Programmed Aging Theory Information  Aging Theory Resources


This section provides links or cites for scientific papers and other resources on aging theories, underlying evolution theory, and scientific observations that are germane to aging.

Aging Theories - One-Page Summary 2015

Theories of Biological Aging and Implications for Public Health - Presentation 2019

Programmed Aging and the Emerging Reality of Anti-Aging Medicine - Presentation 2016

Introduction to Biological Aging Theory - E-Book 2020

Aging Theory Questions and Answers 2020

Programmed Aging Theories

Programmed theories of aging are otherwise known as adap-ive or pro-active aging theories. See Aging Theories

Emerging Programmed Aging Mechanisms and their Medical Implications. T. C. Goldsmith Medical Hypotheses 86 (2016) 92-96 PubMed: 26547271 Article compares functional models of programmed and non-programmed theories and describes the different consequences for medical research.

Solving the Programmed/ Non-Programmed Aging Conundrum. T. C. Goldsmith Current Aging Science 2015;8:34-40 Article suggesting there is no current science basis for rejecting programmed aging on evolutionary grounds.

Aging is a Specific Biological Function Rather than the Result of a Disorder in Complex Living Systems: Biochemical Evidence in Support of Weismann's Hypothesis, V. P. Skulachev. 1997. Biochemistry Moscow 62(11) 1191-5  PubMed: 9467841 -- Full text:  This paper describes a programmed theory of aging based on evolvability and suggests that gradual aging could have additional evolutionary benefit relative to biological suicide.

Aging, evolvability, and the individual benefit requirement T. Goldsmith. 2008. Journal of Theoretical Biology 252(4) 764-8. PubMed: 18396295 -- Full text:  This article describes a programmed theory of aging and provides an evolutionary rationale based on evolvability.

Mammal aging: Active and Passive Mechanisms and their Medical Implications T. Goldsmith 2009. Bioscience Hypothesis. 2(2) 59-64  Full text:  This paper compares active (programmed) and passive (non-programmed) aging theories in light of various experimental observations and concludes that active aging provides a much better fit to empirical evidence. Active and passive theories lead to very different approaches for medical intervention so this issue is important to medical research on age-related diseases.


The Evolution of Aging 3nd edition. T. Goldsmith. 2015.  200 p  E-Book (PDF) provides extensive descriptions of programmed and non-programmed aging theories, underlying evolution controversies, historical perspective, and applicable genetics and observational evidence. Evolution of Aging Paperback version. ISBN: 0978870956 180 p illus $5.49 2014


Empirical evidence for various evolutionary hypotheses on species demonstrating increasing mortality with increasing chronological age in the wild. G. Libertini. 2008. The Scientific World Journal 8:182-93, PubMed: 18301820. This article concludes that empirical evidence favors programmed aging.


Senescence as an adaptation to limit the spread of disease. J. Mitteldorf, J. Pepper. 2009. Journal of Theoretical Biology 260(2) 186-95 PubMed: 19481552  Full text: Paper describing an adaptive theory of aging based on group benefit.


 Aging as an Evolved Characteristic – Weismann’s Theory Reconsidered T. C. Goldsmith 2002.  Article discusses evolutionary disadvantages of immortality and suggests that aging is a necessary adaptation. Full text:


Shattered: Medawar's Test Tubes and their Enduring Legacy of Chaos - J. Bowles, Quarterly Review of Biology 73:3-49. (2000)  PubMed: 10790771  Presents extensive criticism of Medawar's 1952 paper which is the basis for traditional theories of aging.


Evolution Controversies and the Theory of Aging. T. Goldsmith 2009. Article describes social and perception factors that affect scientific theories of evolution and dependent aging theories. Full text:


Aging is not a Process of Wear and Tear J. Mitteldorf. Arguments against the popular generic damage theories.


The Evolution of Ageing L. Milewski. Article favoring adaptive (programmed) aging.


Rationale for Complex Programmed Life Span Regulation in Mammals T. Goldsmith. Discussion of different evolutionary value of life concepts and associated theories of aging.

Non-Programmed Theories of Aging 

(Aging is not an adaptation or primary purpose of an organism design characteristic.)

 An Unsolved Problem of Biology Medawar, P.B., 1952. H.K. Lewis & Co., London.  Medawar’s presentation developed the idea that individual members of a non-aging population would have progressively less impact on evolution with increasing calendar age and introduced the mutation accumulation theory of aging.  See summary and critique here.


Pleiotropy, natural selection and the evolution of senescence, Williams, G. 1957.  Evolution 11, 398-411   Williams’ antagonistic pleiotropy theory is one of the most respected of the traditional theories. See summary and critique here.


The evolution of ageing and longevity. T.B.L. Kirkwood & F.R.S. Holliday (1979). Proceedings of the Royal Society of London B 205: 531-546.  The disposable soma theory. See summary and critique here.


No Truth to the Fountain of Youth - Olshansky, Hayflick, and Carnes, Scientific American June 2002 – This article provides warnings against common ineffective anti-aging remedies.   Aging is an “inescapable biological reality”; caused by the accumulation of random damage to the building blocks of life. This paper takes the position that it is "impossible" for aging to be an adaptation or evolved characteristic because of  "the way evolution works." Most proponents of non-programmed aging also base their position entirely on evolution theory considerations and without consideration of empirical evidence. This article is representative of wide consensus in 2002. However, currently there is little specific scientific objection to particular programmed aging theories or their evolutionary basis.

Experimental Research

Regulation of lifespan by sensory perception in Caenorhabditis elegans.  J. Apfeld, C. Kenyon. 1999. Nature 16;402(6763):804-9.  PubMed: 10617200 Research suggesting that life span in the roundworm is regulated by environmental cues -- supports programmed aging theories.


Hormonal Inhibition of Feeding and Death in Octopus: Control by Optic Gland Secretion. J. Wodinsky. 1977 Science 198(4320) 948-951 PubMed: 17787564 Article describing a biological suicide mechanism that involves hormonal signaling and links to the nervous system. Female octopi die of starvation following breeding because they stop eating.


Regulation of Drosophila life span by olfaction and food-derived odors. S. Liebert, et al. 2007. Science 315(5815) 133-7  PubMed: 17272684 Paper describes a complex evolved life span regulation system mediated by external signals.


Reversing the Negative Genomic Effects of Aging with Short-Term Calorie Restriction - S. Spindler, University of California; Scientific World October 12, 2001 -- see overview at


The retardation of aging in mice by dietary restriction: longevity, cancer, immunity and lifetime energy intake. Weindruch R, Walford RL, Fligiel S, Guthrie D., J Nutrition 1986; 116: 641-54.  One of many reports documenting slowing of aging by means of caloric restriction. 


Daf-2, an insulin receptor-like gene that regulates longevity and diapause in Caenorhabditis elegans. Kimura KK, Tissenbaum HA, Liu Y, Ruvkun G. Science 1997; 277: 942.  A report of the discovery of a gene controlling aging in the roundworm.


The Serious Search for an Anti-Aging Pill - M. A. Lane, et al Scientific American Aug 2002, -- Describes experiments with D2G to simulate the effect of caloric restriction. 


The search for anti-aging agents: NIH Interventions Testing Program.


Google Calico Aging Research Company. 

Traditional Evolutionary Mechanics Theory

These books are available from many publishers.

The Origin of Species - Charles Darwin, 1859 Free online:

Huxley J.S. 1942. Evolution: the modern synthesis. Allen & Unwin, London. 2nd ed 1963; 3rd ed 1974.

Alternative Evolutionary Mechanics Theories


Wagner A. 2005. Robustness and evolvability in living systems. (Princeton Studies in Complexity) Princeton University Press. ISBN 0691122407.

Wagner, A., 2005. Robustness, evolvability and neutrality. FEBS Letters 579, 1772–1778.

Wagner G.P. and L. Altenberg 1996. Complex adaptations and the evolution of evolvability. Evolution 50:967–976.

Kin Selection

Hamilton, W. D. (1963). "The evolution of altruistic behavior". American Naturalist 97: 354–356. doi:10.1086/497114. 

Hamilton, W. D. (1964). "The Genetical Evolution of Social Behavior". Journal of Theoretical Biology 7 (1): 1–52. doi:10.1016/0022-5193(64)90038-4. 

Smith, J. M. (1964). "Group Selection and Kin Selection". Nature 201 (4924): 1145–1147. doi:10.1038/2011145a0. 

Group Selection

Wynne-Edwards, V.C. (1962). Animal Dispersion in Relation to Social Behaviour. Edinburgh: Oliver & Boyd.

Wynne-Edwards, V. C. (1986) Evolution Through Group Selection, Blackwell. ISBN 0-632-01541-1

Koeslag, J.H. (2003). Evolution of cooperation: cooperation defeats defection in the cornfield model. J. theor. Biol. 224, 399-410

Wade, M. J. 1977. An experimental study of group selection. Evolution 31:134–153

Gene-Centered Theories

Dawkins, R. (1976) The Selfish Gene. Oxford University Press, Oxford. ISBN 0-19-286092-5

Dawkins, R. (1982) The Extended Phenotype. Oxford University Press, Oxford. ISBN 0-19-288051-9

Pittendrigh, C. (1958) Adaptation, natural selection, and behavior. In A. Roe and G. G. Simpson, eds., Behavior and Evolution, New Haven: Yale University Press, pp 390–416.

Other Resources

Progeria Research Foundation


Werner's Syndrome Overview


Information on Long-lived Animals with Negligible Senescence


 Longevity Records: Life Spans of Mammals, Birds, Amphibians, Reptiles, and Fish, Max Plank Institute, ISBN 87-7838-539-3 -- The oldest lake sturgeon caught so far was 152 years old.

Confirmation on longevity in Sebastes diploproa (Pisces: Scorpaenidae) from 210Pb/226Ra measurements in otoliths. Bennett, J.T. et al. 1982. Maritime Biology. 71, 209-215.  Describes measurements of age in caught specimens of  Rockfish.

Genes VIII Benjamin Lewin, 2004 Oxford University Press ISBN: 019879276X 990pp.  A very comprehensive college level textbook on genetics that has an associated subscription to online updates. Newer versions may be available.


Human Genome Project.  The HGP is an approximately $3 billion government effort to fully sequence the human genetic code and identify all of the genes in human DNA.  The effort which began in 1990 was substantially completed in 2003.  The second link is for online copies of the actual preliminary project reports dated 2001.

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