Carrying Capacity of The Environment
Carrying Capacity of The Environment
Carrying Capacity of The Environment
Cang Hui, Centre for Invasion Biology, Department of Mathematical Sciences, Stellenbosch University, Matieland, South Africa; and
African Institute for Mathematical Sciences, Muizenburg, South Africa
Ó 2015 Elsevier Ltd. All rights reserved.
This article is a revision of the previous edition article by M.E. Geores, volume 10, pp. 7038–7039, Ó 2001, Elsevier Ltd.
Abstract
Catton defines carrying capacity as the maximum persistently supportable load of a focal population in a bounded envi-
ronment. Here, definitions of carrying capacity for natural populations, ecosystems and humans, as well as biocapacity, are
presented, with issues fully discussed pertaining to their proper interpretation, models and estimation, debate and derived
paradoxes. With the explosion of the human population, the threat of overstepping the earth’s carrying capacity has become
a sword of Damocles, hanging over humanity and urging it to choose a sustainable path of development. Carrying capacity
has become arguably the most important concept in the era of Anthropocene.
Humans are fascinated by growth dynamics, from the weight productivity of that habitat (Rees, 1992). The concept of
of an infant to the size of an economy. Two intrinsic ques- carrying capacity is rooted in the logistic equation depicting
tions on any growth dynamics are how fast and how long population dynamics under simple density-dependent regu-
the system can grow. That is, the rate and the bound of lation and has been developed across different hierarchical
growth. An unbounded growth is infeasible due to limited levels of living systems and in many sectors of human society
resources, and carrying capacity is a measure of this limita- (Seidl and Tisdell, 1999; Monte-Luna et al., 2004). In the
tion. In 1798, Thomas Malthus proposed an exponential Malthusian equation, the probability that an organism will
growth curve in his Essay on the Principle of Population, stating reproduce or die is assumed to remain constant and is inde-
that a population with constant fecundity greater than its pendent of the density of the population. Obviously, this can
mortality will grow exponentially till it causes inevitable only be true when there is no competition among the indi-
conflicts with the limited supply of resources. Deminishing viduals, normally when the population density is low. The
resources and the expanding population will inevitably growth of any population in a limited environment will
lead to scramble competition. Malthus’s idea has further eventually halt due to the shortage of resources. When the
inspired Charles Darwin to adopt resource competition as demand of the existing population on the resource (i.e.,
one prerequisite of evolution via natural selection. Darwin the population size times the basic per capita resource
said in his Autobiography that, “I happened to read for amuse- consumption rate required for maintaining life) is equal to
ment Malthus on Population and being well prepared to the rate of resource supply, the population will reach its satu-
appreciate the struggle for existence which everywhere goes ration level. This saturation level is decided by both the
on from long-continued observation of the habits of animals resource supply and the per capita resource consumption
and plants, it at once struck me that under these circum- rate, called the carrying capacity of the environment for the
stances favourable variations would tend to be preserved, focal population. Using the language of differential equa-
and unfavourable ones to be destroyed. The result of this tions in mathematics, the Verhulst logistic equation reads
would be the formation of new species.” Such struggle of as follows:
life will ensure only the survival of the fittest, or equivalently
dN K N
as Russel Wallace put, the elimination of the unfit. Such ¼ rN
dt K
a density-dependent mechanism of population regulation
that intense competition happens when resource demand where N is the population size; dN/dt the derivative of pop-
approaches supply was formally formulated by Pierre ulation size, indicating the change rate of the population size; r
Verhulst in 1838, by adding a parameter K into the the intrinsic population growth rate, indicating how fast the
Malthusian equation to curb the unbounded growth. This population can grow at small size; K the carrying capacity. The
equation is now well-known as the logistic equation, and the Malthusian equation only includes the first part of the right
parameter K the carrying capacity. term, dN/dt ¼ rN, meaning that the population change rate is
proportional to its current population size. It also means that
the per capita rate of increase (dN/dt)/N, often an indicator of
Population Carrying Capacity population fitness, is a constant and thus density independent.
In the logistic equation, the per capita rate of increase,
Catton (1986), defined an environment’s carrying capacity as r(K – N)/K, is a declining function of current population size N,
its maximum persistently supportable load. In ecology, thus becoming density dependent. In Figure 1, we can clearly
carrying capacity is normally defined as the maximum popu- see the difference between Malthusian’s exponential growth
lation of a given species that can be supported indefinitely curve and the saturation form of the logistic equation. Clearly,
in a defined habitat without permanently impairing the when K approaches infinity, the logistic equation becomes the
International Encyclopedia of the Social & Behavioral Sciences, 2nd edition, Volume 3 http://dx.doi.org/10.1016/B978-0-08-097086-8.91002-X 155
International Encyclopedia of the Social & Behavioral Sciences, Second Edition, 2015, 155–160
156 Carrying Capacity of the Environment
International Encyclopedia of the Social & Behavioral Sciences, Second Edition, 2015, 155–160
Carrying Capacity of the Environment 157
Rees (1992) suggested that carrying capacity is the number of biomass) and biodiversity (often measured by species richness
individuals of a given species that a given habitat can support or the Shannon index that considers the relative abundance of
without being permanently damaged. If the population of species). The directionality of succession can potentially be
a given species exceeds the carrying capacity of a given habitat, indicated by the increase of net primary production. Per capita
then either the resources required to meet the needs of that community productivity can be very high during the initial
species will be depleted, or the wastes produced by that species phases of succession, and decreases progressively as an upper
will build to the point of poisoning members of the species, or limit to biomass is reached, and this phenomenon may be
both, and the population will crash. In livestock management, expressed as a sigmoid curve roughly akin to the logistic growth
carrying capacity is affected by many factors, such as habitat, curve of a population. The upper limit of net primary produc-
climate, vegetation, soil profile, food quantity and quality tion, or the maximum potential biomass, has been suggested to
and accessibility, inter- and intracompetition, foraging, para- indicate the carrying capacity of biomass in a community
sitism, diseases, population density, social behavior, and (Monte-Luna et al., 2004).
anthropogenic disturbance. In terms of nutrition intake, If the number of species in a habitat is relatively low, immi-
carrying capacity has been defined as the accessible and utiliz- gration and diversification through disruptive selection will
able energy in the habitat divided by per capita energy take place, which progresses until the region attains
consumption (Ayllon et al., 2012). In terms of space, carrying a maximum supportable load of species (Monte-Luna et al.,
capacity is also affected by aggregation or social group 2004). This can also define the directionality of succession
behavior, group size, dispersal distance, and habitat suitability, and, thus, the carrying capacity of biodiversity, depicting the
and can be defined as the amount of habitat available divided maximum number of species or the ‘biodiversity ceiling’ that
by the expected individual territory area for a given life stage an environment can support. In fact, the form of species diver-
(Ayllon et al., 2012). It is closely related to subsistence density, sification on evolutionary timescales resembles the logistic
tolerance density, security density, maximum harvest density, population growth. By analogy, the number of species resem-
minimum impact density in livestock and wildlife manage- bles the number of individuals or biomass, and the difference
ment (Hobbs and Hanley, 1990). between rate of speciation and extinction the difference
Carrying capacity is not a static number but is affected by between fecundity and mortality (i.e., the intrinsic growth
the abundance and distribution of limited resources and by rate). The carrying capacity of biodiversity may be regulated
how individuals compete for these limiting resources (Ayllon by both physical factors such as climate and habitat heteroge-
et al., 2012). This notion is especially relevant in organisms neity and biotic interactions between species. Once the biodi-
that compete via both exploitation and interference because versity reaches its ceiling, interspecific interactions can lead to
of behavioral responses, such as competition avoidance, competitive exclusion of some taxa, and adaptive niche parti-
induced by aggressive interactions typically result in a much tioning (Monte-Luna et al., 2004). This can be further related
reduced exploitation of the limited resource than could be to the r/K selection theory in ecology. It specifies a life-
accounted for by resource depletion alone (Ayllon et al., history tradeoff between the values of r and K that a species
2012). In territorial species, the behavioral adjustment of the can possess. Species with high r and low K are opportunistic
size and shape of territories has profound consequences for and often occur at the early stage of succession, whereas species
their population regulation, demography, and spatial ecology. with low r and high K are good competitors for limiting
In practice, carrying capacity is determined by the resource in resources and often occur at the late stage of succession. An
the least supply, ‘the weakest link’ as it were. For production intermediate level of disturbance in a landscape, arguably,
purposes, accurate estimation of carrying capacity will affect can create patches at different levels of succession, thus
the maximum sustainable yield, which is thought to be ob- promoting the coexistence of these two types of species at
tained when the population is maintained at half of the the regional scale.
carrying capacity (Ayllon et al., 2012). By contrast, in conserva-
tion efforts, the maximum carrying capacity is the desired
target. Human Carrying Capacity
International Encyclopedia of the Social & Behavioral Sciences, Second Edition, 2015, 155–160
158 Carrying Capacity of the Environment
International Encyclopedia of the Social & Behavioral Sciences, Second Edition, 2015, 155–160
Carrying Capacity of the Environment 159
International Encyclopedia of the Social & Behavioral Sciences, Second Edition, 2015, 155–160
160 Carrying Capacity of the Environment
sapiens 200 000 years to reach its first billion in the early nine- Bibliography
teenth century, the second billion took only another 100 years.
Standing at 7 billion currently, the human population is pro- Arrow, K., Bolin, B., Constanza, R., Dasgupta, P., Folke, C., Holling, C.S.,
jected to reach 10 billion in the middle of this century. The Jansson, B.O., Levin, S., Maler, K.G., Perrings, C., Pimentel, D., 1995. Economic
growth, carrying capacity, and the environment. Science 286, 520–521.
sheer number and the increasing per capita consumption rate
Ayllon, D., Almodovar, A., Nicola, G.G., Parra, I., Elvira, B., 2012. Modelling carrying
in this era of Anthropocene place an enormous pressure on capacity dynamics for the conservation and management of territorial salmonids.
the planet’s resource supply, both renewable and unrenewable. Fisheries Research 134–136, 95–103.
Humanity as a single species consumes more than one-fifth of Berck, P., Levy, A., Chowdhury, K., 2012. An analysis of the world’s environment and
the planet’s net primary production each year. Will this great population dynamics with varying carrying capacity, concerns and scepticism.
Ecological Economics 73, 103–112.
era of modern humanity continue to grow indefinitely; fluc- Catton, W., 1986. Carrying Capacity and the Limits to Freedom. Paper Prepared for
tuate around a sustainable level after a soft landing; or follow Social Ecology Session 1. XI World Congress of Sociology, New Delhi, India.
the boom–bust pattern of many earlier civilizations? The Cohen, J.E., 1995. Population growth and Earth’s human carrying capacity. Science
concept of carrying capacity has become a great reminder, 269, 341–346.
Daily, G.C., Ehrlich, P.R., 1992. Population, Sustainability, and Earth’s Carrying
a type of ‘sword of Damocles,’ hanging over humanity and
Capacity. Paper No. 0046. Department of Biological Sciences, Stanford University,
driving it onto a sustainable path. Stanford.
Second, as human population continues to expand, the Franck, S., von Bloh, W., Muller, C., Bondeau, A., Sakschewski, B., 2011. Harvesting
planet’s pristine ecosystems are degrading at a rapid rate due the Sun: new estimations of the maximum population of planet Earth. Ecological
to man-made global environmental changes – climate change, Modelling 222, 2019–2026.
Hobbs, N.T., Hanley, T.A., 1990. Habitat evaluation – do use availability data reflect
habitat loss, biological invasion, soil erosion, and pollution, carrying capacity. Journal of Wildlife Management 54, 515–522.
to name a few. Ongoing agricultural intensification, overfishing, Hui, C., 2006. Carrying capacity, population equilibrium, and environment’s maximal
industrialization, and urbanization further transfer how semi- load. Ecological Modelling 192, 317–320.
natural ecosystems function and behave, creating many novel Monte-Luna, P.D., Brook, B.W., Zetina-Rejon, M.J., Cruz-Escalona, V.H., 2004. The
carrying capacity of ecosystems. Global Ecology and Biogeography 13, 485–495.
ecosystems that face high disturbance and gene flow. The
Rees, W.E., 1992. Ecological footprint and appropriated carrying capacity: what urban
planet’s greatest asset – biodiversity, 100 million species strong – economics leaves out. Environmental Urbanization 4, 121–130.
is quickly losing its grip. The rapid loss of species is estimated to Rees, W.E., Wackernagel, M., 1994. Ecological footprints and appropriated carrying
be between 1000 and 10 000 times higher than the natural capacity: measuring the natural capital requirements of the human economy. In:
extinction rate, reaching 10 000 species extinction per year Jansson, A.M., Hammer, M., Folke, C., Constanza, R. (Eds.), Investing in Natural
Capital: The Ecological Economics Approach to Sustainability. Island Press,
many of which disappear before they have even been formally Washington, DC, pp. 362–390.
described. Human society relies heavily on the health of Seidl, I., Tisdell, C.A., 1999. Carrying capacity reconsidered: from Mathus’ population
ecosystem functions, the provision of ecosystem services and theory to cultural carrying capacity. Ecological Economics 31, 395–408.
the benefit from biological diversity for cleaning air and water, Yue, D., Xu, X., Hui, C., Xiong, Y., Han, X., Ma, J., 2011. Biocapacity supply and
demand in Northwestern China: a spatial appraisal. Ecological Economics 70,
stabilizing weather, maintaining soil fertility, dissipating waste,
988–994.
controlling pests, pollinating crops, generating power and Yue, D., Guo, J., Hui, C., 2013. Scale dependency of biocapacity and the fallacy of
discovering new antibodies, and providing food, timber, cloth, unsustainable development. Journal of Environment Management 126, 13–19.
medicine, industrial material (coal, oil, gas, rubber, plastics,
chemicals), together with minerals. We are not making anything Relevant Websites
new but utilizing products and services of the planet’s ecosystem
and physical system. To sustain humanity, we need to manage http://www.carryingcapacity.com.au/
the planet’s biosphere within its bearable margin to avoid http://www.carryingcapacity.org/
disruptive regime shift and massive extinction, although some http://www.facingthefuture.org/
argue that we have already crossed the point of no return. On http://www.sustainablescale.org/
the positive side, the concept of carrying capacity is a powerful
tool for assessing and conserving natural capital.
International Encyclopedia of the Social & Behavioral Sciences, Second Edition, 2015, 155–160