Editing Population size

{{Short description|Total number of individuals in a defined group or area}}
{{More citations needed|date=December 2017}}
In [[population genetics]] and [[population ecology]], '''population size''' (usually denoted ''N'') is a [[countable quantity]] representing the number of individual [[organism]]s in a [[population]]. Population size is directly associated with amount of [[genetic drift]], and is the underlying cause of effects like [[population bottleneck]]s and the [[founder effect]].<ref name="Wright_1929">{{Cite journal|last=Wright|first=Sewall | name-list-style = vanc |date=November 1929 |title=The Evolution of Dominance |journal=The American Naturalist|language=en|volume=63|issue=689|pages=556–561|doi=10.1086/280290 |bibcode=1929ANat...63..556W |s2cid=85301374 }}</ref> Genetic drift is the major source of decrease of genetic diversity within populations which drives fixation and can potentially lead to [[speciation]] events.<ref name="Wright_1929" />

== Genetic drift ==
Of the five conditions required to maintain [[Hardy–Weinberg principle|Hardy-Weinberg Equilibrium]], infinite population size will always be violated; this means that some degree of genetic drift is always occurring.<ref name="Wright_1929"/> [[Small population size|Smaller population size]] leads to increased [[genetic drift]], it has been hypothesized that this gives these groups an evolutionary advantage for acquisition of genome complexity.<ref name=":2">{{cite journal | vauthors = Rozen DE, Habets MG, Handel A, de Visser JA | title = Heterogeneous adaptive trajectories of small populations on complex fitness landscapes | journal = PLOS ONE | volume = 3 | issue = 3 | pages = e1715 | date = March 2008 | pmid = 18320036 | pmc = 2248617 | doi = 10.1371/journal.pone.0001715 | bibcode = 2008PLoSO...3.1715R | doi-access = free }}</ref> An alternate hypothesis posits that while genetic drift plays a larger role in small populations developing complexity, selection is the mechanism by which large populations develop complexity.<ref name=":1">{{cite journal | vauthors = LaBar T, Adami C | title = Different Evolutionary Paths to Complexity for Small and Large Populations of Digital Organisms | journal = PLOS Computational Biology | volume = 12 | issue = 12 | pages = e1005066 | date = December 2016 | pmid = 27923053 | pmc = 5140054 | doi = 10.1371/journal.pcbi.1005066 | arxiv = 1604.06299 | bibcode = 2016PLSCB..12E5066L | doi-access = free }}</ref>

=== Population bottlenecks and founder effect ===
[[Population bottleneck]]s occur when population size reduces for a short period of time, decreasing the genetic diversity in the population.

The [[founder effect]] occurs when few individuals from a larger population establish a new population and also decreases the genetic diversity, and was originally outlined by [[Ernst Mayr]].<ref>{{cite journal | vauthors = Provine WB | title = Ernst Mayr: Genetics and speciation | journal = Genetics | volume = 167 | issue = 3 | pages = 1041–6 | date = July 2004 | doi = 10.1093/genetics/167.3.1041 | pmid = 15280221 | pmc = 1470966 }}</ref> The founder effect is a unique case of genetic drift, as the smaller founding population has decreased genetic diversity that will move alleles within the population more rapidly towards [[Fixation (population genetics)|fixation]].

=== Modeling genetic drift ===
Genetic drift is typically modeled in lab environments using bacterial populations or digital simulation. In digital organisms, a generated population undergoes evolution based on varying parameters, including differential fitness, variation, and heredity set for individual organisms.<ref name=":1" />

Rozen et al. use separate bacterial strains on two different mediums, one with simple nutrient components and one with nutrients noted to help populations of bacteria evolve more heterogeneity.<ref name=":2" />  A digital simulation based on the bacterial experiment design was also used, with assorted assignations of fitness and effective population sizes comparable to those of the bacteria used based on both small and large population designations<ref name=":2" /> Within both simple and complex environments, smaller populations demonstrated greater population variation than larger populations, which showed no significant fitness diversity.<ref name=":2" /> Smaller populations had increased fitness and adapted more rapidly in the complex environment, while large populations adapted faster than small populations in the simple environment.<ref name=":2" /> These data demonstrate that the consequences of increased variation within small populations is dependent on the environment: more challenging or complex environments allow variance present within small populations to confer greater advantage.<ref name=":2" /> Analysis demonstrates that smaller populations have more significant levels of fitness from heterogeneity within the group regardless of the complexity of the environment; adaptive responses are increased in more complex environments.<ref name=":2" /> Adaptations in asexual populations are also not limited by mutations, as genetic variation within these populations can drive adaptation.<ref>{{cite journal | vauthors = Lang GI, Botstein D, Desai MM | title = Genetic variation and the fate of beneficial mutations in asexual populations | journal = Genetics | volume = 188 | issue = 3 | pages = 647–61 | date = July 2011 | pmid = 21546542 | pmc = 3176544 | doi = 10.1534/genetics.111.128942 }}</ref> Although small populations tend to face more challenges because of limited access to widespread beneficial mutation adaptation within these populations is less predictable and allows populations to be more plastic in their environmental responses.<ref name=":2" /> Fitness increase over time in small asexual populations is known to be strongly positively correlated with population size and mutation rate, and fixation probability of a beneficial mutation is inversely related to population size and mutation rate.<ref>{{cite journal | vauthors = Gerrish PJ, Lenski RE | title = The fate of competing beneficial mutations in an asexual population | journal = Genetica | volume = 102-103 | issue = 1–6 | pages = 127–44 | date = 1998 | pmid = 9720276 | doi = 10.1023/a:1017067816551 | s2cid = 15148583 }}</ref>

LaBar and Adami use digital haploid organisms to assess differing strategies for accumulating genomic complexity. This study demonstrated that both drift and selection are effective in small and large populations, respectively, but that this success is dependent on several factors.<ref name=":1" /> Data from the observation of insertion mutations in this digital system demonstrate that small populations evolve larger genome sizes from fixation of deleterious mutations and large populations evolve larger genome sizes from fixation of beneficial mutations.<ref name=":1" />  Small populations were noted to have an advantage in attaining full genomic complexity due to drift-driven phenotypic complexity.<ref name=":1" /> When deletion mutations were simulated, only the largest populations had any significant fitness advantage.<ref name=":1" /> These simulations demonstrate that smaller populations fix deleterious mutations by increased genetic drift.<ref name=":1" /> This advantage is likely limited by high rates of extinction.<ref name=":1" /> Larger populations evolve complexity through mutations that increase expression of particular genes; removal of deleterious alleles does not limit developing more complex genomes in the larger groups and a large number of insertion mutations that resulted in beneficial or non-functional elements within the genome were not required.<ref name=":1" /> When deletion mutations occur more frequently, the largest populations have an advantage that suggests larger populations generally have an evolutionary advantage for development of new traits.<ref name=":1" />

== Critical Mutation Rate ==
Critical mutation rate, or [[Error threshold (evolution)|error threshold]], limits the number of mutations that can exist within a self-replicating molecule before genetic information is destroyed in later generations.<ref>{{cite journal | vauthors = Eigen M | title = Selforganization of matter and the evolution of biological macromolecules | journal = Die Naturwissenschaften | volume = 58 | issue = 10 | pages = 465–523 | date = October 1971 | pmid = 4942363 | doi = 10.1007/bf00623322 | bibcode = 1971NW.....58..465E | s2cid = 38296619 }}</ref>

Contrary to the findings of previous studies,<ref>{{cite journal | vauthors = Gillespie JH | title = Is the population size of a species relevant to its evolution? | journal = Evolution; International Journal of Organic Evolution | volume = 55 | issue = 11 | pages = 2161–9 | date = November 2001 | pmid = 11794777 | jstor = 2680348 | doi=10.1111/j.0014-3820.2001.tb00732.x| doi-access = free }}</ref> critical mutation rate has been noted to be dependent on population size in both haploid and diploid populations.<ref name=":0">{{cite journal | vauthors = Aston E, Channon A, Day C, Knight CG | title = Critical mutation rate has an exponential dependence on population size in haploid and diploid populations | journal = PLOS ONE | volume = 8 | issue = 12 | pages = e83438 | date = 2013-12-27 | pmid = 24386200 | pmc = 3873944 | doi = 10.1371/journal.pone.0083438 | bibcode = 2013PLoSO...883438A | doi-access = free }}</ref> When populations have fewer than 100 individuals, critical mutation rate can be exceeded, but will lead to loss of genetic material which results in further population decline and likelihood of extinction.<ref name=":0" /> This ‘speed limit’ is common within small, adapted asexual populations and is independent of mutation rate.<ref>{{cite journal | vauthors = Arjan JA, Visser M, Zeyl CW, Gerrish PJ, Blanchard JL, Lenski RE | title = Diminishing returns from mutation supply rate in asexual populations | journal = Science | volume = 283 | issue = 5400 | pages = 404–6 | date = January 1999 | doi = 10.1126/science.283.5400.404 | pmid = 9888858 | jstor = 2896813 | bibcode = 1999Sci...283..404A }}</ref>

== Effective population size (''N<sub>e</sub>'') ==
The [[effective population size]] (''N<sub>e</sub>'') is defined as "the number of breeding individuals in an [[idealized population]] that would show the same amount of dispersion of [[allele frequencies]] under random [[genetic drift]] or the same amount of [[inbreeding]] as the population under consideration."  ''N<sub>e</sub>'' is usually less than ''N'' (the absolute population size) and this has important applications in [[conservation genetics]].<ref>{{cite journal | vauthors = Husemann M, Zachos FE, Paxton RJ, Habel JC | title = Effective population size in ecology and evolution | language = En | journal = Heredity | volume = 117 | issue = 4 | pages = 191–2 | date = October 2016 | pmid = 27553454 | pmc = 5026761 | doi = 10.1038/hdy.2016.75 }}</ref>

[[Overpopulation (biology)|Overpopulation]] may indicate any case in which the population of any species of animal may exceed the [[carrying capacity]] of its [[ecological niche]].<ref>{{cite journal | author = Population Reference Bureau PRB | title = What is overpopulation? | journal = Population Education Interchange | volume = 17 | issue = 4 | pages = 1–2 | date = December 1988 | pmid = 12281798 }}</ref>

== See also ==
* [[Carrying capacity]]
* [[Holocene extinction event]]
* [[Lists of organisms by population]]
* [[Overpopulation (biology)|Overpopulation]]
* [[Population growth rate]]

== References ==
{{Reflist}}

{{modelling ecosystems|expanded=other}}

{{Authority control}}

{{DEFAULTSORT:Population Size}}
[[Category:Ecological metrics]]
[[Category:Population genetics]]
[[Category:Countable quantities]]