Selection shadow

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(a) The survival rate of a population decreases as individuals age, but the reproduction rate remains constant. (b) The reproduction probability peaks early in life at sexual maturity and then steadily decreases as an individual ages, with the remaining share of the population decreasing with age as they enter the selection shadow. Selection shadow.svg
(a) The survival rate of a population decreases as individuals age, but the reproduction rate remains constant. (b) The reproduction probability peaks early in life at sexual maturity and then steadily decreases as an individual ages, with the remaining share of the population decreasing with age as they enter the selection shadow.

The selection shadow is a concept involved with the evolutionary theories of aging that states that selection pressures on an individual decrease as an individual ages and passes sexual maturity, resulting in a "shadow" of time where selective fitness is not considered. Over generations, this results in maladaptive mutations that accumulate later in life due to aging being non-adaptive toward reproductive fitness. [1] The concept was first worked out by J. B. S. Haldane and Peter Medawar in the 1940s, with Medawar creating the first graphical model. [1]

Contents

Model

The model developed by Medawar states that due to the dangerous conditions and pressures from the environment, including predators and diseases, most individuals in the wild die not long after sexual maturity. Therefore, there is a low probability for individuals to survive to an advanced age and suffer the effects related to aging. In conjunction with this, the effects of natural selection decrease as age increases, so that later individual performance is ignored by selection forces. [1] This results in beneficial mutations not being selected for if they only have a positive result later in life, along with later in life deleterious mutations not being selected against. Due to the fitness of an individual not being affected once it is past its reproductive prime, later mutations and effects are considered to be in the "shadow" of selection. [2]

This concept would later be adapted into Medawar's 1952 mutation accumulation hypothesis, which was itself expanded upon by George C. Williams in his 1957 antagonistic pleiotropy hypothesis. [1]

A classical requirement and constraint of the model is that the number of individuals within a population that live to reach senescence must be small in number. If this is not true for a population, then the effects of old age will not be under a selection shadow and instead affect adaptation and evolution of the population as a whole. At the same time, however, this requirement has been challenged by increasing evidence of senescence being more common in wild populations than previously expected, especially among birds and mammals, while the effects of the selection shadow remain present. [3]

Medawar's Test Tube model

Peter Brian Medawar, the scientist/biologist behind the test tube theory. This theory was created to demonstrate how older test tubes are continuously replaced when they are broken, thus decreasing older test tube population numbers as time progresses. The test tubes were symbols for individual subjects in a species. Peter Brian Medawar.jpg
Peter Brian Medawar, the scientist/biologist behind the test tube theory. This theory was created to demonstrate how older test tubes are continuously replaced when they are broken, thus decreasing older test tube population numbers as time progresses. The test tubes were symbols for individual subjects in a species.

Medawar developed a theoretical model to demonstrate his thought process which explained that most animals will die before aging will be the ultimate cause for death in that animal. This would be from environmental factors such as large storms, drought, and fires, and predation. Medawar wanted to demonstrate this possibility by using test tubes to get his point across. The test tubes would be used to represent a population of species. [4] If one of these test tubes were to theoretically break, this would represent an individual animal dying. Randomly, test tubes would then be broken in the population to keep his model realistic. The broken test tubes would be replaced with a new one, which represents a new animal being born into the population. [4] Over time, the model showed that test tubes over a certain age would decline in the population as new test tubes were put in. The overall results in Medawar’s thought model demonstrated an exponential decline in the survivor curve which resulted in the population having a half life. [4] The amount of older animals, or test tubes in the population would then be harder to maintain and ultimately die. Medawar created this model to ultimately explain what would realistically happen in actual life.

Criticism

Some scientists, however, have criticized the idea of aging being non-adaptive, instead adopting the theory of "death by design". This theory follows the work of August Weismann, which states that aging specifically evolved as an adaptation, and disagrees with Medawar's model as a perceived oversimplification of the impact older organisms have on evolution. It is also claimed that older organisms have a higher reproductive capacity due to being better fit in order to reach their age, rather than their capacity being equal as in Medawar's calculations. [5]

Related Research Articles

Natural selection Mechanism of evolution by differential survival and reproduction of individuals

Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Charles Darwin popularised the term "natural selection", contrasting it with artificial selection, which in his view is intentional, whereas natural selection is not.

Senescence Deterioration of function with age

Senescence or biologicalaging is the gradual deterioration of functional characteristics in living organisms. The word senescence can refer to either cellular senescence or to senescence of the whole organism. Organismal senescence involves an increase in death rates and/or a decrease in fecundity with increasing age, at least in the latter part of an organism's life cycle.

Survival of the fittest Phrase to describe the mechanism of natural selection

"Survival of the fittest" is a phrase that originated from Darwinian evolutionary theory as a way of describing the mechanism of natural selection. The biological concept of fitness is defined as reproductive success. In Darwinian terms the phrase is best understood as "Survival of the form that will leave the most copies of itself in successive generations."

Adaptation Trait with a current functional role in the life history of an organism maintained and evolved by natural selection

In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the population during that process. Thirdly, it is a phenotypic trait or adaptive trait, with a functional role in each individual organism, that is maintained and has evolved through natural selection.

Reproductive success The passing of genes on to the next generation in a way that they too can pass on those genes

Reproductive success is an individual's production of offspring per breeding event or lifetime. This is not limited by the number of offspring produced by one individual, but also the reproductive success of these offspring themselves. Reproductive success is different from fitness in that individual success is not necessarily a determinant for adaptive strength of a genotype since the effects of chance and the environment have no influence on those specific genes. Reproductive success turns into a part of fitness when the offspring are actually recruited into the breeding population. If offspring quantity is not correlated with quality this holds up, but if not then reproductive success must be adjusted by traits that predict juvenile survival in order to be measured effectively. Quality and quantity is about finding the right balance between reproduction and maintenance and the disposable soma theory of aging tells us that a longer lifespan will come at the cost of reproduction and thus longevity is not always correlated with high fecundity. Parental investment is a key factor in reproductive success since taking better care to offspring is what often will give them a fitness advantage later in life. This includes mate choice and sexual selection as an important factor in reproductive success, which is another reason why reproductive success is different from fitness as individual choices and outcomes are more important than genetic differences. As reproductive success is measured over generations, Longitudinal studies are the preferred study type as they follow a population or an individual over a longer period of time in order to monitor the progression of the individual(s). These long term studies are preferable since they negate the effects of the variation in a single year or breeding season.

Pleiotropy Influence of a single gene on multiple phenotypic traits

Pleiotropy occurs when one gene influences two or more seemingly unrelated phenotypic traits. Such a gene that exhibits multiple phenotypic expression is called a pleiotropic gene. Mutation in a pleiotropic gene may have an effect on several traits simultaneously, due to the gene coding for a product used by a myriad of cells or different targets that have the same signaling function.

In population genetics and population ecology, population size is the number of individual organisms in a population. Population size is directly associated with amount of genetic drift, and is the underlying cause of effects like population bottlenecks and the founder effect. Genetic drift is the major source of decrease of genetic diversity within populations which drives fixation and can potentially lead to speciation events.

Grandmother hypothesis

The grandmother hypothesis is a hypothesis to explain the existence of menopause in human life history by identifying the adaptive value of extended kin networking. It builds on the previously postulated "mother hypothesis" which states that as mothers age, the costs of reproducing become greater, and energy devoted to those activities would be better spent helping her offspring in their reproductive efforts. It suggests that by redirecting their energy onto those of their offspring, grandmothers can better ensure the survival of their genes through younger generations. By providing sustenance and support to their kin, grandmothers not only ensure that their genetic interests are met, but they also enhance their social networks which could translate into better immediate resource acquisition. This effect could extend past kin into larger community networks and benefit wider group fitness.

The age of onset is the age at which an individual acquires, develops, or first experiences a condition or symptoms of a disease or disorder. For instance, the general age of onset for the spinal disease scoliosis is "10-15 years old," meaning that most people develop scoliosis when they are of an age between ten and fifteen years.

Inbreeding depression is the reduced biological fitness in a given population as a result of inbreeding, or breeding of related individuals. Population biological fitness refers to an organism's ability to survive and perpetuate its genetic material. Inbreeding depression is often the result of a population bottleneck. In general, the higher the genetic variation or gene pool within a breeding population, the less likely it is to suffer from inbreeding depression.

Life history theory is an analytical framework designed to study the diversity of life history strategies used by different organisms throughout the world, as well as the causes and results of the variation in their life cycles. It is a theory of biological evolution that seeks to explain aspects of organisms' anatomy and behavior by reference to the way that their life histories—including their reproductive development and behaviors, post-reproductive behaviors, and lifespan —have been shaped by natural selection. A life history strategy is the "age- and stage-specific patterns" and timing of events that make up an organism's life, such as birth, weaning, maturation, death, etc. These events, notably juvenile development, age of sexual maturity, first reproduction, number of offspring and level of parental investment, senescence and death, depend on the physical and ecological environment of the organism.

Enquiry into the evolution of ageing aims to explain why a detrimental process such as aging would evolve, and why there is so much variability in the lifespans of living organisms. The classical theories of evolution suggest that environmental factors, such as predation, accidents, disease, 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. Natural selection will instead strongly favor genes which ensure early maturation and rapid reproduction, and the selection for genetic traits which promote molecular and cellular self-maintenance will decline with age for most organisms.

Red Queen hypothesis Concept in evolutionary biology

The Red Queen hypothesis, also referred to as Red Queen's, the Red Queen effect, the Red Queen model, Red Queen's race, and Red Queen dynamics, is a hypothesis in evolutionary biology which proposes that species must constantly adapt, evolve, and proliferate in order to survive while pitted against ever-evolving opposing species. The hypothesis was intended to explain the constant (age-independent) extinction probability as observed in the paleontological record caused by co-evolution between competing species; however, it has also been suggested that the Red Queen hypothesis explains the advantage of sexual reproduction at the level of individuals, and the positive correlation between speciation and extinction rates in most higher taxa.

Antagonistic pleiotropy hypothesis proposed evolutionary explanation for senescence

The antagonistic pleiotropy hypothesis was first proposed by George C. Williams in 1957 as an evolutionary explanation for senescence. Pleiotropy is the phenomenon where one gene controls for more than one phenotypic trait in an organism. Antagonistic pleiotropy is when one gene controls for more than one trait, where at least one of these traits is beneficial to the organism's fitness early on in life and at least one is detrimental to the organism's fitness later on due to a decline in the force of natural selection. The theme of G.C. William's idea about antagonistic pleiotropy was that if a gene caused both increased reproduction in early life and aging in later life, then senescence would be adaptive in evolution. For example, one study suggests that since follicular depletion in human females causes both more regular cycles in early life and loss of fertility later in life through menopause, it can be selected for by having its early benefits outweigh its late costs.

Host–parasite coevolution Mutually adaptive genetic change of a host and a parasite

Host–parasite coevolution is a special case of coevolution, where a host and a parasite continually adapt to each other. This can create an evolutionary arms race between them. A more benign possibility is of an evolutionary trade-off between transmission and virulence in the parasite, as if it kills its host too quickly, the parasite will not be able to reproduce either. Another theory, the Red Queen hypothesis, proposes that since both host and parasite have to keep on evolving to keep up with each other, and since sexual reproduction continually creates new combinations of genes, parasitism favours sexual reproduction in the host.

Outline of evolution Hierarchical outline list of articles related to evolution

The following outline is provided as an overview of and topical guide to evolution:

The disposable soma theory of aging states that organisms age due to an evolutionary trade-off between growth, reproduction, and DNA repair maintenance. Formulated by Thomas Kirkwood, the disposable soma theory explains that an organism only has a limited amount of resources that it can allocate to its various cellular processes. Therefore, a greater investment in growth and reproduction would result in reduced investment in DNA repair maintenance, leading to increased cellular damage, shortened telomeres, accumulation of mutations, compromised stem cells, and ultimately, senescence. Although many models, both animal and human, have appeared to support this theory, parts of it are still controversial. Specifically, while the evolutionary trade-off between growth and aging has been well established, the relationship between reproduction and aging is still without scientific consensus, and the cellular mechanisms largely undiscovered.

This glossary of evolutionary biology is a list of definitions of terms and concepts used in the study of evolutionary biology, population biology, speciation, and phylogenetics, as well as sub-disciplines and related fields. For additional terms from related glossaries, see Glossary of genetics, Glossary of ecology, and Glossary of biology.

Extrinsic mortality is the sum of the effects of external factors, such as sunlight and pollutants that contribute to senescence and eventually death. This is opposed to intrinsic mortality, which is the sum of the effects of internal factors, such as mutation due to DNA replication errors. Extrinsic mortality plays a significant role in evolutionary theories of aging, as well as the discussion of health barriers across socioeconomic borders.

Mutation accumulation theory

The mutation accumulation theory of ageing was first proposed by Peter Medawar in 1952 as an evolutionary explanation for biological ageing and the associated decline in fitness that accompanies it. Medawar used the term 'senescence' to refer to this process. The theory explains that, in the case where harmful mutations are only expressed later in life, when reproduction has ceased and future survival is increasingly unlikely, then these mutations are likely to be unknowingly passed on to future generations. In this situation the force of natural selection will be weak, and so insufficient to consistently eliminate these mutations. Medawar posited that over time these mutations would accumulate due to genetic drift and lead to the evolution of what is now referred to as ageing.

References

  1. 1 2 3 4 Fabian, Daniel; Flatt, Thomas (2011). "The Evolution of Aging". Scitable . Nature Publishing Group . Retrieved May 20, 2014.
  2. Flatt, Thomas; Schmidt, Paul S. (July 18, 2009). "Integrating evolutionary and molecular genetics of aging" (PDF). Biochimica et Biophysica Acta (BBA) - General Subjects. Elsevier. 1790 (10): 951–962. doi:10.1016/j.bbagen.2009.07.010. PMC   2972575 . PMID   19619612 . Retrieved May 20, 2014.
  3. Turbill, Christopher; Ruf, Thomas (August 6, 2010). "Senescence Is More Important in the Natural Lives of Long- Than Short-Lived Mammals". PLOS One . Public Library of Science. 5 (8): e12019. Bibcode:2010PLoSO...512019T. doi: 10.1371/journal.pone.0012019 . PMC   2917356 . PMID   20700508.
  4. 1 2 3 "Medawar's Hypothesis - Aging has a negligible effect on fitness". www.programmed-aging.org. Retrieved 2020-04-13.
  5. Lachowicz, Miroslaw; Miękisz, Jacek (April 20, 2009). From Genetics to Mathematics. Singapore: World Scientific. p. 39. ISBN   978-9812837257 . Retrieved May 25, 2014.