Evolution of ageing - Knowledge (XXG)

404:. However, telomerase fails us as we age; it becomes less able to repair telomeres, and our whole body starts falling apart. This means that our cells can no longer divide or divide with errors, and some believe that this contributes to the process of aging. New research has also shown that there is an association between telomere shortening and mitochondrial dysfunction. Nevertheless, over-expression of telomerase increases the chances of cancer. If telomeres stay in repair, there is a greater chance of longevity, but there is also more cell division and a greater chance of mutation, which could result in cancer. Therefore, a long-lived cell is just a time bomb. Enhancing telomerase activity is, therefore, not a solution; it only allows the cells to live longer. 94:

likely an organism is to survive and reproduce. It is based on the environment and is also relative to other individuals in the population. Examples of life history traits include; age and size at first reproduction, number of size and offsprings produced, and the period of reproductive lifespan. Organisms put energy into growth, reproduction, and maintenance by following a particular pattern which changes throughout their lifetime due to the trade-offs that exist between the different energy allocations. Investment in current vs future reproduction, for example, comes at the expense of the other. Natural selection, however is not so effective on organisms as they age.

555:. Intrinsic mortality is defined as mortality due to ageing, the physiological decline due to innate processes, whereas extrinsic mortality is the result of environmental factors such as for example predation, starvation, accidents and others. Flying animals such as bats, for example, have fewer predators, and therefore have a low extrinsic mortality. Birds are warm-blooded and similar in size to many small mammals, yet often live 5–10 times as long. They face less predation than ground-dwelling mammals, and thus have lower extrinsic mortality. 233:. Antagonistic pleiotropy on the other hand deals with one gene that creates two traits with one being beneficial and the other detrimental. In essence, this refers to genes that offer benefits early in life, but later accumulate a cost. In other words, antagonistic pleiotropy is when the resultant relationship between two traits is negative. It's when one phenotypic trait positively affects current reproduction at the expense of later accelerated senescence, growth, and maintenance. 192:, which is the translation of genetic information into a phenotypic character. Evolution is the change in a heritable trait in a population across generations since mutations generate variations in the heritable traits; they are considered the raw material for evolution. Therefore, beneficial mutation accumulations during the developmental processes could generate more phenotypic variations, which increases their gene frequency and affect the capacity of phenotypic evolution. 171:

cells, or to not produce toxic metabolism? Why did menopause evolve? Because selection is more efficient on traits that appear early in life. Mutations that have an effect early in life will increase fitness much more than mutations that manifest late. Most people have already reproduced before any disease manifest; this means that parents will pass their alleles to their offsprings before they show any fitness problems, and it is therefore "too late" for selection.

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species, having earlier ages of sexual maturity, have less need for longevity and thus did not evolve or retain the more-effective repair mechanisms. Damage therefore accumulates more rapidly, resulting in earlier manifestations and shorter lifespan. Since there are a wide variety of ageing manifestations that appear to have very different causes, it is likely that there are many different maintenance and repair functions.

367: 5148: 274:, presumes that the body must budget the resources available to it. The body uses resources derived from the environment for metabolism, for reproduction, and for repair and maintenance, and the body must compromise when there is a finite supply of resources. The theory states that this compromise causes the body to reallocate energy to the repair function that causes the body to gradually deteriorate with age. 5158: 455:

population decreases with age as they enter the selection shadow. The model also supports Medewars' theory that due to dangerous and unpredicted conditions in the environment such as diseases, climate changes and predators, many individuals die not too long after sexual maturation. Consequently, the probability of an individual surviving and suffering from age related effects is relatively low.

707:, short stature, and early graying and loss of hair. Once the individual reaches the twenties, there is generally a change in hair color, skin, and voice. The average life expectancy of someone with this disease is around 46 years. This condition can also affect the weight distribution between the arms, legs, and torso. Those who have Werner syndrome are at an increased risk for cataracts, 544: 289:, which lengthens life. However, dietary restriction has not been shown to increase lifetime reproductive success (fitness), because when food availability is lower, reproductive output is also lower. Moreover, calories are not the only resource of possibly limited supply to an organism that could have an effect on multiple dimensions of fitness. 188:

fitness. Previously done experiments have shown that most mutation accumulations are deleterious, and just a few are beneficial. Mutations of genes that interact with one another during the developmental process create biological and, thus, phenotypical diversities. Mutations is genetic information that are expressed among organisms via

256:. His team found that they were able to breed flies that lived more than twice as long as the flies they started with, but to their surprise, the long-lived, inbred flies actually laid more eggs than the short-lived flies. This was another setback for pleiotropy theory, though Rose maintains it may be an experimental artifact. 746:

is a rare autosomal recessive disorder that is characterized by short stature, chromosomal instability, predisposition to cancer, and sun-sensitive skin. Those with Bloom syndrome can also have learning disabilities and have an increased risk of developing chronic obstructive pulmonary disease (COPD)

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and have a series of symptoms that cause abnormalities in the joints, hair, skin, eyes, and face. Most who have the disease only live to about age 13. Although the term progeria applies strictly speaking to all diseases characterized by premature aging symptoms, and is often used as such, it is often

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Medewar developed a model that highlights this, showing the decrease in the survival rate of a population as an individual ages, however the reproduction rate stays constant. The reproduction probability typically peaks during sexual maturity and decreases as an individual ages, while the rest of the

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Selective shadowing is one of the evolutionary theories of aging based on the presumption that selection of an individual generally decreases once they essentially pass the sexual mature phase. As a result, this forms a shadow without the account of sexual fitness, which is no longer considered as an

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postulates that DNA damage is ubiquitous in the biological world and is the primary cause of ageing. The theory is based on the idea that ageing occurs over time due to the damage of the DNA. As an example, studies of mammalian brain and muscle have shown that DNA repair capability is relatively high

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in 1957. Williams noted that senescence may be causing many deaths even if animals are not 'dying of old age.' He began his hypothesis with the idea that ageing can cause earlier senescence due to the competitive nature of life. Even a small amount of ageing can be fatal; hence natural selection does

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is low for traits that would maintain viability past the time when most animals would have died anyway. Metabolic diseases come along due to the low demand for physical activity in modern civilization compared to times where humans had to forage in the wild for survival. With the selective shadow now

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Redwood AB, Perkins SM, Vanderwaal RP, Feng Z, Biehl KJ, Gonzalez-Suarez I, Morgado-Palacin L, Shi W, Sage J, Roti-Roti JL, Stewart CL, Zhang J, Gonzalo S (27 October 2014). "A dual role for A-type lamins in DNA double-strand break repair". Cell Cycle. 10 (15): 2549–2560. doi:10.4161/cc.10.15.16531.

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When examining the body-size vs. lifespan relationship, one also observes that predatory mammals tend to live longer than prey mammals in a controlled environment, such as a zoo or nature reserve. The explanation for the long lifespans of primates (such as humans, monkeys, and apes) relative to body

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The somatic mutation theory of ageing states that accumulation of mutations in somatic cells is the primary cause of aging. A comparison of somatic mutation rate across several mammal species found that the total number of accumulated mutations at the end of lifespan was roughly equal across a broad

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Mutations happen, and they are completely random with respect to a need in the environment and fitness. Mutations can either be beneficial in which they increase an organism's fitness, neutral in which they do not affect an organism's fitness or deleterious where they negatively affect an organism's

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have shown an inverse relationship between the mean optimal age at maturity and mutation rates per gene. Mutation accumulation affects the allocation of energy, and time that are directed towards growth and reproduction over the lifetime of an organism, especially the period of reproductive lifespan

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is a process that allows organisms to better adapt to the environment, it is the survival of the fittest which are predicted to produce more offsprings. Natural selection acts on life history traits in order to optimize reproductive success and lifetime fitness. Fitness in this context refers to how

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Lenart and Vašku (2016) have also invoked evolvability as the main mechanism driving evolution of ageing. However, they proposed that even though the actual rate of aging can be an adaptation the aging itself is inevitable. In other words, evolution can change the speed of aging but some ageing no

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A non-programmed theory of mammal ageing states that different species possess different capabilities for maintenance and repair. Longer-lived species possess many mechanisms for offsetting damage due to causes such as oxidation, telomere shortening, and other deteriorative processes. Shorter-lived

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A caveat to this theory suggests that this reallocation of energy is based on time instead of limiting resources. This concept focuses on the evolutionary pressure to reproduce in a set, optimal time period that is dictated by age and ecological niche. The way that this is successful is through the

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The two theories; non-adaptive, and adaptive, are used to explain the evolution of senescence, which is the decline in reproduction with age. The non-adaptive theory assumes that the evolutionary deterioration of human age occurs as a result of accumulation of deleterious mutations in the germline.

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Liu B, Wang J, Chan KM, Tjia WM, Deng W, Guan X, Huang J-d, Li KM, Chau PY, Chen DJ, Pei D, Pendas AM, Cadiñanos J, López-Otín C, Tse HF, Hutchison C, Chen J, Cao Y, Cheah KSE, Tryggvason K, Zhou Z (26 June 2005). "Genomic instability in laminopathy-based premature aging". Nature Medicine. 11 (7):

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Although antagonistic pleiotropy is a prevailing theory today, this is largely by default, and has not been well verified. Research has shown that this is not true for all genes and may be thought of as partial validation of the theory, but it cuts the core premise: that genetic trade-offs are the

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concept. The development of human civilization has shifted the selective shadow as the conditions that humans now live in include improved quality of victuals, living conditions, and healthcare. This improved healthcare includes modern medicine such as antibiotics and new medical technology. A few

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Those who believe in the idea that ageing is an unavoidable side effect of some necessary function (antagonistic pleiotropy or disposable soma theories) logically tend to believe that attempts to delay ageing would result in unacceptable side effects to the necessary functions. Altering ageing is

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There is no theory that explains how these deleterious mutations affect fitness on different ages and the evolution of senescence. Their idea was that ageing was a matter of neglect, as nature is a highly competitive place. Almost all animals die in the wild from predators, disease, or accidents,

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In the same way, many beneficial mutations are selected against if they have a positive effect on an individual later on in life. For instance if a beneficial or deleterious mutation occurs only after an individual's reproductive phase, then it will not affect fitness, which therefore can not be

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Senescence is considered a by-product of physiology because our cell metabolism creates products that are toxic, we get mutations when we age, and we don't have enough stem cells that regenerate. Why did selection not find and favor mutations in ways that allow us, for example, to regenerate our

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have shown that mutation accumulation drives the combination of alleles which have "age-specific additive effects" that cause a decline in stress response and ultimately an age-related decline in fitness. The number of germ cell divisions per generation is variable among lineages, and relates to

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Those who believe in programmed ageing suppose that ways might be found to interfere with the operation of the part of the ageing mechanism that appears to be common to multiple symptoms, essentially "slowing down the clock" and delaying multiple manifestations. Such effect might be obtained by

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Group selection is based on the idea that all members of a given group will either succeed or fail together depending on the circumstance. With this mechanism, genetic drift occurs collectively to all in the group and sets them apart from other groups of its own species. This is different than

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Just like DNA mutation and expression have phenotypic effects on organisms, DNA damage and mutation accumulation also have phenotypic consequences in older humans. Damage to macromolecules such as DNA, RNA, and proteins along with the deterioration of tissues and organs are the basis of aging.

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suggests that the repair mechanisms are controlled by a common control mechanism capable of sensing conditions, such as caloric restriction, and may be responsible for lifespan in particular species. In this theory, the survival techniques are based on control mechanisms instead of individual

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Yang (2013)'s model is also based on the idea that ageing accelerates the accumulation of novel adaptive genes in local populations. However, Yang changed the terminology of "evolvability" into "genetic creativity" throughout his paper to facilitate the understanding of how ageing can have a

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evolution in a modern theoretical framework. In 1889, he theorized that ageing was part of life's program to make room for the next generation in order to sustain the turnover that is necessary for evolution. The idea that the ageing characteristic was selected (an adaptation) because of its

653:. Although progeria can cause physical abnormalities on a child, it does not impact their motor skills or intellectual advancement. Those who have HGPS are prone to suffer from neurological and cardiovascular disorders. HGPS is caused by a point mutation in the gene that encodes 54:) suggest that environmental factors, such as predation, accidents, disease, and/or starvation, ensure that most organisms living in natural settings will not live until old age, and so there will be very little pressure to conserve genetic changes that increase longevity. 201:

range of lifespans. The authors state that this strong relationship between somatic mutation rate and lifespan across different mammalian species suggests that evolution may constrain somatic mutation rates, perhaps by selection acting on different DNA repair pathways.

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Individual organisms are ordinarily mortal; they age and die, while the germlines which connect successive generations are potentially immortal. The basis for this difference is a fundamental problem in biology. The Russian biologist and historian

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Skulachev (1997) has suggested that programmed ageing assists the evolution process by providing a gradually increasing challenge or obstacle to survival and reproduction, and therefore enhancing the selection of beneficial characteristics.

447:"The key conceptual insight that allowed Medawar, Williams, and others, to develop the evolutionary theory of aging is based on the notion that the force of natural selection, a measure of how effectively selection acts on survival rate or 416:

Theories, such as Weismann's "programmed death" theory, suggest that deterioration and death due to ageing are a purposeful result of an organism's evolved design, and are referred to as theories of programmed ageing or adaptive ageing.

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Evolvability is the concept that a species should profit from faster genetic adaptation to its present environment. In the following examples, this is used to argue that eliminating old individuals might benefit the species overall.

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guard their grandchildren; there is cooperative breeding in some mammals, many insects and about 200 species of birds; sex differences in the survival of anthropoid primates tend to correlate with the care to offspring; or an

699:. It affects about 1 in 200,000 people in the United States. This syndrome starts to affect individuals during the teenage years, preventing teens from growing at puberty. There are four common traits of Werner's syndrome: 625:

Progeroid syndromes are genetic diseases that are linked to premature aging. Progeroid syndromes are characterized by having features that resemble those of physiological aging such as hair loss and cardiovascular disease.

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have shown that the age of expression of novel deleterious mutations, defines the effects they contribute on mortality. Overall, however; although their frequency increases, their effects and variation decreases with age.

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Cagan, Alex; Baez-Ortega, Adrian; Brzozowska, Natalia; Abascal, Federico; Coorens, Tim H. H.; Sanders, Mathijs A.; Lawson, Andrew R. J.; Harvey, Luke M. R.; Bhosle, Shriram; Jones, David; Alcantara, Raul E. (April 2022).

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Those believing in default theories of multiple maintenance mechanisms tend to believe that ways might be found to enhance the operation of some of those mechanisms. Perhaps they can be assisted by antioxidants or other

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due to the fact that mutation accumulation accelerates senescence, this means that organisms must reach the optimum age of maturity at a younger age as their reproductive lifespan is shortened with accumulated mutation.

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applied specifically in reference to Hutchinson–Gilford progeria syndrome (HGPS). Children diagnosed with HGPS develop prominent facial features such as a small face, thin lips, small chin, and protruding

359:, which is also known as cyclin-dependent kinase inhibitor 1. This ensures that the cell cannot enter the next stage of cell division unless the DNA damage is repaired. However, the p21 cells can trigger 517:

Goldsmith (2008) proposed that though increasing the generation rate and evolution rate is beneficial for a species, it is also important to limit lifespan so older individuals will not dominate the

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Gavrilova NS, Gavrilov LA, Evdokushkina GN, Semyonova VG, Gavrilova AL, Evdokushkina NN, et al. (August 1998). "Evolution, mutations, and human longevity: European royal and noble families".

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Skulachev VP (November 1997). "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".

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complex, combined with evidence that WRN deficient cells produce extensive deletions at sites of joining of non-homologous DNA ends, suggests a role for WRN protein in the DNA repair process of

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have high telomerase activity, they live long, and were thought by some to never get cancer; and therefore possibly be an exception to this hypothesis. Naked mole rats do get cancer, however.

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and causes the progressive loss of cellular and tissue functions that define aging. As a response to DNA damage, one of the responses triggered by oxidative stress is the activation of the

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Gavrilova NS, Gavrilov LA, Semyonova VG, Evdokushkina GN (June 2004). "Does exceptional human longevity come with a high cost of infertility? Testing the evolutionary theories of aging".

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are two factors that manifest with DNA damage; therefore, we need to understand the change in the association between DNA damage and DNA repair as we age in order to be aware of

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age-related mutated alleles is known as mutation accumulation. Note that somatic mutations are not heritable, they are only a source of developmental variation. Studies done on

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individual ages. This supports the idea that the force of natural selection declines as a function of age, which was first introduced by Peter B. Medewar and J.B.S Haldane.

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ageing could evolve if there is little migration among populations. Weismann later abandoned his theory and after some time followed up with his "programmed death" theory.

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allocation of time and energy in damage repair at the cellular level resulting in an accumulation of damage and a decreased lifespan relative to organisms with longer

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is a rare autosomal recessive disorder that affects the skin. It is characterized by the sparse hair, juvenile cataracts, skeletal abnormalities, and stunted growth.

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infant is often attended by more than 10 people. Lee developed a formal theory integrating selection due to transfers (at all ages) with selection due to fertility.

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that cause acceleration of many or most symptoms of ageing during childhood. It affects about 1 in 4-8 million births. Those who have this disease are known for

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Charlesworth B (May 2001). "Patterns of age-specific means and genetic variances of mortality rates predicted by the mutation-accumulation theory of ageing".

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Thompson LH, Schild D. Recombinational DNA repair and human disease. Mutat Res. 2002 Nov 30;509(1-2):49-78. doi: 10.1016/s0027-5107(02)00224-5. PMID 12427531

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Drost JB, Lee WR (1995). "Biological basis of germline mutation: comparisons of spontaneous germline mutation rates among drosophila, mouse, and human".

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Gavrilova NS, Gavrilov LA (2005). "Human longevity and reproduction: An evolutionary perspective.". In Voland E, Chasiotis A, Schiefenhoevel W (eds.).

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Rifkin SA, Houle D, Kim J, White KP (November 2005). "A mutation accumulation assay reveals a broad capacity for rapid evolution of gene expression".

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sequences that protect the ends of our chromosome; they are sensitive to oxidative stress and degrade during chromosomal replication. Telomerase is a

363:. Apoptosis or programmed cell death is associated with gradual degradation of the immune system, skeletal muscle, and aging-associated malfunction. 5133: 4588: 175:

These deleterious mutations start expressing themselves late in life, by the time we are weak/wobbly and have already reproduced, this means that

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Site provides comprehensive information on programmed ageing, the programmed/non-programmed controversy, and underlying evolution controversies.

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Gensler HL (1981). "Low level of U.V.-induced unscheduled DNA synthesis in postmitotic brain cells of hamsters: possible relevance to aging".

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Goldsmith TC (June 2008). "Aging, evolvability, and the individual benefit requirement; medical implications of aging theory controversies".

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Mortality is the number of deaths, in a particular group, over a specific time period. There are two types of mortality: intrinsic and

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is a homozygous or heterozygous mutation that results in short stature, abnormalities in head size, and slow growth and development.

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which lowers the average age of death. Therefore, there is not much reason why the body should remain fit for the long haul because

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during early development when cells are dividing mitotically, but declines substantially as cells enter the post-mitotic state.

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selected against. Subsequently, these later mutations and effects are considered to be in the "shadow region" of selection."

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Werner syndrome, also known as "adult progeria", is another single-gene genetic disease. it is caused by a mutation in the

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Lampidis TJ, Schaiberger GE (December 1975). "Age-related loss of DNA repair synthesis in isolated rat myocardial cells".

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fooling a sense function. One such effort is an attempt to find a "mimetic" that would "mime" the anti-ageing effect of

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Leroi AM, Chippindale AK, Rose MR (August 1994). "Long-term laboratory evolution of a genetic life-history tradeoff in

1283:"Fitness change in relation to mutation number in spontaneous mutation accumulation lines of Chlamydomonas reinhardtii" 5197: 5161: 4755: 3650: 384: 347:

The effect of reducing expression of DNA repair capability is increased accumulation of DNA damage. This impairs gene

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Describes empirical data, evolutionary rationale, and historical perspective supporting programmed ageing in mammals.

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Medvedev ZA (December 1981). "On the immortality of the germ line: genetic and biochemical mechanism. A review".

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Lenart P, Bienertová-Vašků J (August 2017). "Keeping up with the Red Queen: the pace of aging as an adaptation".

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Delaney, M. A.; Ward, J. M.; Walsh, T. F.; Chinnadurai, S. K.; Kerns, K.; Kinsel, M. J.; Treuting, P. M. (2016).

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deleterious effect was largely discounted for much of the 20th century, but a theoretical model suggests that

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will instead strongly favor genes which ensure early maturation and rapid reproduction, and the selection for

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Article compares programmed and non-programmed maintenance theories of ageing in light of empirical evidence.

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indicate the presence of unique information maintenance and restoration processes at the different stages of

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Species-specific rates of aging are due to deleterious changes which manifest after the reproductive phase. "

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for long lifespan. Based on antagonistic pleiotropy, Rose expected that this would surely reduce their

1572:"Antagonistic pleiotropy as a widespread mechanism for the maintenance of polymorphic disease alleles" 5033: 4928: 4862: 4663: 4578: 4494: 4406: 4389: 4316: 4306: 4016: 3924: 3908: 3861: 3489: 2757: 2675: 2555: 2504: 2268: 1755: 1518: 1458: 1403: 1348: 1101: 901: 731:. WRN protein also appears to play a role in resolving recombination intermediate structures during 5192: 5096: 5054: 5004: 4923: 4771: 4763: 4693: 4673: 4553: 4463: 4216: 4159: 4021: 4004: 3982: 3836: 3685: 3502: 787: 581: 552: 538: 448: 286: 5157: 600:; he saw these as processes within the germ cells that were capable of restoring the integrity of 117:

genome size; for humans; 401 germ cell divisions occur per generation in males and 31 in females.

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Chen D, Guarente L (February 2007). "SIR2: a potential target for calorie restriction mimetics".

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Theories of ageing affect efforts to understand and find treatments for age-related conditions:

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which promote molecular and cellular self-maintenance will decline with age for most organisms.

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Williams GC (December 1957). "Pleiotropy, Natural Selection, and the Evolution of Senescence".

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and have increased sensitivity to DNA damaging agents. In HGPS, the inability to adequately

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control". Damage to mtDNA is therefore another contributing factor to phenotypes related to

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size is that they manage to achieve lower extrinsic mortality due to their intelligence.

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Gensler HL, Bernstein H (September 1981). "DNA damage as the primary cause of aging".

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is permanent unless a mutation that modifies the effects of the primary locus occurs.

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Sahin E, Colla S, Liesa M, Moslehi J, Müller FL, Guo M, et al. (February 2011).

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therefore "impossible", and study of ageing mechanisms is of only academic interest.

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maintenance mechanism, which you see in the non-programmed theory of mammal ageing.

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Grandmotherhood - The Evolutionary Significance of the Second Half of Female Life

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Hamilton WD (September 1966). "The moulding of senescence by natural selection".

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replicative and other synthetic systems alone cannot explain the immortality of

487: 84: 3191: 3174: 2769: 2668:

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2480: 1834: 1817: 1530: 1451:

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Often also postreproductive individuals make intergenerational transfers:

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3228: 3215: 3175:"A report of two cases of Werner's syndrome and review of the literature" 2173: 2106: 2088: 921: 720: 635: 401: 389: 3571: 2280: 1744:"Disposable Soma Theory and the Evolution of Maternal Effects on Ageing" 1415: 336:

and develop lifestyles that could possibly promote a healthy life span.

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Work that describes Weismann's theory about making room for the young.

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Williams eventually proposed his own hypothesis called antagonistic

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cannot act on them because reproduction has ended. Studies done on

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shorter-term benefit than the word "evolvability" would imply.

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from the types of damage that cause irreversible ageing in

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Study of the evolutionary development of ageing processes

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in 1952. This theory formed in the previous decade with

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without having to actually radically restrict diet.

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The classical theories of evolution ( 3433: 3423: 3144: 3142: 3086: 3056: 3054: 3052: 3033:"Hutchinson-Gilford progeria syndrome" 2976:"Hutchinson-Gilford progeria syndrome" 1570:Carter AJ, Nguyen AQ (December 2011). 563:Potential immortality of the germ line 4897:Dialogues Concerning Natural Religion 3447:"Modern Biological Theories of Aging" 3222:. 15 Spec No 2 (suppl_2): R151–R161. 1330: 1328: 1326: 1087: 1085: 1083: 801:List of life extension-related topics 7: 3666:The Case for Programmed Mammal Aging 2861:Mechanisms of Ageing and Development 2589:Flatt T, Schmidt PS (October 2009). 1867:Mechanisms of Ageing and Development 1183:"Horizons in the evolution of aging" 1181:Flatt T, Partridge L (August 2018). 1041: 1039: 887: 885: 735:repair of DNA double-strand breaks. 529:matter how slow will always occur. 4302:Evolutionary developmental biology 3726:Antagonistic pleiotropy hypothesis 3408:10.1016/b978-0-12-420190-3.00023-5 1672:10.1111/j.1558-5646.1994.tb05309.x 860:10.1111/j.1558-5646.1957.tb02911.x 711:, different types of cancers, and 211:Antagonistic pleiotropy hypothesis 25: 5156: 5147: 5146: 3847:List of longest-living organisms 3632:The Evolutionary Theory of Aging 2835:"NCI Dictionary of Cancer Terms" 2469:Journal of Bioscience Hypotheses 2330:10.32607/20758251-2017-9-4-31-41 2208:"Programmed cell death in aging" 1801:Weindruch R, Walford IL (1986). 572:considered that the accuracy of 4959:Extended evolutionary synthesis 4148:Gene-centered view of evolution 3065:(Second ed.). Boca Raton. 2908:Disease Models & Mechanisms 1987:The Quarterly Review of Biology 944:An Unsolved Problem of Biology. 547:Constant failure rate over time 412:Programmed maintenance theories 266:Disposable soma theory of aging 5087:Hologenome theory of evolution 4954:History of molecular evolution 4180:Evolutionarily stable strategy 4069:Last universal common ancestor 3402:. Elsevier. pp. 383–396. 3179:Journal of Orthopaedic Surgery 2750:Journal of Theoretical Biology 2548:Journal of Theoretical Biology 1631:10.1016/B0-08-043076-7/03374-X 1094:Journal of Theoretical Biology 1: 5203:Theories of biological ageing 4881:Renaissance and Enlightenment 3128:780–785. doi:10.1038/nm1266. 2644:Evolutionary Ecology Research 1026:Evolutionary Ecology Research 946:Published for the college by 5092:Missing heritability problem 4719:Gamete differentiation/sexes 3746:Free-radical theory of aging 3373:10.1016/j.molmed.2006.12.004 3361:Trends in Molecular Medicine 2873:10.1016/0047-6374(81)90052-X 2607:10.1016/j.bbagen.2009.07.010 2568:10.1016/0022-5193(66)90184-6 2134:10.1016/0014-4827(75)90276-1 2042:10.1016/0531-5565(81)90046-2 1769:10.1371/journal.pone.0145544 1362:10.1371/journal.pone.0034146 594:recombination during meiosis 127:Mutation accumulation theory 3336:"Rothmund-Thomson syndrome" 2077:The Journal of Cell Biology 1954:10.1101/cshperspect.a025130 1250:10.1534/genetics.108.088526 642:single-gene genetic disease 385:Telomeres in the cell cycle 136:The first modern theory of 5224: 4724:Life cycles/nuclear phases 4276:Trivers–Willard hypothesis 3736:DNA damage theory of aging 3192:10.1177/230949900301100222 2770:10.1016/j.jtbi.2008.02.035 2481:10.1016/j.bihy.2008.12.002 2206:Tower J (September 2015). 2122:Experimental Cell Research 1835:10.1177/019262339602400618 1805:. Springfield, IL: Thomas. 1531:10.1038/s41586-022-04618-z 1005:. Oxford: Clarendon Press. 766: 733:homologous recombinational 729:non-homologous end joining 688: 659:non-homologous end joining 633: 536: 502: 471: 436: 378: 341:DNA damage theory of aging 299:DNA damage theory of aging 296: 263: 208: 124: 5142: 4222:Parent–offspring conflict 4027:Earliest known life forms 3915: 3791:Stem cell theory of aging 3559:TheScientificWorldJournal 2839:National Cancer Institute 2805:10.1007/s10522-016-9674-4 2224:10.1016/j.arr.2015.04.002 1879:10.1016/j.mad.2005.03.012 1200:10.1186/s12915-018-0562-z 979:10.1016/j.arr.2019.100947 757:Rothmund–Thomson syndrome 739:Other progeroid syndromes 375:Telomere theory of ageing 244:In breeding experiments, 140:ageing was formulated by 5075:Cultural group selection 4939:The eclipse of Darwinism 4911:On the Origin of Species 4886:Transmutation of species 3220:Human Molecular Genetics 3001:A dictionary of genetics 2723:Biochemistry. Biokhimiia 2380:10.1177/0300985816630796 2030:Experimental Gerontology 1589:10.1186/1471-2350-12-160 681:-based premature aging. 663:homologous recombination 5080:Dual inheritance theory 4919:History of paleontology 3766:Network theory of aging 3672:Life Table for USA 2005 3636:João Pedro de Magalhães 3512:10.1196/annals.1297.095 3340:Genetics Home Reference 3292:Genetics Home Reference 3154:Genetics Home Reference 3037:Genetics Home Reference 2980:Genetics Home Reference 2689:10.1073/pnas.1530303100 2517:10.1196/annals.1354.002 2212:Ageing Research Reviews 1656:Drosophila melanogaster 1472:10.1073/pnas.0703349104 966:Ageing Research Reviews 293:DNA damage/error theory 235:Antagonistic pleiotropy 205:Antagonistic pleiotropy 181:Drosophila melanogaster 113:Drosophila melanogaster 100:antagonistic pleiotropy 66:Theories and hypotheses 48:antagonistic pleiotropy 4768:Punctuated equilibrium 4089:Non-adaptive radiation 4037:Evolutionary arms race 3857:Regeneration (biology) 3817:Biological immortality 3445:Jin K (October 2010). 2662:Lee RD (August 2003). 2161:Nucleic Acids Research 1720:10.4161/cc.10.22.18302 1625:. pp. 13897–902. 1114:10.1006/jtbi.2001.2296 671:chromosome aberrations 548: 371: 260:Disposable soma theory 241:root cause of ageing. 5060:Evolutionary medicine 4934:Mendelian inheritance 4642:Biological complexity 4630:Programmed cell death 4322:Phenotypic plasticity 4042:Evolutionary pressure 4032:Evidence of evolution 3930:Timeline of evolution 3776:Programmed cell death 3761:Negligible senescence 3546:Encyclopedia of Aging 3136:. S2CID 11798376 2638:Mitteldorf J (2006). 2437:10.1093/gerona/glw047 1822:Toxicologic Pathology 1149:10.1002/em.2850250609 806:Negligible senescence 546: 369: 121:Mutation accumulation 96:Mutation accumulation 44:mutation accumulation 5188:Evolutionary biology 5034:Teleology in biology 4929:Blending inheritance 4307:Genetic assimilation 4170:Artificial selection 3909:Evolutionary biology 3862:Rejuvenation (aging) 3642:Programmed-Aging.Org 3061:McDonald RB (2019). 2367:Veterinary Pathology 2089:10.1083/jcb.74.1.274 1816:Weindruch R (1996). 1576:BMC Medical Genetics 334:age-related diseases 5097:Molecular evolution 5055:Ecological genetics 4924:Transitional fossil 4714:Sexual reproduction 4554:endomembrane system 4483:pollinator-mediated 4439:dolphins and whales 4217:Parental investment 3837:Indefinite lifespan 3741:Evolution of ageing 3711:(biology of ageing) 3572:10.1100/tsw.2002.96 3494:2004NYASA1019..513G 2762:2008JThBi.252..764G 2680:2003PNAS..100.9637L 2560:1966JThBi..12...12H 2509:2006NYASA1067....1H 2281:10.1038/nature09787 2273:2011Natur.470..359S 1760:2016PLoSO..1145544V 1523:2022Natur.604..517C 1463:2007PNAS..10412235N 1457:(30): 12235–12242. 1445:Nei M (July 2007). 1416:10.1038/nature04114 1408:2005Natur.438..220R 1353:2012PLoSO...734146D 1143:(Suppl 26): 48–64. 1106:2001JThBi.210...47C 1001:Weismann A (1889). 942:Medawar PB (1952). 906:1977Natur.270..301K 788:calorie restriction 715:. The finding that 621:Progeroid syndromes 582:sexual reproduction 553:extrinsic mortality 539:Life history theory 484:bottlenose dolphins 310:cellular metabolism 287:caloric restriction 32:evolution of ageing 5198:Theories of ageing 5070:Cultural evolution 4185:Fisher's principle 4114:Handicap principle 4104:Parallel evolution 3968:Adaptive radiation 3653:2007-07-25 at the 3229:10.1093/hmg/ddl214 3108:PMC 3180193. 2920:10.1242/dmm.024711 2174:10.1093/nar/gkm744 675:repair DNA damages 570:Zhores A. Medvedev 549: 372: 308:(mtDNA) regulates 217:George C. Williams 164:selection pressure 18:Evolution of aging 5170: 5169: 4786:Uniformitarianism 4739:Sex-determination 4244:Sexual dimorphism 4239:Natural selection 4143:Unit of selection 4109:Signalling theory 3875: 3874: 3852:Maximum life span 3807:Adaptive mutation 3451:Aging and Disease 3417:978-0-12-420190-3 3150:"Werner syndrome" 3072:978-0-8153-4567-1 3010:978-0-19-937686-5 2729:(11): 1191–1195. 2674:(16): 9637–9642. 2267:(7334): 359–365. 2167:(22): 7475–7484. 1713:(22): 3853–3856. 1640:978-0-08-043076-8 1517:(7906): 517–524. 1402:(7065): 220–223. 1299:10.1111/evo.13360 1293:(12): 2918–2929. 1065:10.1111/evo.13408 900:(5635): 301–304. 751:Cockayne syndrome 646:failure to thrive 463:Natural selection 326:Neurodegeneration 306:Mitochondrial DNA 177:Natural selection 91:Natural selection 56:Natural selection 30:Enquiry into the 16:(Redirected from 5215: 5160: 5150: 5149: 4949:Modern synthesis 4709:Multicellularity 4704:Mosaic evolution 4589:auditory ossicle 4271:Social selection 4254:Flowering plants 4249:Sexual selection 3902: 3895: 3888: 3879: 3786:Selection shadow 3771:Plant senescence 3756:Immunosenescence 3702: 3695: 3688: 3679: 3615: 3594: 3584: 3574: 3549: 3540: 3531: 3505: 3476: 3466: 3441: 3435: 3431: 3429: 3421: 3385: 3384: 3356: 3350: 3349: 3347: 3346: 3332: 3326: 3325: 3323: 3322: 3308: 3302: 3301: 3299: 3298: 3288:"Bloom syndrome" 3284: 3278: 3277: 3275: 3274: 3260: 3254: 3251: 3242: 3241: 3231: 3211: 3205: 3204: 3194: 3170: 3164: 3163: 3161: 3160: 3146: 3137: 3125: 3116: 3105: 3099: 3098: 3092: 3084: 3063:Biology of aging 3058: 3047: 3046: 3044: 3043: 3029: 3023: 3022: 2996: 2990: 2989: 2987: 2986: 2972: 2966: 2965: 2963: 2962: 2948: 2942: 2941: 2931: 2899: 2893: 2892: 2856: 2850: 2849: 2847: 2846: 2831: 2825: 2824: 2788: 2782: 2781: 2745: 2739: 2738: 2718: 2712: 2711: 2701: 2691: 2659: 2653: 2651: 2635: 2629: 2628: 2618: 2586: 2580: 2579: 2543: 2537: 2536: 2492: 2486: 2484: 2464: 2458: 2457: 2439: 2415: 2409: 2408: 2382: 2358: 2352: 2351: 2341: 2309: 2303: 2302: 2292: 2252: 2246: 2245: 2235: 2218:(Pt A): 90–100. 2203: 2197: 2196: 2186: 2176: 2152: 2146: 2145: 2117: 2111: 2110: 2100: 2068: 2062: 2061: 2025: 2019: 2018: 1982: 1976: 1975: 1965: 1933: 1927: 1926: 1906: 1900: 1898: 1862: 1856: 1855: 1837: 1813: 1807: 1806: 1798: 1792: 1791: 1781: 1771: 1739: 1733: 1732: 1722: 1698: 1692: 1691: 1666:(4): 1244–1257. 1651: 1645: 1644: 1618: 1612: 1611: 1601: 1591: 1567: 1561: 1560: 1550: 1501: 1495: 1494: 1484: 1474: 1442: 1436: 1435: 1391: 1385: 1384: 1374: 1364: 1332: 1321: 1320: 1310: 1278: 1272: 1271: 1261: 1244:(4): 2061–2073. 1229: 1223: 1222: 1212: 1202: 1178: 1169: 1168: 1132: 1126: 1125: 1089: 1078: 1077: 1067: 1043: 1034: 1033: 1020:Yang JN (2013). 1017: 1008: 1006: 998: 992: 991: 981: 957: 951: 940: 934: 933: 914:10.1038/270301a0 889: 880: 879: 843: 439:Selection shadow 433:Selective shadow 318:oxidative stress 150:selection shadow 146:J. B. S. Haldane 21: 5223: 5222: 5218: 5217: 5216: 5214: 5213: 5212: 5173: 5172: 5171: 5166: 5138: 5065:Group selection 5038: 4963: 4867: 4794: 4756:Tempo and modes 4750: 4605: 4509: 4326: 4285: 4161: 4154: 4131:Species complex 3944: 3935:History of life 3911: 3906: 3876: 3871: 3795: 3712: 3706: 3655:Wayback Machine 3623: 3618: 3597: 3552: 3543: 3534: 3479: 3444: 3432: 3422: 3418: 3397: 3393: 3391:Further reading 3388: 3358: 3357: 3353: 3344: 3342: 3334: 3333: 3329: 3320: 3318: 3310: 3309: 3305: 3296: 3294: 3286: 3285: 3281: 3272: 3270: 3262: 3261: 3257: 3252: 3245: 3213: 3212: 3208: 3172: 3171: 3167: 3158: 3156: 3148: 3147: 3140: 3126: 3119: 3106: 3102: 3085: 3073: 3060: 3059: 3050: 3041: 3039: 3031: 3030: 3026: 3011: 2998: 2997: 2993: 2984: 2982: 2974: 2973: 2969: 2960: 2958: 2950: 2949: 2945: 2901: 2900: 2896: 2858: 2857: 2853: 2844: 2842: 2833: 2832: 2828: 2790: 2789: 2785: 2747: 2746: 2742: 2720: 2719: 2715: 2661: 2660: 2656: 2637: 2636: 2632: 2601:(10): 951–962. 2588: 2587: 2583: 2545: 2544: 2540: 2494: 2493: 2489: 2466: 2465: 2461: 2417: 2416: 2412: 2360: 2359: 2355: 2311: 2310: 2306: 2254: 2253: 2249: 2205: 2204: 2200: 2154: 2153: 2149: 2119: 2118: 2114: 2070: 2069: 2065: 2027: 2026: 2022: 1984: 1983: 1979: 1948:(10): a025130. 1935: 1934: 1930: 1908: 1907: 1903: 1864: 1863: 1859: 1815: 1814: 1810: 1800: 1799: 1795: 1754:(1): e0145544. 1741: 1740: 1736: 1700: 1699: 1695: 1653: 1652: 1648: 1641: 1620: 1619: 1615: 1569: 1568: 1564: 1503: 1502: 1498: 1444: 1443: 1439: 1393: 1392: 1388: 1334: 1333: 1324: 1280: 1279: 1275: 1231: 1230: 1226: 1180: 1179: 1172: 1134: 1133: 1129: 1091: 1090: 1081: 1045: 1044: 1037: 1019: 1018: 1011: 1000: 999: 995: 959: 958: 954: 941: 937: 891: 890: 883: 845: 844: 833: 829: 797: 771: 765: 741: 719:interacts with 713:atherosclerosis 709:type 2 diabetes 693: 691:Werner syndrome 687: 685:Werner Syndrome 655:lamin A protein 638: 632: 623: 618: 592:are created by 565: 541: 535: 507: 501: 476: 474:Group selection 470: 468:Group selection 465: 441: 435: 414: 406:Naked mole rats 387: 377: 301: 295: 272:Thomas Kirkwood 268: 262: 246:Michael R. Rose 213: 207: 198: 190:gene expression 134: 129: 123: 76:August Weismann 73: 68: 52:disposable soma 28: 23: 22: 15: 12: 11: 5: 5221: 5219: 5211: 5210: 5205: 5200: 5195: 5190: 5185: 5183:Life extension 5175: 5174: 5168: 5167: 5165: 5164: 5154: 5143: 5140: 5139: 5137: 5136: 5131: 5126: 5121: 5116: 5115: 5114: 5104: 5099: 5094: 5089: 5084: 5083: 5082: 5077: 5072: 5062: 5057: 5052: 5046: 5044: 5040: 5039: 5037: 5036: 5031: 5030: 5029: 5024: 5019: 5018: 5017: 5007: 5002: 4997: 4992: 4987: 4977: 4971: 4969: 4965: 4964: 4962: 4961: 4956: 4951: 4946: 4941: 4936: 4931: 4926: 4921: 4916: 4915: 4914: 4905:Charles Darwin 4902: 4901: 4900: 4888: 4883: 4877: 4875: 4869: 4868: 4866: 4865: 4860: 4855: 4850: 4845: 4843:Non-ecological 4840: 4835: 4830: 4825: 4820: 4815: 4810: 4804: 4802: 4796: 4795: 4793: 4792: 4783: 4774: 4760: 4758: 4752: 4751: 4749: 4748: 4743: 4742: 4741: 4736: 4731: 4726: 4721: 4711: 4706: 4701: 4696: 4691: 4686: 4681: 4676: 4671: 4666: 4661: 4660: 4659: 4649: 4644: 4639: 4634: 4633: 4632: 4627: 4616: 4614: 4607: 4606: 4604: 4603: 4602: 4601: 4596: 4594:nervous system 4591: 4586: 4581: 4573: 4572: 4571: 4566: 4561: 4556: 4551: 4546: 4536: 4531: 4526: 4520: 4518: 4511: 4510: 4508: 4507: 4502: 4497: 4492: 4487: 4486: 4485: 4475: 4474: 4473: 4468: 4467: 4466: 4461: 4451: 4446: 4441: 4436: 4431: 4430: 4429: 4424: 4414: 4404: 4399: 4398: 4397: 4387: 4382: 4377: 4372: 4371: 4370: 4360: 4355: 4354: 4353: 4343: 4337: 4335: 4328: 4327: 4325: 4324: 4319: 4314: 4309: 4304: 4299: 4293: 4291: 4287: 4286: 4284: 4283: 4278: 4273: 4268: 4267: 4266: 4261: 4256: 4246: 4241: 4236: 4231: 4226: 4225: 4224: 4219: 4209: 4204: 4199: 4198: 4197: 4187: 4182: 4177: 4172: 4166: 4164: 4156: 4155: 4153: 4152: 4151: 4150: 4140: 4135: 4134: 4133: 4128: 4118: 4117: 4116: 4106: 4101: 4096: 4094:Origin of life 4091: 4086: 4081: 4079:Microevolution 4076: 4074:Macroevolution 4071: 4066: 4061: 4060: 4059: 4049: 4044: 4039: 4034: 4029: 4024: 4019: 4014: 4012:Common descent 4009: 4008: 4007: 3997: 3992: 3990:Baldwin effect 3987: 3986: 3985: 3980: 3970: 3965: 3960: 3954: 3952: 3946: 3945: 3943: 3942: 3937: 3932: 3927: 3922: 3916: 3913: 3912: 3907: 3905: 3904: 3897: 3890: 3882: 3873: 3872: 3870: 3869: 3864: 3859: 3854: 3849: 3844: 3842:Life extension 3839: 3834: 3829: 3824: 3819: 3814: 3809: 3803: 3801: 3800:Related topics 3797: 3796: 3794: 3793: 3788: 3783: 3778: 3773: 3768: 3763: 3758: 3753: 3751:Hayflick limit 3748: 3743: 3738: 3733: 3728: 3722: 3720: 3714: 3713: 3707: 3705: 3704: 3697: 3690: 3682: 3676: 3675: 3669: 3663: 3657: 3645: 3639: 3629: 3622: 3621:External links 3619: 3617: 3616: 3606:(4): 799–804. 3595: 3550: 3541: 3532: 3503:10.1.1.10.7390 3488:(1): 513–517. 3477: 3442: 3416: 3394: 3392: 3389: 3387: 3386: 3351: 3327: 3303: 3279: 3255: 3243: 3206: 3185:(2): 224–233. 3165: 3138: 3117: 3100: 3071: 3048: 3024: 3009: 2991: 2967: 2943: 2914:(7): 719–735. 2894: 2867:(4): 331–359. 2851: 2826: 2799:(4): 693–709. 2793:Biogerontology 2783: 2756:(4): 764–768. 2740: 2713: 2654: 2630: 2581: 2538: 2487: 2459: 2410: 2373:(3): 691–696. 2353: 2304: 2247: 2198: 2147: 2128:(2): 412–416. 2112: 2083:(1): 274–286. 2063: 2036:(2): 199–207. 2020: 1999:10.1086/412317 1993:(3): 279–303. 1977: 1928: 1901: 1873:(9): 913–922. 1857: 1828:(6): 742–745. 1808: 1793: 1734: 1693: 1646: 1639: 1613: 1562: 1496: 1437: 1386: 1322: 1273: 1224: 1170: 1127: 1079: 1058:(2): 303–317. 1035: 1009: 993: 952: 935: 881: 854:(4): 398–411. 830: 828: 825: 824: 823: 818: 813: 808: 803: 796: 793: 792: 791: 783: 779: 767:Main article: 764: 763:Biogerontology 761: 744:Bloom syndrome 740: 737: 689:Main article: 686: 683: 640:Progeria is a 634:Main article: 631: 628: 622: 619: 617: 614: 564: 561: 537:Main article: 534: 531: 503:Main article: 500: 497: 472:Main article: 469: 466: 464: 461: 437:Main article: 434: 431: 413: 410: 398:ribonucleotide 392:are recurring 381:Hayflick limit 376: 373: 297:Main article: 294: 291: 264:Main article: 261: 258: 209:Main article: 206: 203: 197: 194: 133: 130: 125:Main article: 122: 119: 72: 69: 67: 64: 60:genetic traits 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 5220: 5209: 5206: 5204: 5201: 5199: 5196: 5194: 5191: 5189: 5186: 5184: 5181: 5180: 5178: 5163: 5159: 5155: 5153: 5145: 5144: 5141: 5135: 5132: 5130: 5127: 5125: 5122: 5120: 5117: 5113: 5110: 5109: 5108: 5107:Phylogenetics 5105: 5103: 5100: 5098: 5095: 5093: 5090: 5088: 5085: 5081: 5078: 5076: 5073: 5071: 5068: 5067: 5066: 5063: 5061: 5058: 5056: 5053: 5051: 5048: 5047: 5045: 5041: 5035: 5032: 5028: 5025: 5023: 5020: 5016: 5013: 5012: 5011: 5010:Structuralism 5008: 5006: 5003: 5001: 4998: 4996: 4993: 4991: 4988: 4986: 4985:Catastrophism 4983: 4982: 4981: 4978: 4976: 4973: 4972: 4970: 4966: 4960: 4957: 4955: 4952: 4950: 4947: 4945: 4944:Neo-Darwinism 4942: 4940: 4937: 4935: 4932: 4930: 4927: 4925: 4922: 4920: 4917: 4913: 4912: 4908: 4907: 4906: 4903: 4899: 4898: 4894: 4893: 4892: 4889: 4887: 4884: 4882: 4879: 4878: 4876: 4874: 4870: 4864: 4861: 4859: 4858:Reinforcement 4856: 4854: 4851: 4849: 4846: 4844: 4841: 4839: 4836: 4834: 4831: 4829: 4826: 4824: 4821: 4819: 4816: 4814: 4811: 4809: 4806: 4805: 4803: 4801: 4797: 4791: 4790:Catastrophism 4787: 4784: 4782: 4781:Macromutation 4778: 4777:Micromutation 4775: 4773: 4769: 4765: 4762: 4761: 4759: 4757: 4753: 4747: 4744: 4740: 4737: 4735: 4732: 4730: 4727: 4725: 4722: 4720: 4717: 4716: 4715: 4712: 4710: 4707: 4705: 4702: 4700: 4697: 4695: 4692: 4690: 4687: 4685: 4684:Immune system 4682: 4680: 4677: 4675: 4672: 4670: 4667: 4665: 4662: 4658: 4655: 4654: 4653: 4650: 4648: 4645: 4643: 4640: 4638: 4635: 4631: 4628: 4626: 4623: 4622: 4621: 4618: 4617: 4615: 4613: 4608: 4600: 4597: 4595: 4592: 4590: 4587: 4585: 4582: 4580: 4577: 4576: 4574: 4570: 4567: 4565: 4562: 4560: 4557: 4555: 4552: 4550: 4547: 4545: 4544:symbiogenesis 4542: 4541: 4540: 4537: 4535: 4532: 4530: 4527: 4525: 4522: 4521: 4519: 4517: 4512: 4506: 4503: 4501: 4498: 4496: 4493: 4491: 4488: 4484: 4481: 4480: 4479: 4476: 4472: 4469: 4465: 4462: 4460: 4457: 4456: 4455: 4452: 4450: 4447: 4445: 4442: 4440: 4437: 4435: 4432: 4428: 4425: 4423: 4420: 4419: 4418: 4415: 4413: 4410: 4409: 4408: 4405: 4403: 4400: 4396: 4393: 4392: 4391: 4388: 4386: 4383: 4381: 4378: 4376: 4373: 4369: 4366: 4365: 4364: 4361: 4359: 4356: 4352: 4349: 4348: 4347: 4344: 4342: 4339: 4338: 4336: 4334: 4329: 4323: 4320: 4318: 4315: 4313: 4310: 4308: 4305: 4303: 4300: 4298: 4295: 4294: 4292: 4288: 4282: 4279: 4277: 4274: 4272: 4269: 4265: 4262: 4260: 4257: 4255: 4252: 4251: 4250: 4247: 4245: 4242: 4240: 4237: 4235: 4232: 4230: 4227: 4223: 4220: 4218: 4215: 4214: 4213: 4212:Kin selection 4210: 4208: 4207:Genetic drift 4205: 4203: 4200: 4196: 4193: 4192: 4191: 4188: 4186: 4183: 4181: 4178: 4176: 4173: 4171: 4168: 4167: 4165: 4163: 4157: 4149: 4146: 4145: 4144: 4141: 4139: 4136: 4132: 4129: 4127: 4124: 4123: 4122: 4119: 4115: 4112: 4111: 4110: 4107: 4105: 4102: 4100: 4097: 4095: 4092: 4090: 4087: 4085: 4082: 4080: 4077: 4075: 4072: 4070: 4067: 4065: 4062: 4058: 4055: 4054: 4053: 4050: 4048: 4045: 4043: 4040: 4038: 4035: 4033: 4030: 4028: 4025: 4023: 4020: 4018: 4015: 4013: 4010: 4006: 4003: 4002: 4001: 3998: 3996: 3993: 3991: 3988: 3984: 3981: 3979: 3976: 3975: 3974: 3971: 3969: 3966: 3964: 3961: 3959: 3956: 3955: 3953: 3951: 3947: 3941: 3938: 3936: 3933: 3931: 3928: 3926: 3923: 3921: 3918: 3917: 3914: 3910: 3903: 3898: 3896: 3891: 3889: 3884: 3883: 3880: 3868: 3865: 3863: 3860: 3858: 3855: 3853: 3850: 3848: 3845: 3843: 3840: 3838: 3835: 3833: 3830: 3828: 3825: 3823: 3820: 3818: 3815: 3813: 3810: 3808: 3805: 3804: 3802: 3798: 3792: 3789: 3787: 3784: 3782: 3779: 3777: 3774: 3772: 3769: 3767: 3764: 3762: 3759: 3757: 3754: 3752: 3749: 3747: 3744: 3742: 3739: 3737: 3734: 3732: 3729: 3727: 3724: 3723: 3721: 3719: 3715: 3710: 3703: 3698: 3696: 3691: 3689: 3684: 3683: 3680: 3673: 3670: 3667: 3664: 3661: 3658: 3656: 3652: 3649: 3646: 3643: 3640: 3637: 3633: 3630: 3628: 3625: 3624: 3620: 3613: 3609: 3605: 3601: 3600:Human Biology 3596: 3592: 3588: 3583: 3578: 3573: 3568: 3564: 3560: 3556: 3551: 3547: 3542: 3538: 3533: 3529: 3525: 3521: 3517: 3513: 3509: 3504: 3499: 3495: 3491: 3487: 3483: 3478: 3474: 3470: 3465: 3460: 3456: 3452: 3448: 3443: 3439: 3427: 3419: 3413: 3409: 3405: 3401: 3396: 3395: 3390: 3382: 3378: 3374: 3370: 3366: 3362: 3355: 3352: 3341: 3337: 3331: 3328: 3317: 3313: 3307: 3304: 3293: 3289: 3283: 3280: 3269: 3265: 3259: 3256: 3250: 3248: 3244: 3239: 3235: 3230: 3225: 3221: 3217: 3210: 3207: 3202: 3198: 3193: 3188: 3184: 3180: 3176: 3169: 3166: 3155: 3151: 3145: 3143: 3139: 3135: 3131: 3124: 3122: 3118: 3115: 3111: 3104: 3101: 3096: 3090: 3082: 3078: 3074: 3068: 3064: 3057: 3055: 3053: 3049: 3038: 3034: 3028: 3025: 3020: 3016: 3012: 3006: 3002: 2995: 2992: 2981: 2977: 2971: 2968: 2957: 2953: 2947: 2944: 2939: 2935: 2930: 2925: 2921: 2917: 2913: 2909: 2905: 2898: 2895: 2890: 2886: 2882: 2878: 2874: 2870: 2866: 2862: 2855: 2852: 2840: 2836: 2830: 2827: 2822: 2818: 2814: 2810: 2806: 2802: 2798: 2794: 2787: 2784: 2779: 2775: 2771: 2767: 2763: 2759: 2755: 2751: 2744: 2741: 2736: 2732: 2728: 2724: 2717: 2714: 2709: 2705: 2700: 2695: 2690: 2685: 2681: 2677: 2673: 2669: 2665: 2658: 2655: 2649: 2645: 2641: 2634: 2631: 2626: 2622: 2617: 2612: 2608: 2604: 2600: 2596: 2592: 2585: 2582: 2577: 2573: 2569: 2565: 2561: 2557: 2553: 2549: 2542: 2539: 2534: 2530: 2526: 2522: 2518: 2514: 2510: 2506: 2502: 2498: 2491: 2488: 2482: 2478: 2474: 2470: 2463: 2460: 2455: 2451: 2447: 2443: 2438: 2433: 2429: 2425: 2421: 2414: 2411: 2406: 2402: 2398: 2394: 2390: 2386: 2381: 2376: 2372: 2368: 2364: 2357: 2354: 2349: 2345: 2340: 2335: 2331: 2327: 2323: 2319: 2315: 2308: 2305: 2300: 2296: 2291: 2286: 2282: 2278: 2274: 2270: 2266: 2262: 2258: 2251: 2248: 2243: 2239: 2234: 2229: 2225: 2221: 2217: 2213: 2209: 2202: 2199: 2194: 2190: 2185: 2180: 2175: 2170: 2166: 2162: 2158: 2151: 2148: 2143: 2139: 2135: 2131: 2127: 2123: 2116: 2113: 2108: 2104: 2099: 2094: 2090: 2086: 2082: 2078: 2074: 2067: 2064: 2059: 2055: 2051: 2047: 2043: 2039: 2035: 2031: 2024: 2021: 2016: 2012: 2008: 2004: 2000: 1996: 1992: 1988: 1981: 1978: 1973: 1969: 1964: 1959: 1955: 1951: 1947: 1943: 1939: 1932: 1929: 1924: 1920: 1917:(1–4): 3–14. 1916: 1912: 1905: 1902: 1896: 1892: 1888: 1884: 1880: 1876: 1872: 1868: 1861: 1858: 1853: 1849: 1845: 1841: 1836: 1831: 1827: 1823: 1819: 1812: 1809: 1804: 1797: 1794: 1789: 1785: 1780: 1775: 1770: 1765: 1761: 1757: 1753: 1749: 1745: 1738: 1735: 1730: 1726: 1721: 1716: 1712: 1708: 1704: 1697: 1694: 1689: 1685: 1681: 1677: 1673: 1669: 1665: 1661: 1657: 1650: 1647: 1642: 1636: 1632: 1628: 1624: 1617: 1614: 1609: 1605: 1600: 1595: 1590: 1585: 1581: 1577: 1573: 1566: 1563: 1558: 1554: 1549: 1544: 1540: 1536: 1532: 1528: 1524: 1520: 1516: 1512: 1508: 1500: 1497: 1492: 1488: 1483: 1478: 1473: 1468: 1464: 1460: 1456: 1452: 1448: 1441: 1438: 1433: 1429: 1425: 1421: 1417: 1413: 1409: 1405: 1401: 1397: 1390: 1387: 1382: 1378: 1373: 1368: 1363: 1358: 1354: 1350: 1347:(4): e34146. 1346: 1342: 1338: 1331: 1329: 1327: 1323: 1318: 1314: 1309: 1304: 1300: 1296: 1292: 1288: 1284: 1277: 1274: 1269: 1265: 1260: 1255: 1251: 1247: 1243: 1239: 1235: 1228: 1225: 1220: 1216: 1211: 1206: 1201: 1196: 1192: 1188: 1184: 1177: 1175: 1171: 1166: 1162: 1158: 1154: 1150: 1146: 1142: 1138: 1131: 1128: 1123: 1119: 1115: 1111: 1107: 1103: 1099: 1095: 1088: 1086: 1084: 1080: 1075: 1071: 1066: 1061: 1057: 1053: 1049: 1042: 1040: 1036: 1031: 1027: 1023: 1016: 1014: 1010: 1004: 997: 994: 989: 985: 980: 975: 971: 967: 963: 956: 953: 949: 945: 939: 936: 931: 927: 923: 919: 915: 911: 907: 903: 899: 895: 888: 886: 882: 877: 873: 869: 865: 861: 857: 853: 849: 842: 840: 838: 836: 832: 826: 822: 819: 817: 814: 812: 809: 807: 804: 802: 799: 798: 794: 789: 784: 780: 776: 775: 774: 770: 762: 760: 758: 754: 752: 748: 747:and disease. 745: 738: 736: 734: 730: 726: 722: 718: 714: 710: 706: 702: 698: 692: 684: 682: 680: 676: 672: 668: 664: 660: 656: 652: 647: 643: 637: 629: 627: 620: 615: 613: 611: 610:somatic cells 607: 603: 599: 595: 591: 587: 586:gametogenesis 583: 579: 575: 571: 562: 560: 556: 554: 545: 540: 532: 530: 526: 522: 520: 515: 511: 506: 498: 496: 494: 489: 485: 480: 475: 467: 462: 460: 456: 452: 450: 445: 440: 432: 430: 426: 423: 418: 411: 409: 407: 403: 399: 395: 391: 386: 382: 374: 368: 364: 362: 358: 354: 350: 349:transcription 345: 342: 337: 335: 331: 327: 323: 319: 315: 311: 307: 300: 292: 290: 288: 283: 281: 275: 273: 267: 259: 257: 255: 251: 247: 242: 238: 236: 232: 228: 227: 221: 218: 212: 204: 202: 196:Somatic cells 195: 193: 191: 185: 182: 178: 172: 168: 165: 159: 156: 151: 147: 143: 142:Peter Medawar 139: 131: 128: 120: 118: 115: 114: 109: 105: 101: 97: 92: 88: 86: 81: 77: 71:The beginning 70: 65: 63: 61: 57: 53: 49: 45: 41: 37: 33: 19: 5119:Polymorphism 5102:Astrobiology 5050:Biogeography 5005:Saltationism 4995:Orthogenesis 4980:Alternatives 4909: 4895: 4828:Cospeciation 4823:Cladogenesis 4772:Saltationism 4729:Mating types 4652:Color vision 4637:Avian flight 4619: 4559:mitochondria 4297:Canalisation 4175:Biodiversity 3920:Introduction 3822:CGK733 fraud 3740: 3603: 3599: 3562: 3558: 3545: 3536: 3485: 3481: 3457:(2): 72–74. 3454: 3450: 3399: 3367:(2): 64–71. 3364: 3360: 3354: 3343:. 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K. 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