrathymically expressed c-kit ligand (stem cell factor) is a major factor driving the expansion of very immature thymocytes in vivo. Immunity 3, 313, 1995.
135. Grabstein, K.H., Waldschmidt, T.J., Finkelman, F.D., et al. Inhibition of murine B and T lymphopoiesis in vivo by an anti-interleukin 7 monoclonal antibody. J. Exp. Med. 178, 257, 1993.
136. Bhatia, S.K., Tygrett, L.T., Grabstein, K.H., and Waldschmidt, T.J. The effect of in vivo IL-7 deprivation on T cell maturation. J. Exp. Med. 181, 1399, 1995.
137. Peschon, J.J., Morrissey, P.J., Grabstein, K.H., et al. Early lymphocyte expansion is severely impaired in interleukin 7-deficient mice. J. Exp. Med. 180, 1955, 1994.
138. von Freeden-Jeffrey, U., Vieira, P., Lucian, L.A., McNeil, T., Burdach, S.E.G., and Murray, R. Lymphopenia in interleukin (IL)-7 gene-deleted mice identifies IL-7 as a nonredundant cytokine. J. Exp. Med. 181, 1519, 1995.
139. Sudo, T., Nishikawa, S., Ohno, N., et al. Expression and function of the interleukin 7 receptor in murine lymphocytes. Proc. Natl. Acad. Sci. U.S.A. 90, 9125, 1993.
140. Rich, B.E., and Leder, P. Transgenic expression of interleukin 7 restores T cell populations in nude mice. J. Exp. Med. 181, 1223, 1995.
141. Abdul-Hai, A., Or, R., Slavin, S., et al. Stimulation of immune reconstitution by interleukin-7 after syngeneic bone marrow transplantation in mice. Exp. Hematol. 24, 1416, 1996. [For erratum, see Exp. Hematol. 24, 1540, 1996.]
142. Giroir, B.P., Brown, T., and Beutler, B. Constitutive synthesis of tumor necrosis factor in the thymus. Proc. Natl. Acad. Sci. U.S.A. 89, 4864, 1992.
143. Kohchi, C., Noguchi, K., Tanabe, Y., Nizuno, D.-I., and Soma, G.-I. Constitutive expression of TNF-α and -β genes in mouse embryo: roles of cytokines as regulator and effector on development. Int. J. Biochem. 26, 111, 1994.
144. De Kossodo, S., Grau, G.E., Daneva, T., et al. Tumor necrosis factor alpha is involved in mouse growth and lymphoid tissue development. J. Exp. Med. 176, 1259, 1992.
145. Zúniga-Pflücker, J.C., and Lenardo, M.J. Requirement for TNF-α and IL-1α in fetal thymocyte commitment and differentiation. Science 268, 1906, 1995.
146. Dunon, D., and Imhof, B.A. Mechanisms of thymus homing. Blood 81, 1, 1993.
147. + cytolytic cells. Nature 344, 742, 1990.
148. + T cells. Science 248, 1227, 1990.
149. Vicari, A.P., Figueroa, D.J., Hedrick, J.A., et al. TECK: a novel CC chemokine specifically expressed by thymic dendritic cells and potentially involved in T cell development. Immunity 7, 291, 1997.
150. Imai, T., Baba, M., Nishimura, M., Kakizaki, M., Takagi, S., and Yoshie, O. The T cell-directed CC chemokine TARC is a highly specific biological ligand for CC chemokine receptor 4. J. Biol. Chem. 272, 15036, 1997.
151. Imai, T., Yoshida, T., Baba, M., Nishimura, M., Kakizaki, M., and Yoshie, O. Molecular cloning of a novel T cell-directed CC chemokine expressed in thymus by signal sequence trap using Epstein-Barr virus vector. J. Biol. Chem. 271, 21514, 1996.
152. Touw, I., Pouwels, K., van Agthoren, T., et al. Interleukin-7 is a growth factor of precursor B and T acute lymphoblastic leukemia. Blood 5, 2097, 1990.
153. Ropke, C. Thymic epithelial cell culture. Microsc. Res. Tech. 38, 276, 1997.
154. George, A.J., and Ritter, M.A. Thymic involution with ageing: obsolescence or good housekeeping? Immunol. Today 17, 267, 1996.
155. Aronson, M. Hypothesis: involution of the thymus with aging - programmed and beneficial. Thymus 18, 7, 1991.