Эффект Аллее: зависимость приспособленности от плотности популяции.

Population phenomenon in biology
The Allee effect is a phenomenon in biology characterized by a correlation between population size or density and the mean individual fitness (often measured as per capita population growth rate) of a population or species.

Although the concept of Allee effect had no title at the time, it was first described in the 1930s by its namesake, Warder Clyde Allee. Through experimental studies, Allee was able to demonstrate that goldfish have a greater survival rate when there are more individuals within the tank. This led him to conclude that aggregation can improve the survival rate of individuals, and that cooperation may be crucial in the overall evolution of social structure. The term "Allee principle" was introduced in the 1950s, a time when the field of ecology was heavily focused on the role of competition among and within species. The classical view of population dynamics stated that due to competition for resources, a population will experience a reduced overall growth rate at higher density and increased growth rate at lower density. In other words, individuals in a population would be better off when there are fewer individuals around due to a limited amount of resources (see logistic growth). However, the concept of the Allee effect introduced the idea that the reverse holds true when the population density is low. Individuals within a species often require the assistance of another individual for more than simple reproductive reasons in order to persist. The most obvious example of this is observed in animals that hunt for prey or defend against predators as a group.

The generally accepted definition of Allee effect is positive density dependence, or the positive correlation between population density and individual fitness. It is sometimes referred to as "undercrowding" and it is analogous (or even considered synonymous by some) to "depensation" in the field of fishery sciences. Listed below are a few significant subcategories of the Allee effect used in the ecology literature.

The component Allee effect is the positive relationship between any measurable component of individual fitness and population density. The demographic Allee effect is the positive relationship between the overall individual fitness and population density. The distinction between the two terms lies on the scale of the Allee effect: the presence of a demographic Allee effect suggests the presence of at least one component Allee effect, while the presence of a component Allee effect does not necessarily result in a demographic Allee effect. For example, cooperative hunting and the ability to more easily find mates, both influenced by population density, are component Allee effects, as they influence individual fitness of the population. At low population density, these component Allee effects would add up to produce an overall demographic Allee effect (increased fitness with higher population density). When population density reaches a high number, negative density dependence often offsets the component Allee effects through resource competition, thus erasing the demographic Allee effect. Allee effects might occur even at high population density for some species. For example, wind pollinated plants would have a lower fitness in sparse populations due to the lower likelihood of pollen successfully landing on a conspecific. Cooperative defense Another possible benefit of aggregation is to protect against predation by group anti predator behavior. Many species exhibit higher rates of predator vigilance behavior per individual at lower density. This increased vigilance might result in less time and energy spent on foraging, thus reducing the fitness of an individual living in smaller groups. One striking example of such shared vigilance is exhibited by meerkats. Meanwhile, other species move in synchrony to confuse and avoid predators such as schools of sardines and flocks of starlings. The confusion effect that this herding behavior would have on predators will be more effective when more individuals are present. While compelling, this case of environmental conditioning by salmon has not been rigorously supported by empirical evidence.

Classic economic theory predicts that human exploitation of a population is unlikely to result in species extinction because the escalating costs to find the last few individuals will exceed the fixed price one achieves by selling the individuals on the market. However, when rare species are more desirable than common species, prices for rare species can exceed high harvest costs. This phenomenon can create an "anthropogenic" Allee effect where rare species go extinct but common species are sustainably harvested. The anthropogenic Allee effect has become a standard approach for conceptualizing the threat of economic markets on endangered species. However, the original theory was posited using a one dimensional analysis of a two dimensional model. This overall fitness decrease of a species is caused by an accumulation of deleterious mutations throughout the population. Genetic variation within a species could range from beneficial to detrimental. Nevertheless, in a smaller sized gene pool, there is a higher chance of a stochastic event in which deleterious alleles become fixed (genetic drift). While evolutionary theory states that expressed deleterious alleles should be purged through natural selection, purging would be most efficient only at eliminating alleles that are highly detrimental or harmful. Mildly deleterious alleles such as those that act later in life would be less likely to be removed by natural selection, and conversely, newly acquired beneficial mutations are more likely to be lost by random chance in smaller genetic pools than larger ones. Although the long term population persistence of several species with low genetic variation has recently prompted debate on the generality of inbreeding depression, there are various empirical evidences for genetic Allee effects. One such case was observed in the endangered Florida panther (Puma concolor coryi). The Florida panther experienced a genetic bottleneck in the early 1990s where the population was reduced to ≈25 adult individuals. This reduction in genetic diversity was correlated with defects that include lower sperm quality, abnormal testosterone levels, cowlicks, and kinked tails. In response, a genetic rescue plan was put in motion and several female pumas from Texas were introduced into the Florida population. This action quickly led to the reduction in the prevalence of the defects previously associated with inbreeding depression. Although the timescale for this inbreeding depression is larger than of those more immediate Allee effects, it has significant implications on the long term persistence of a species.

Demographic stochasticity refers to variability in population growth arising from sampling random births and deaths in a population of finite size. In small populations, demographic stochasticity will decrease the population growth rate, causing an effect similar to the Allee effect, which will increase the risk of population extinction. Whether or not demographic stochasticity can be considered a part of Allee effect is somewhat contentious however. The most current definition of Allee effect considers the correlation between population density and mean individual fitness. Therefore, random variation resulting from birth and death events would not be considered part of Allee effect as the increased risk of extinction is not a consequence of the changing fates of individuals within the population. Meanwhile, when demographic stochasticity results in fluctuations of sex ratios, it arguably reduces the mean individual fitness as population declines. For example, a fluctuation in small population that causes a scarcity in one sex would in turn limit the access of mates for the opposite sex, decreasing the fitness of the individuals within the population. This type of Allee effect will likely be more prevalent in monogamous species than polygynous species.

Clumping results due to individuals aggregating in response to: local habitat or landscape differences, daily and seasonal weather changes, reproductive processes, or as the result of social attractions.