Aging Theory Resources

 Back to Aging Info

This is the reference list from the book: The Evolution of Aging 2nd ed T. C. Goldsmith 2006. This short book describes traditional and new theories of aging and is available as a paperback (ISBN: 0978870905), or as a downloadable (PDF) file, (free).  A summary of the evolvability theory of aging and the research implications of various theories is also available at   

 Adaptive (Programmed) Theories of Aging

Adaptive theories of aging are also known as theories of genetically programmed aging or pro-active aging. According to these theories organisms are purposely designed to age or possess some other design characteristic (such as biological suicide) that actively and purposely limits life span. 

Evolution Controversies and the Theory of Aging, T. Goldsmith 2009 Article describes how a 150-year-old controversy regarding the mechanics of evolution is still inhibiting medical research.

Aging, Evolvability, and the Individual Benefit Requirement; Medical Implications of Aging Theory Controversies, Theodore C. Goldsmith, 2008, Journal of Theoretical Biology, DOI: 10.1016/j.jtbi.2008.02.035

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 Moscow State University -- Article suggesting that gradual aging introduces a competitive evolutionary advantage.

Aging Selected for its Own Sake- J. Mitteldorf; Temple University – This article describes an adaptive theory of aging and provides criticisms of the main non-adaptive theories.

 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.

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

Traditional Theories of Aging 

The traditional aging theories hold that aging is not an adaptation or genetically programmed.

 An Unsolved Problem of Biology Medawar, P.B., 1952. H.K. Lewis & Co., London.  Medawar’s paper 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.

 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 (Fifty-one traditional scientists endorsed this position paper to the effect that aging is not and cannot be an evolved characteristic or adaptation.)

 Evolutionary Theories of Aging and Longevity  - L. A. Gavrilov et al; University of Chicago Center on Aging -- Links to many articles by this team - Overview of evolutionary theories of aging including Weismann (adaptive) and mutation accumulation / antagonistic pleiotropy (non-adaptive).  Describes negative impact of some aging theories on research - Cautions that all aging theories are just theories and should not unduly influence research.


 These books are available from many publishers.

 On The Origin of Species - Charles Darwin, 1859

 The Descent of Man – Charles Darwin, 1871 

 Anti-Aging Research

 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.  

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.

 Information on Bighorn Sheep:

 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. ($130). 

 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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