Carrying Capacity of The Environment

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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

habitat is not equal to the maximum population equilibrium


that can survive through the inflow of propagules, known as
the rescue effect. This highlights the difference between carrying
capacity and the maximum population equilibrium. Indeed,
many preindustrial cities often benefited from being the hub
of trade routes and supported a large population beyond their
capacity.
In 1992, Lev R. Ginzburg considered incorporating addi-
tional mortality into the logistic equation without changing
any other aspects of the environment. One of his proposals
is: dN/dt ¼ rN(1  N/K)  mN, where mN(m > 0) is the addi-
tional mortality term. As the resources do not change in this
hypothetical scenario, intuitively the population with higher
mortality might attain the same K, but only slower. However,
the new population equilibrium is K(r  m)/r(m < r), lower
than the carrying capacity K. Once again, the Ginzburg paradox
is due to the confusion between the concept of carrying
capacity and population equilibrium. To illustrate, let us
Figure 1 An illustration of population dynamics governed by the consider a thought experiment (Hui, 2006). There are three
Malthusian equation and the logistic equation. Parameter values: eggs in a six-position egg box (Figure 2). If we eat one and
r ¼ N0 ¼ 1 and K ¼ 1000. then put a new one in it everyday, the size of the egg population
will be maintained at three. Now what is the carrying capacity
Malthusian equation. The exact solution to the logistic equa- of this egg population, three or six? Now, if we eat two eggs per
tion exists, day, the population equilibrium will become zero but the
carrying capacity remains six. According to our knowledge,
K$N0 $expðr$tÞ
NðtÞ ¼ the above equation does not have any mechanistic problem
K þ N0 ðexpðr$tÞ  1Þ
but has been widely used as a theoretical foundation in spatial
where N0 is the population size when time t ¼ 0. When time t and metapopulation ecology. This new stable equilibrium is
increases, the population size will converge to the carrying called the local carrying capacity of a metapopulation, whereas
capacity K (Figure 1). The logistic equation is the simplest other scientists call this equilibrium local density. Once again,
model, which contains a carrying capacity for the population. the concept of equilibrium was confused with carrying
In fact, carrying capacity exists in many models that implement capacity. This confusion also appears in some textbooks, yet
the negative density-dependence term, i.e., (dN/dt)/N is many others do differentiate between these two concepts. The
a declining function of N. carrying capacity of the egg box example should be the ‘envi-
Because the carrying capacity is also the asymptote, or the ronment’s maximal load,’ i.e., the six positions in the egg
equilibrium of the population size in the logistic equation, box, and the equilibrium of egg population is three if we eat
the concept of carrying capacity has often been confused with one and add one per day. If we can clarify these two concepts:
population equilibrium, leading to unnecessary confusion carrying capacity and population equilibrium (Hui, 2006),
and so-called paradoxes. In practice, many studies have consid- there is no contradiction between intuition and there should
ered the maximum population equilibrium or maximum pop- be no Ginzburg’s paradox.
ulation size as the carrying capacity, making the distinction Ecological carrying capacity is a measure of the amount of
even muddier. In 1969, Richard Levins hypothesized that if r renewable resources in the environment in units of the number
in the logistic equation is negative, then we will have a positive of organisms these resources can support. Normally, K is a func-
dN/dt when N > K and thus the population size will increase tion of both the species and the environment, and is expected
indefinitely. This is clearly counterintuitive. Although con- to change only in evolutionary time (Daily and Ehrlich, 1992).
straining r to be only positive can make Levins’ paradox disap-
pear, many declining populations are indeed experiencing the
situation of r < 0, thus such a constraint could exclude many
biologically meaningful scenarios. Although constraining r
and K to be the same sign could resolve Levins’ paradox,
such a constraint nonetheless incurs a new problem on the
meaning of negative carrying capacity (K < 0). The negative
carrying capacity could be a measure of just how unfavorable
the environment is (Hui, 2006). An important finding from
metapopulation theory is that the source–sink dynamics are
such that where a local population is in a ‘sink habitat’ a posi-
tive equilibrium can still be maintained due to contributions
from migrants from source populations. These sink patches
have a negative carrying capacity but support a positive popu- Figure 2 A population of three eggs at the equilibrium in an egg box
lation equilibrium. Clearly, the carrying capacity of a sink with the six-holder carrying capacity.

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

Our planet is practically a closed system in terms of physical


Carrying Capacity of Ecosystems resources, and all human activity depends on these limited
resources. Due to the continue growing of the world popula-
The directionality of community succession is a powerful tion and the rapid diminishing of pristine ecosystems from
concept for conservation biology, analogous to the irrevers- our exploration for consumption, the concept of carrying
ibility of time in physics, and has revolutionized the capacity is essential and should be considered paramount
understanding of complex adaptive systems. By definition, in our future planning. The Club of Rome has warned us
succession is ‘an orderly process of community change’ after the possible consequences of world population growth and
disturbance. Knowing the directionality of succession is neces- industrialization for food production and resource exhaus-
sary for (1) distinguishing new from mature communities (i.e., tion. They projected that within 50 years from now, the
defining the age of a community), (2) understanding how planet would reach its limit of growth (Figure 3), and thus
communities evolve and respond to disturbance (e.g., habitat advocated an urgent mutually beneficial integration of
loss and climate change), and (3) designing more efficient economics and ecology. Approaching carrying capacity
conservation and restoration plans. Two important concepts means the deterioration of ecosystem services, the loss of
of an ecosystem are productivity (often measured by the total biodiversity and habitat heterogeneity and the break down

International Encyclopedia of the Social & Behavioral Sciences, Second Edition, 2015, 155–160
158 Carrying Capacity of the Environment

agriculture and other forms of food supply compatible with


the aims of world development programs.
In a Malthusian framework, economic growth generates
population growth; the population growth increases pressure
on the natural resource base; the deterioration of the natural
resource base decreases per capita food output; and the reduced
per capita food limits population growth which, in turn, limits
exploitation of the environment (Berck et al., 2012). This
moderating feedback loop (density dependence) can be
diverged by technological improvements. Population pressure
on the resource base stimulates technological improvements
in food production and hence facilitates continued population
growth, which intensifies exploitation of the resource base
(Berck et al., 2012). Alternatively, as income goes up there is
increasing environmental degradation up to a point, after
which environmental quality improves, causing an inverted-
U shape (Arrow et al., 1995). Moreover, the detrimental impact
from human society to ecosystems is often discounted over
Figure 3 World human population from 10 000 BC to AD 2050 when time and distance in decision-making, i.e., discounting future
the carrying capacity of 10 billion is projected to be reached. and faraway activities than can reduce carrying capacity. This
encourages behavior that may reduce carrying capacity for
future generations (Daily and Enrlick, 1992). Globalization
of ecosystem resilience (Arrow et al., 1995). This could lead further creates the illusion that all regions could simulta-
to irreversible change, enlarged uncertainty to the biosphere, neously sustain populations that sum to more than global K,
and even a regime shift of the world’s living and climate a typical Netherlands fallacy.
systems. This concern has greatly influenced the popularity
of environmentalism and the creation of the United Nations
Environment Program. Biocapacity
Carrying capacity is commonly assumed to be constant in
population dynamics models used for resource management. Since the concept of sustainable development was put forward,
Human carrying capacity, however, is not a fixed, static, or it has become an ideal development mode and a common
a simple relationship, but is dynamic and adaptive. Humans policy goal. To date, many indicators have been developed to
can have more than one K value for a given resource at a given assess the status of sustainable development, such as the life-
time, and the carrying capacity can vary markedly with culture cycle assessment, human development index by the United
and level of economic development (Daily and Enrlick, 1992). Nations Development Programme, barometer of sustainability,
Earth’s human carrying capacity cannot be characterized by index of sustainable economic welfare, environmental pressure
a simple monotonic decline of the natural capital as the indicator, genuine progress indicator, sustainable technology
number of humans increase. For instance, Cohen (1995) development, environmental sustainability index, and ecolog-
studied a dynamic K model, with the change of carrying ical footprint (Rees, 1992). The latter has gained popularity
capacity affected by the population change rate. due to its compatibility with the data format commonly
Human carrying capacity describes the number of human derived from economic and social surveys. The ecological foot-
beings that can be supported on a sustainable basis in a given print for a particular population is defined as the total area of
area (or on the whole Earth) within natural resource limits productive land and water ecosystems required to produce
and by human choices concerning social, cultural, and sufficient resources and assimilate wastes (Rees and
economic conditions (Franck et al., 2011). This concept is Wackernagel, 1994). Biocapacity can be defined as the locally
twofold: Biophysical carrying capacity is the maximum popu- available carrying capacity of the ecosystem for generating
lation that can be supported by the resources of planet Earth resources and absorbing wastes (Rees and Wackernagel,
at a given level of technology, whereas social carrying capacity 1994) and is constrained by the carbon sequestration rate of
is the sustainable biophysical carrying capacity within a given the ecosystem. According to the global average rate of carbon
social organization (Franck et al., 2011). In the case of very sequestration, we need to have about 17 ha of land and ocean
efficient agriculture the carrying capacity is only determined area to absorb 1 metric ton of carbon emission. Ecological foot-
by the ratio of the total productive area and the nonagricul- print and biocapacity, thus, represent the demand on and the
tural used area (per person). For 2005 land cover, it can supply from a regional ecosystem, respectively. As both ecolog-
support 11.4 billion (109) (Franck et al., 2011), ranging ical footprint and biocapacity are measured in the same unit
from 7.7 to 12 billion in literature (Cohen, 1995). It has (the global hectare: gha), it is straightforward to calculate
been predicted to be much higher than this range under regional ecological budget as surplus and deficit, after discount-
certain scenarios, ranging from 30 to 100 billion (Franck ing the influence of trading. This specifies whether a regional
et al., 2011). It is the limiting resource that constrains human population is potentially self-sufficient or is at least partially
carrying capacity. Although there exists the technology and reliant on imported biocapacity. An ecological surplus has
economic willingness to develop intensive sustainable been proposed as a minimum criterion for sustainability.

International Encyclopedia of the Social & Behavioral Sciences, Second Edition, 2015, 155–160
Carrying Capacity of the Environment 159

Due to it being highly operable and easy to understand for


public and policy makers, with the data required accessible
from government yearbooks, to date they have been applied
at a variety of spatial scales, from municipality/provincial level
to national/global extents, covering all aspects of socioeco-
nomic sectors, such as industry, education, agriculture,
tourism, and waste management.
As a continuously developing field, the ecological footprint
methodology has been widely criticized and amended. For
instance, it has been considered a static indicator of weak
sustainability as no dynamics and bounds are imposed on
the level of ecosystem services and their demands. This has
been partially solved by time series analysis and extrapolation
(Yue et al., 2013). The introduction of spatial features, with the
help of the geographic information system (GIS), has largely
released the methodology from this constraint. To calculate
the biocapacity of a region, one first needs to estimate the avail-
able areas of biologically productive land and water. Specifi-
cally, this biologically productive area can normally be Figure 4 The scale dependency of the biocapacity in Jinghe River
divided into six or more main categories (cropland, grazing Watershed, Northwest China.
land, fishing land, forest, built-up area, and barren land), and
the sizes of these six land covers can then be either extracted
from government agencies or calculated using remote sensing
images with the aid of GIS (Yue et al., 2013):
X
K¼ Ai $ Yi $ Ei
i

where Ai is the biologically productive area of land cover


category i; yi is the yield factor of land category i and is calcu-
lated annually as the ratio of the local yield of a generic product
to the global average yield of the same product. The yield factor
converts local biologically productive land into units of global
average productivity and thus facilitates comparisons across
regions; Ei represents the equivalence factor of land cover
category i and is a scaling factor needed for converting a specific
land use type into a universal unit of biologically productive
area (gha). Equivalence factor is calculated each year as the
ratio of the global average productivity of a specific land type to
the average productivity of all biologically productive land on
the earth.
An important issue that is associated with area-based
information is the scale dependency of spatial features. Figure 5 The relationship between population density and biocapacity
Evidently, the shape and size of different land covers are sensi- per hectare for 320 counties in Northwest China.
tive to the spatial scale as most landscape features are scale
dependent and have self-similar, fractal structures. Put simply,
results for biocapacity can be influenced by the resolution of overshoot, indicating that regional development is not
the map (Yue et al., 2013). Since area-based information ecologically self-sustainable. If the biocapacity pressure index
has been widely implemented for estimating the sizes of is less than one, the region is potentially self-sufficient (Yue
different land covers and therefore the biocapacity, it is et al., 2011).
important to at least be aware of the scale dependency of bio-
capacity and its impact on regional sustainability assessment
(Figure 4). Moreover, there is often a power–law relationship Relevance in Anthropocene
between the biocapacity and population density (Figure 5),
suggesting that biocapacity serves not only as a support for The growing number of studies on carrying capacity and its
social development and human well-being but also sets an further development into different research fields have made
ecological limit for human activities (Yue et al., 2011). The this concept the most important issue in this century, for
biocapacity pressure index is the regional ecofootprint to bio- obvious reasons. First, the world has entered a rather peaceful
capacity ratio and reflects the pressure posed by the human era after the World War II, allowing both the living condition
population on the local ecosystem. If the biocapacity pressure to proliferate and the economy to boom at a staggering pace.
index is greater than one, the population is in ecological Although it took the modern human population of Homo

International Encyclopedia of the Social & Behavioral Sciences, Second Edition, 2015, 155–160
160 Carrying Capacity of the Environment

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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.

See also: Environment and Development; Environmental


Movements; Food Security and ‘Green Revolution’; Limits to
Growth; Sustainable Development: An Economic Perspective.

International Encyclopedia of the Social & Behavioral Sciences, Second Edition, 2015, 155–160

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