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]
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 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.[ citation needed ]
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]
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 is intentional, whereas natural selection is not.
Senescence or biological aging is the gradual deterioration of functional characteristics in living organisms. Whole organism senescence involves an increase in death rates or a decrease in fecundity with increasing age, at least in the later part of an organism's life cycle. However, the resulting effects of senescence can be delayed. The 1934 discovery that calorie restriction can extend lifespans by 50% in rats, the existence of species having negligible senescence, and the existence of potentially immortal organisms such as members of the genus Hydra have motivated research into delaying senescence and thus age-related diseases. Rare human mutations can cause accelerated aging diseases.
Population genetics is a subfield of genetics that deals with genetic differences within and among populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and population structure.
"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 in successive generations will leave most copies of itself."
In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection 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 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.
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.
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 age between ten and fifteen years.
Inbreeding depression is the reduced biological fitness that has the potential to result from inbreeding. The loss of genetic diversity that is seen due to inbreeding, results from small population size. 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, though inbreeding and outbreeding depression can simultaneously occur.
Philopatry is the tendency of an organism to stay in or habitually return to a particular area. The causes of philopatry are numerous, but natal philopatry, where animals return to their birthplace to breed, may be the most common. The term derives from the Greek roots philo, "liking, loving" and patra, "fatherland", although in recent years the term has been applied to more than just the animal's birthplace. Recent usage refers to animals returning to the same area to breed despite not being born there, and migratory species that demonstrate site fidelity: reusing stopovers, staging points, and wintering grounds.
Life history theory (LHT) 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, or aging, aims to explain why a detrimental process such as ageing would evolve, and why there is so much variability in the lifespans of organisms. The classical theories of evolution 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. 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.
The antagonistic pleiotropy hypothesis (APT) is a theory in evolutionary biology that suggests certain genes may confer beneficial effects early in an organism's life, enhancing reproductive success, while also causing detrimental effects later in life, contributing to the aging process.
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.
In biogerontology, 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 British biologist 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 genetics and evolutionary biology is a list of definitions of terms and concepts used in the study of genetics and evolutionary biology, as well as sub-disciplines and related fields, with an emphasis on classical genetics, quantitative genetics, population biology, phylogenetics, speciation, and systematics. It has been designed as a companion to Glossary of cellular and molecular biology, which contains many overlapping and related terms; other related glossaries include Glossary of biology and Glossary of ecology.
Extrinsic mortality is the sum of the effects of external factors, such as predation, starvation and other environmental factors not under control of the individual that cause death. This is opposed to intrinsic mortality, which is the sum of the effects of internal factors contributing to normal, chronologic aging, such as, for example, mutations due to DNA replication errors, and which determined species maximum lifespan. Extrinsic mortality plays a significant role in evolutionary theories of aging, as well as the discussion of health barriers across socioeconomic borders.
The mutation accumulation theory of aging was first proposed by Peter Medawar in 1952 as an evolutionary explanation for biological aging 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 aging.
Few animals have a menopause: humans are joined by just four other species in which females live substantially longer than their ability to reproduce. The others are all cetaceans: beluga whales, narwhals, orcas and short-finned pilot whales. There are various theories on the origin and process of the evolution of menopause. These attempt to suggest evolutionary benefits to the human species stemming from the cessation of women's reproductive capability before the end of their natural lifespan. Explanations can be categorized as adaptive and non-adaptive: