Drift Load
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Drift load, a type of
genetic load Genetic load is the difference between the fitness of an average genotype in a population and the fitness of some reference genotype, which may be either the best present in a population, or may be the theoretically optimal genotype. The average ...
, is the decline in fitness in a
population Population is a set of humans or other organisms in a given region or area. Governments conduct a census to quantify the resident population size within a given jurisdiction. The term is also applied to non-human animals, microorganisms, and pl ...
due to the fixation of
deleterious mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mitosis ...
s by
genetic drift Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene va ...
. Drift load generally decreases as a function of
effective population size The effective population size (''N'e'') is the size of an idealised population that would experience the same rate of genetic drift as the real population. Idealised populations are those following simple one- locus models that comply with ass ...
, and is a mechanism of
inbreeding depression Inbreeding depression is the reduced biological fitness caused by loss of genetic diversity as a consequence of inbreeding, the breeding of individuals closely related genetically. This loss of genetic diversity results from small population siz ...
. If all individuals in a population become homozygous for one
allele An allele is a variant of the sequence of nucleotides at a particular location, or Locus (genetics), locus, on a DNA molecule. Alleles can differ at a single position through Single-nucleotide polymorphism, single nucleotide polymorphisms (SNP), ...
, that allele is
fixed Fixed may refer to: * ''Fixed'' (EP), EP by Nine Inch Nails * ''Fixed'' (film), an upcoming animated film directed by Genndy Tartakovsky * Fixed (typeface), a collection of monospace bitmap fonts that is distributed with the X Window System * Fi ...
, and any other alleles at that locus are lost from that population by the random
evolutionary process Evolution is the change in the heritable characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, resulting in certai ...
of genetic drift. As fixation of deleterious alleles occurs at more loci and variability is lost, drift load increases. With increasing drift load, populations become less fit with each generation, a
mutational meltdown In evolutionary genetics, mutational meltdown is a sub class of extinction vortex in which the environment and genetic predisposition mutually reinforce each other. Mutational meltdown (not to be confused with the concept of an error catastroph ...
, which can lead to an
extinction vortex Extinction vortices are a class of models through which conservation biologists, geneticists and ecologists can understand the dynamics of and categorize extinctions in the context of their causes. This model shows the events that ultimately lea ...
. Strategies to address drift load in
endangered species An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching, inv ...
is a goal of
conservation genetics Conservation genetics is an interdisciplinary subfield of population genetics that aims to understand the dynamics of genes in a population for the purpose of natural resource management, conservation of genetic diversity, and the prevention of ...
.


Calculating drift load

For realistic values of N_e, the effective population size, and \lambda_D, the mean effect on fitness of all fixed deleterious mutations, the loss of fitness in each generation due to drift load, \Delta , can be calculated: \Delta \cong \frac Where the total number of new deleterious mutations per diploid zygote is U_D. This demonstrates that more fixed deleterious alleles in a population lead to greater reductions in fitness, unless counteracted by large effective populations or weakly deleterious effects overall.


Decreasing drift load

Factors that decrease drift load, or counteract its negative effects on population fitness, include
polyploidy Polyploidy is a condition in which the cells of an organism have more than two paired sets of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one fro ...
,
gene flow In population genetics, gene flow (also known as migration and allele flow) is the transfer of genetic variation, genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent ...
and
purifying selection In natural selection, negative selection or purifying selection is the selective removal of alleles that are deleterious. This can result in stabilising selection through the purging of deleterious genetic polymorphisms that arise through random ...
. Drift load increases more rapidly in small populations, so factors that tend to maintain large populations such as high reproductive rates and low levels of biotic and
abiotic stress Abiotic stress is the negative impact of non-living factors on the living organisms in a specific environment. The non-living variable must influence the environment beyond its normal range of variation to adversely affect the population performan ...
will reduce the incidence of drift load. As a target of conservationists seeking to ensure endangered animals' long-term survival, drift load has been directly addressed in genetic restoration projects around the world. Some researchers caution that long-term benefits of genetic restoration and assisted gene flow may be doubtful, and these programs have risks that are not yet well-understood. Nonetheless, several projects have reported success in reducing drift load in wild populations.


Florida Panther

By 1995, the
Florida panther The Florida panther is a North American cougar (''P. c. couguar'') population in South Florida. It lives in pinelands, tropical hardwood hammocks and mixed freshwater swamp forests. Its range includes the Big Cypress National Preserve, Everglade ...
(''Felis concolor coryi''), with a local range between two large cities and divided by highways, was reduced to a population of 20-25 individuals in the wild. As a result of inbreeding depression and drift load, Florida panthers exhibited low genetic variability and overall fitness, as shown in congenital heart defects, high parasite and pathogen load and impaired male fertility and sperm viability. Though protected from hunting as an endangered species, models predicted the risk of extinction within 20 years as 95%. A genetic restoration project was developed to achieve gene flow and an increase in genetic variability. Eight female panthers from Texas (''
Felis concolor stanleyana ''Felis'' is a genus of small and medium-sized cat species native to most of Africa and south of 60° latitude in Europe and Asia to Indochina. The genus includes the domestic cat. The smallest of the seven ''Felis'' species is the black-footed ...
'') were transported to join the remaining 22 Florida panthers and 4 panthers originally from the Everglades in their habitat. The results after 15 years of monitoring were a tripled population size, doubled genetic heterozygosity, a lowered incidence of congenital disorders and inbreeding correlates, and improved survival and fitness.


Common European adder

A population of adders (''Vipera berus berus'') in Smygehuk, Sweden had become reproductively isolated by habitat encroachment to a small strip of coastal grassland. Their numbers had declined and so had their fitness; they exhibited severe inbreeding depression and drift load, demonstrated by a high proportion of deformed or stillborn offspring and very low genetic variability. Researchers released 20 male adders from another Swedish population into the Smygehuk habitat, and after four breeding seasons captured the visiting adders and transported them back to their native range. The population grew dramatically, from about 18 to about 64 individuals during the study. Incidence of congenital defects and stillbirth fell sharply, and genetic variability greatly increased, as measured by polymorphism in
MHC class I MHC class I molecules are one of two primary classes of major histocompatibility complex (MHC) molecules (the other being MHC class II) and are found on the cell surface of all nucleated cells in the bodies of vertebrates. They also occur on ...
genes.


Scandinavian arctic fox

The Scandinavian
arctic fox The Arctic fox (''Vulpes lagopus''), also known as the white fox, polar fox, or snow fox, is a small species of fox native to the Arctic regions of the Northern Hemisphere and common throughout the Tundra#Arctic tundra, Arctic tundra biome. I ...
(''Vulpes lagopus'') was hunted almost to extinction for the fur trade, thereafter going through a genetic bottleneck that, in the southernmost population in Norway led to an average
inbreeding coefficient The coefficient of relationship is a measure of the degree of consanguinity (or biological relationship) between two individuals. The term coefficient of relationship was defined by Sewall Wright in 1922, and was derived from his definition of ...
higher (''f'' = 0.14) than that of half-sibling crosses (''f'' = 0.125). The release of three out-bred males from a captive breeding program in 2010 into this population resulted, in five years, in a doubling of the population and a large decrease in its average inbreeding coefficient to ''f'' = 0.08, with kits' first-year survival rate almost doubling, and a 41% increase in allelic richness. Five years after their introduction, 89% of litters descended from the three immigrants.


References

{{reflist Wikipedia Student Program Population genetics Extinction Evolutionary biology Endangered species Mutation Natural selection