An evolutionary perspective on the mechanisms of immunosenescence

Aw, 2007, Immunosenescence: emerging challenges for an ageing population, Immunology, 120, 435, 10.1111/j.1365-2567.2007.02555.x

Pawelec, 2005, Human immunosenescence: is it infectious?, Immunol. Rev., 205, 257, 10.1111/j.0105-2896.2005.00271.x

Williams, 2003, Antagonistic pleiotropy, mortality source interactions, and the evolutionary theory of senescence, Evolution, 57, 1478, 10.1111/j.0014-3820.2003.tb00356.x

Williams, 2006, The shaping of senescence in the wild, Trends Ecol. Evol., 21, 458, 10.1016/j.tree.2006.05.008

Sorci, 2009, Inflammation and oxidative stress in vertebrate host-parasite systems, Philos. Trans. R. Soc. Lond. B Biol. Sci., 364, 71, 10.1098/rstb.2008.0151

Pawelec, 1998, Impact of aging on innate immunity, J. Leukoc. Biol., 64, 703, 10.1002/jlb.64.6.703

Siegrist, 2009, B-cell responses to vaccination at the extremes of age, Nat. Rev. Immunol., 9, 185, 10.1038/nri2508

Goronzy, 2005, T cell development and receptor diversity during aging, Curr. Opin. Immunol., 17, 468, 10.1016/j.coi.2005.07.020

Medawar, 1952

Kirkwood, 2000, Why do we age?, Nature, 408, 233, 10.1038/35041682

Williams, 1957, Pleiotropy, natural selection and the evolution of senescence, Evolution, 11, 398, 10.2307/2406060

Kirkwood, 1977, Evolution of aging, Nature, 270, 301, 10.1038/270301a0

Stearns, 2000, Experimental evolution of aging, growth, and reproduction in fruitflies, Proc. Natl. Acad. Sci. U. S. A., 97, 3309, 10.1073/pnas.060289597

Rose, 1980, A test of evolutionary theories of senescence, Nature, 287, 141, 10.1038/287141a0

Zwaan, 1995, Direct selection on life span in Drosophila melanogaster, Evolution, 49, 649, 10.2307/2410318

Austad, 1993, Retarded senescence in an insular population of opossums, J. Zool. (Lond.), 229, 695, 10.1111/j.1469-7998.1993.tb02665.x

Bryant, 2004, Comparative studies of senescence in natural populations of guppies, Am. Nat., 163, 55, 10.1086/380650

Ricklefs, 2008, The evolution of senescence from a comparative perspective, Funct. Ecol., 22, 379, 10.1111/j.1365-2435.2008.01420.x

Drenos, 2005, Modelling the disposable soma theory of aging, Mech. Aging Dev., 126, 99, 10.1016/j.mad.2004.09.026

Kirkwood, 1991, Evolution of senescence: late survival sacrificed for reproduction, Philos. Trans. R. Soc. Lond. B. Biol. Sci., 332, 15, 10.1098/rstb.1991.0028

Charlesworth, 2000, Fisher, Medawar, Hamilton and the evolution of aging, Genetics, 156, 927, 10.1093/genetics/156.3.927

Hamilton, 1966, The moulding of senescence by natural selection, J. Theor. Biol., 12, 12, 10.1016/0022-5193(66)90184-6

Sozou, 2004, To age or not to age, Proc. B. Biol. Sci., 271, 457, 10.1098/rspb.2003.2614

Lee, 2003, Rethinking the evolutionary theory of aging: transfers, not births, shape senescence in social species, Proc. Natl. Acad. Sci. U. S. A., 100, 9637, 10.1073/pnas.1530303100

Vaupel, 2004, The case for negative senescence, Theor. Popul. Biol., 65, 339, 10.1016/j.tpb.2003.12.003

Garigan, 2002, Genetic analysis of tissue aging in Caenorhabditis elegans: a role for heat-shock factor and bacterial proliferation, Genetics, 161, 1101, 10.1093/genetics/161.3.1101

Pletcher, 2007, Chemical complexity and the genetics of aging, Annu. Rev. Ecol. Evol. Syst., 38, 299, 10.1146/annurev.ecolsys.38.091206.095634

Zerofsky, 2005, Aging of the innate immune response in Drosophila melanogaster, Aging Cell, 4, 103, 10.1111/j.1474-9728.2005.00147.x

Amdam, 2004, Hormonal control of the yolk precursor vitellogenin regulates immune function and longevity in honeybees, Exp. Gerontol., 39, 767, 10.1016/j.exger.2004.02.010

Holmes, 2004, Declining immunity with age in the wild, Sci. Aging Knowl. Environ., 2004, pe22, 10.1126/sageke.2004.21.pe22

Palacios, 2007, Immunosenescence in some but not all immune components in a free-living vertebrate, the tree swallow, Proc. Biol. Sci., 274, 951, 10.1098/rspb.2006.0192

Adams, 2008, Advanced age in horses affects divisional history of T cells and inflammatory cytokine production, Mech. Ageing Dev., 129, 656, 10.1016/j.mad.2008.09.004

Blount, 2005, Age-related alterations to immune parameters in Labrador retriever dogs, Vet. Immunol. Immunopathol., 108, 399, 10.1016/j.vetimm.2005.06.015

Campbell, 2004, Age-related differences in parameters of feline immune status, Vet. Immunol. Immunopathol., 100, 73, 10.1016/j.vetimm.2004.03.002

Katepalli, 2008, The effect of age and telomere length on immune function in the horse, Dev. Comp. Immunol., 32, 1409, 10.1016/j.dci.2008.06.007

Jackson, 2009, Review series on helminths, immune modulation and the hygiene hypothesis: immunity against helminths and immunological phenomena in modern human populations: coevolutionary legacies?, Immunology, 126, 18, 10.1111/j.1365-2567.2008.03010.x

Lee, 2008, Delayed maturation of an IL-12–producing dendritic cell subset explains the early Th2 bias in neonatal immunity, J. Exp. Med., 205, 2269, 10.1084/jem.20071371

Singh, 1996, Neonatal peptide exposure can prime T cells and, upon subsequent immunization, induce their immune deviation: implications for antibody vs. T cell-mediated autoimmunity, J. Exp. Med, 183, 1613, 10.1084/jem.183.4.1613

Chen, 1995, Enhanced type 2 and diminished type 1 cytokines in neonatal tolerance, Transplantation, 59, 933, 10.1097/00007890-199504150-00002

Forsthuber, 1996, Induction of TH1 and TH2 immunity in neonatal mice, Science, 271, 1728, 10.1126/science.271.5256.1728

Yu, 2008, Fetal exposure to high-avidity TCR ligand enhances expansion of peripheral T regulatory cells, J. Immunol., 181, 73, 10.4049/jimmunol.181.1.73

Miller, 1961, Immunological function of the thymus, Lancet, 2, 748, 10.1016/S0140-6736(61)90693-6

Aspinall, 2000, Thymic atrophy in the mouse is a soluble problem of the thymic environment, Vaccine, 18, 1629, 10.1016/S0264-410X(99)00498-3

Savino, 2006, The thymus is a common target organ in infectious diseases, PLoS Pathog., 2, e62, 10.1371/journal.ppat.0020062

Howard, 1999, Leptin protects mice from starvation-induced lymphoid atrophy and increases thymic cellularity in ob/ob mice, J. Clin. Invest., 104, 1051, 10.1172/JCI6762

Kendall, 2000, The thymus in the mouse changes its activity during pregnancy: a study of the microenvironment, J. Anat., 197, 393, 10.1046/j.1469-7580.2000.19730393.x

Gruver, 2008, Cytokines, leptin, and stress-induced thymic atrophy, J. Leukoc. Biol., 84, 915, 10.1189/jlb.0108025

Ciriaco, 2003, Age-related changes in the avian primary lymphoid organs (thymus and bursa of Fabricius), Microsc. Res. Tech., 62, 482, 10.1002/jemt.10416

Plytycz, 1995, Age-dependent changes in thymuses in the European common frog, Rana temporaria, J. Exp. Zool, 273, 451, 10.1002/jez.1402730602

Danilova, 2004, T cells and the thymus in developing zebrafish, Dev. Comp. Immunol., 28, 755, 10.1016/j.dci.2003.12.003

Lam, 2002, Morphologic transformation of the thymus in developing zebrafish, Dev. Dyn., 225, 87, 10.1002/dvdy.10127

Torroba, 2003, Aging of the vertebrate immune system, Microsc. Res. Tech., 62, 477, 10.1002/jemt.10409

Bertho, 1997, Phenotypic and immunohistological analyses of the human adult thymus: evidence for an active thymus during adult life, Cell. Immunol., 179, 30, 10.1006/cimm.1997.1148

Steinmann, 1985, The involution of the aging human thymic epithelium is independent of puberty, Scand. J. Immunol., 22, 563, 10.1111/j.1365-3083.1985.tb01916.x

Sempowski, 2002, T cell receptor excision circle assessment of thymopoiesis in aging mice, Mol. Immunol., 38, 841, 10.1016/S0161-5890(01)00122-5

Gray, 2006, Developmental kinetics, turnover and stimulatory capacity of thymic epithelial cells, Blood, 108, 3777, 10.1182/blood-2006-02-004531

Ortman, 2002, Molecular characterization of the mouse involuted thymus: aberrations in expression of transcription regulators in thymocyte and epithelial compartments, Int. Immunol., 14, 813, 10.1093/intimm/dxf042

Hale, 2006, Thymic output in aged mice, Proc. Natl. Acad. Sci. U. S. A., 103, 8447, 10.1073/pnas.0601040103

Chen, 2009, Foxn1 is required to maintain the postnatal thymic microenvironment in a dosage-sensitive manner, Blood, 113, 567, 10.1182/blood-2008-05-156265

Min, 2003, Neonates support lymphopenia-induced proliferation, Immunity, 18, 131, 10.1016/S1074-7613(02)00508-3

Schuler, 2004, Cutting edge: IL-7-dependent homeostatic proliferation of CD8+ T cells in neonatal mice allows the generation of long-lived natural memory T cells, J. Immunol., 172, 15, 10.4049/jimmunol.172.1.15

Min, 2006, Reassessing the role of growth hormone and sex steroids in thymic involution, Clin. Immunol., 118, 117, 10.1016/j.clim.2005.08.015

Sutherland, 2005, Activation of thymic regeneration in mice and humans following androgen blockade, J. Immunol., 175, 2741, 10.4049/jimmunol.175.4.2741

George, 1996, Thymic involution with aging: obsolescence or good housekeeping?, Immunol. Today, 17, 267, 10.1016/0167-5699(96)80543-3

Dorshkind, 2009, The aging immune system: is it ever too old to become young again?, Nat. Rev. Immunol., 9, 57, 10.1038/nri2471

Aw, 2008, Architectural changes in the thymus of aging mice, Aging Cell, 7, 158, 10.1111/j.1474-9726.2007.00365.x

Dowling, 2009, Why does the thymus involute? A selection-based hypothesis, Trends Immunol., 30, 295, 10.1016/j.it.2009.04.006

Dowling, 2009, Modelling naive T-cell homeostasis: consequences of heritable cellular lifespan during aging, Nat. Rev. Immuol. Cell Biol

Tyan, 1977, Age-related decrease in mouse T cell progenitors, J. Immunol., 118, 846, 10.4049/jimmunol.118.3.846

Sudo, 2000, Age-associated characteristics of murine hematopoietic stem cells, J. Exp. Med., 192, 1273, 10.1084/jem.192.9.1273

Zhu, 2007, Lymphohematopoietic progenitors do not have a synchronized defect with age-related thymic involution, Aging Cell, 6, 663, 10.1111/j.1474-9726.2007.00325.x

Heng, 2005, Effects of castration on thymocyte development in two different models of thymic involution, J. Immunol., 175, 2982, 10.4049/jimmunol.175.5.2982

Min, 2004, Reduction in the developmental potential of intrathymic T cell progenitors with age, J. Immunol., 173, 245, 10.4049/jimmunol.173.1.245

Gui, 2007, The aged thymus shows normal recruitment of lymphohematopoietic progenitors but has defects in thymic epithelial cells, Int. Immunol., 19, 1201, 10.1093/intimm/dxm095

Clise-Dwyer, 2007, Environmental and intrinsic factors lead to antigen unresponsiveness in CD4+ recent thymic emigrants from aged mice, J. Immunol., 178, 1321, 10.4049/jimmunol.178.3.1321

Eaton, 2008, Bone marrow precursor cells from aged mice generate CD4 T cells that function well in primary and memory responses, J. Immunol., 181, 4825, 10.4049/jimmunol.181.7.4825

Aw, 2009, Is thymocyte development functional in the aged?, Impact Aging, 1, 146, 10.18632/aging.100027

Aw, 2009, Phenotypical and morphological changes in the thymic microenvironment from aging mice, Biogerontology, 10, 311, 10.1007/s10522-008-9182-2

Li, 2003, Cellular mechanism of thymic involution, Scand. J. Immunol., 57, 410, 10.1046/j.1365-3083.2003.01206.x

Mackall, 1998, Thymic function in young/old chimeras: substantial thymic T cell regenerative capacity despite irreversible age-associated thymic involution, Eur. J. Immunol., 28, 1886, 10.1002/(SICI)1521-4141(199806)28:06<1886::AID-IMMU1886>3.0.CO;2-M

Leposavic, 2006, Age-associated changes in CD90 expression on thymocytes and in TCR-dependent stages of thymocyte maturation in male rats, Exp. Gerontol., 41, 574, 10.1016/j.exger.2006.03.006

Aspinall, 2007, Old rhesus macaques treated with Interleukin-7 show increased TREC levels and respond well to influenza vaccination, Rejuvenation Res., 10, 5, 10.1089/rej.2006.9098

Alpdogan, 2006, Keratinocyte growth factor (KGF) is required for postnatal thymic regeneration, Blood, 107, 2453, 10.1182/blood-2005-07-2831

Min, 2007, Sustained thymopoiesis and improvement in functional immunity induced by exogenous KGF administration in murine models of aging, Blood, 109, 2529, 10.1182/blood-2006-08-043794

Rossi, 2007, Keratinocyte growth factor (KGF) enhances postnatal T-cell development via enhancements in proliferation and function of thymic epithelial cells, Blood, 109, 3803, 10.1182/blood-2006-10-049767

Klasing, 2004, The costs of immunity, Acta Zoologica Sinica, 50, 961

Hurtado, 2008, The role of helminthes in human evolution: implications for global health in the 21st century, 152

Råberg, 2002, Basal metabolic rate and the evolution of the adaptive immune system, Proc. Biol. Sci., 269, 817, 10.1098/rspb.2001.1953

Franceschi, 2007, Inflammaging and anti-inflammaging: a systemic perspective on aging and longevity emerged from studies in humans, Mech. Ageing Dev., 128, 92, 10.1016/j.mad.2006.11.016

Van Bodegom, 2007, Regulation of human life histories, Annals N.Y. Acad. Sci., 1100, 84, 10.1196/annals.1395.007

Franceschi, 2000, Inflamm-aging. An evolutionary perspective on immunosenescence, Ann. N. Y. Acad. Sci, 908, 208, 10.1111/j.1749-6632.2000.tb06651.x

Licastro, 2005, Innate immunity and inflammation in aging: a key for understanding age-related diseases, Immun. Ageing, 2, 8, 10.1186/1742-4933-2-8

Vasto, 2009, Inflammation, aging and cancer, Mech. Ageing Dev., 130, 40, 10.1016/j.mad.2008.06.003

Ottaviani, 2008, Ecoimmunology: is there any room for the neuroendocrine system?, Bioessays, 30, 868, 10.1002/bies.20801

Finch, 2004, Inflammatory exposure and historical changes in human life-spans, Science, 305, 1736, 10.1126/science.1092556

Crimmins, 2006, Infection, inflammation, height, and longevity, Proc. Natl. Acad. Sci. U. S. A., 103, 498, 10.1073/pnas.0501470103

Gurven, 2008, Aging and inflammation in two epidemiological worlds, J. Gerontol. A Biol. Sci. Med. Sci., 63, 196, 10.1093/gerona/63.2.196

McDade, 2008, Maintenance versus growth: investigating the costs of immune activation among children in lowland Bolivia, Am. J. Phys. Anthropol., 136, 478, 10.1002/ajpa.20831

McDade, 2005, Life history, maintenance, and the early origins of immune function, Am. J. Hum. Biol., 17, 81, 10.1002/ajhb.20095

Moore, 1997, Season of birth predicts mortality in rural Gambia, Nature, 388, 434, 10.1038/41245

Collinson, 2003, Birth season and environmental influences on patterns of thymic growth in rural Gambian infants, Acta Paediatr., 92, 1014, 10.1111/j.1651-2227.2003.tb02568.x

McDade, 2003, Life history theory and the immune system: steps toward a human ecological immunology, Am. J. Phys. Anthropol., 122, 100, 10.1002/ajpa.10398

Zuk, 2002, Immune defense and host life history, Am. Nat., 160, S9, 10.1086/342131

Fedorka, 2004, Immune suppression and the cost of reproduction in the ground cricket, Allonemobius socius, Evolution, 58, 2478, 10.1111/j.0014-3820.2004.tb00877.x

Helle, 2004, Accelerated immunosenescence in preindustrial twin mothers, Proc. Natl. Acad. Sci. U. S. A., 101, 12391, 10.1073/pnas.0402215101

Raghupathy, 1997, Th 1-type immunity is incompatible with successful pregnancy, Immunol. Today, 18, 478, 10.1016/S0167-5699(97)01127-4

Westendorp, 2001, Optimizing human fertility and survival, Nat. Med., 7, 873, 10.1038/90868

Drenos, 2006, Trade-off mediated effects on the genetics of human survival caused by increasingly benign living conditions, Biogerontology, 7, 287, 10.1007/s10522-006-9027-9

Wilson, 2006, A hallmark of balancing selection is present at the promoter region of interleukin 10, Genes Immun., 7, 680, 10.1038/sj.gene.6364336

Medawar, 1953, Some immunological and endocrinological problems raised by the evolution of viviparity in vertebrates, Symp. Soc. Exp. Biol., 7, 320

Lesser, 2006, Naturally occurring genetic variation in the age-specific immune response of Drosophila melanogaster, Aging Cell, 5, 293, 10.1111/j.1474-9726.2006.00219.x

Salminen, 2008, Activation of innate immunity system during aging: NF-kB signaling is the molecular culprit of inflamm-aging, Ageing Res. Rev., 7, 83, 10.1016/j.arr.2007.09.002

Taub, 2008, Neuroendocrine interactions in the immune system, Cell. Immunol., 252, 1, 10.1016/j.cellimm.2008.05.006

Pawelec, 2002, T cells and aging, Front. Biosci., 7, d59