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19.2: Population Growth and Regulation

Population ecologists make use of a variety of methods to model population dynamics. An accurate model should be able to describe the changes occurring in a population and predict future changes.

Population Growth

The two simplest models of population growth use deterministic equations (equations that do not account for random events) to describe the rate of change in the size of a population over time. The first of these models, exponential growth, describes theoretical populations that increase in numbers without any limits to their growth. The second model, logistic growth, introduces limits to reproductive growth that become more intense as the population size increases. Neither model adequately describes natural populations, but they provide points of comparison.

Exponential Growth

Charles Darwin, in developing his theory of natural selection, was influenced by the English clergyman Thomas Malthus. Malthus published his book in 1798 stating that populations with abundant natural resources grow very rapidly; however, they limit further growth by depleting their resources. The early pattern of accelerating population size is called exponential growth.

The best example of exponential growth in organisms is seen in bacteria. Bacteria are prokaryotes that reproduce largely by binary fission. This division takes about an hour for many bacterial species. If 1000 bacteria are placed in a large flask with an abundant supply of nutrients (so the nutrients will not become quickly depleted), the number of bacteria will have doubled from 1000 to 2000 after just an hour. In another hour, each of the 2000 bacteria will divide, producing 4000 bacteria. After the third hour, there should be 8000 bacteria in the flask. The important concept of exponential growth is that the growth rate—the number of organisms added in each reproductive generation—is itself increasing; that is, the population size is increasing at a greater and greater rate. After 24 of these cycles, the population would have increased from 1000 to more than 16 billion bacteria. When the population size, N, is plotted over time, a J-shaped growth curve is produced (Figure 19.2.1a).

The bacteria-in-a-flask example is not truly representative of the real world where resources are usually limited. However, when a species is introduced into a new habitat that it finds suitable, it may show exponential growth for a while. In the case of the bacteria in the flask, some bacteria will die during the experiment and thus not reproduce; therefore, the growth rate is lowered from a maximal rate in which there is no mortality. The growth rate of a population is largely determined by subtracting the death rate, D, (number organisms that die during an interval) from the birth rate, B, (number organisms that are born during an interval). The growth rate can be expressed in a simple equation that combines the birth and death rates into a single factor: r. This is shown in the following formula:

Population growth = rNPopulation growth = rN

The value of r can be positive, meaning the population is increasing in size (the rate of change is positive); or negative, meaning the population is decreasing in size; or zero, in which case the population size is unchanging, a condition known as zero population growth.

Logistic Growth

Extended exponential growth is possible only when infinite natural resources are available; this is not the case in the real world. Charles Darwin recognized this fact in his description of the “struggle for existence,” which states that individuals will compete (with members of their own or other species) for limited resources. The successful ones are more likely to survive and pass on the traits that made them successful to the next generation at a greater rate (natural selection). To model the reality of limited resources, population ecologists developed the logistic growth model.

Carrying Capacity and the Logistic Model

In the real world, with its limited resources, exponential growth cannot continue indefinitely. Exponential growth may occur in environments where there are few individuals and plentiful resources, but when the number of individuals gets large enough, resources will be depleted and the growth rate will slow down. Eventually, the growth rate will plateau or level off (Figure 19.2.1b). This population size, which is determined by the maximum population size that a particular environment can sustain, is called the carrying capacity, or K. In real populations, a growing population often overshoots its carrying capacity, and the death rate increases beyond the birth rate causing the population size to decline back to the carrying capacity or below it. Most populations usually fluctuate around the carrying capacity in an undulating fashion rather than existing right at it.

The formula used to calculate logistic growth adds the carrying capacity as a moderating force in the growth rate. The expression “K – N” is equal to the number of individuals that may be added to a population at a given time, and “K – N” divided by “K” is the fraction of the carrying capacity available for further growth. Thus, the exponential growth model is restricted by this factor to generate the logistic growth equation:

Population growth = rN [K−NK]Population growth = rN [ K−NK ]

Notice that when N is almost zero the quantity in brackets is almost equal to 1 (or K/K) and growth is close to exponential. When the population size is equal to the carrying capacity, or N = K, the quantity in brackets is equal to zero and growth is equal to zero. A graph of this equation (logistic growth) yields the S-shaped curve (Figure 19.2.1b). It is a more realistic model of population growth than exponential growth. There are three different sections to an S-shaped curve. Initially, growth is exponential because there are few individuals and ample resources available. Then, as resources begin to become limited, the growth rate decreases. Finally, the growth rate levels off at the carrying capacity of the environment, with little change in population number over time.

Figure 19.2.1: When resources are unlimited, populations exhibit (a) exponential growth, shown in a J-shaped curve. When resources are limited, populations exhibit (b) logistic growth. In logistic growth, population expansion decreases as resources become scarce, and it levels off when the carrying capacity of the environment is reached. The logistic growth curve is S-shaped.

Role of Intraspecific Competition

The logistic model assumes that every individual within a population will have equal access to resources and, thus, an equal chance for survival. For plants, the amount of water, sunlight, nutrients, and space to grow are the important resources, whereas in animals, important resources include food, water, shelter, nesting space, and mates.

In the real world, phenotypic variation among individuals within a population means that some individuals will be better adapted to their environment than others. The resulting competition for resources among population members of the same species is termed intraspecific competition. Intraspecific competition may not affect populations that are well below their carrying capacity, as resources are plentiful and all individuals can obtain what they need. However, as population size increases, this competition intensifies. In addition, the accumulation of waste products can reduce carrying capacity in an environment.

Examples of Logistic Growth

Yeast, a microscopic fungus used to make bread and alcoholic beverages, exhibits the classical S-shaped curve when grown in a test tube (Figure 19.2.2a). Its growth levels off as the population depletes the nutrients that are necessary for its growth. In the real world, however, there are variations to this idealized curve. Examples in wild populations include sheep and harbor seals (Figure 19.2.2b). In both examples, the population size exceeds the carrying capacity for short periods of time and then falls below the carrying capacity afterwards. This fluctuation in population size continues to occur as the population oscillates around its carrying capacity. Still, even with this oscillation, the logistic model is confirmed.

ART CONNECTION

Figure 19.2.2: (a) Yeast grown in ideal conditions in a test tube shows a classical S-shaped logistic growth curve, whereas (b) a natural population of seals shows real-world fluctuation. The yeast is visualized using differential interference contrast light micrography. (credit a: scale-bar data from Matt Russell)

If the major food source of seals declines due to pollution or overfishing, which of the following would likely occur?

  1. The carrying capacity of seals would decrease, as would the seal population.
  2. The carrying capacity of seals would decrease, but the seal population would remain the same.
  3. The number of seal deaths would increase, but the number of births would also increase, so the population size would remain the same.
  4. The carrying capacity of seals would remain the same, but the population of seals would decrease.
Population Dynamics and Regulation

The logistic model of population growth, while valid in many natural populations and a useful model, is a simplification of real-world population dynamics. Implicit in the model is that the carrying capacity of the environment does not change, which is not the case. The carrying capacity varies annually. For example, some summers are hot and dry whereas others are cold and wet; in many areas, the carrying capacity during the winter is much lower than it is during the summer. Also, natural events such as earthquakes, volcanoes, and fires can alter an environment and hence its carrying capacity. Additionally, populations do not usually exist in isolation. They share the environment with other species, competing with them for the same resources (interspecific competition). These factors are also important to understanding how a specific population will grow.

Population growth is regulated in a variety of ways. These are grouped into density-dependent factors, in which the density of the population affects growth rate and mortality, and density-independent factors, which cause mortality in a population regardless of population density. Wildlife biologists, in particular, want to understand both types because this helps them manage populations and prevent extinction or overpopulation.

Density-dependent Regulation

Most density-dependent factors are biological in nature and include predation, inter- and intraspecific competition, and parasites. Usually, the denser a population is, the greater its mortality rate. For example, during intra- and interspecific competition, the reproductive rates of the species will usually be lower, reducing their populations’ rate of growth. In addition, low prey density increases the mortality of its predator because it has more difficulty locating its food source. Also, when the population is denser, diseases spread more rapidly among the members of the population, which affect the mortality rate.

Density dependent regulation was studied in a natural experiment with wild donkey populations on two sites in Australia.1 On one site the population was reduced by a population control program; the population on the other site received no interference. The high-density plot was twice as dense as the low-density plot. From 1986 to 1987 the high-density plot saw no change in donkey density, while the low-density plot saw an increase in donkey density. The difference in the growth rates of the two populations was caused by mortality, not by a difference in birth rates. The researchers found that numbers of offspring birthed by each mother was unaffected by density. Growth rates in the two populations were different mostly because of juvenile mortality caused by the mother’s malnutrition due to scarce high-quality food in the dense population. Figure 19.2.3 shows the difference in age-specific mortalities in the two populations.

Figure 19.2.3: This graph shows the age-specific mortality rates for wild donkeys from high- and low-density populations. The juvenile mortality is much higher in the high-density population because of maternal malnutrition caused by a shortage of high-quality food.

Density-independent Regulation and Interaction with Density-dependent Factors

Many factors that are typically physical in nature cause mortality of a population regardless of its density. These factors include weather, natural disasters, and pollution. An individual deer will be killed in a forest fire regardless of how many deer happen to be in that area. Its chances of survival are the same whether the population density is high or low. The same holds true for cold winter weather.

In real-life situations, population regulation is very complicated and density-dependent and independent factors can interact. A dense population that suffers mortality from a density-independent cause will be able to recover differently than a sparse population. For example, a population of deer affected by a harsh winter will recover faster if there are more deer remaining to reproduce.

EVOLUTION IN ACTION:Why Did the Woolly Mammoth Go Extinct?

Figure 19.2.4: The three images include: (a) 1916 mural of a mammoth herd from the American Museum of Natural History, (b) the only stuffed mammoth in the world is in the Museum of Zoology located in St. Petersburg, Russia, and (c) a one-month-old baby mammoth, named Lyuba, discovered in Siberia in 2007. (credit a: modification of work by Charles R. Knight; credit b: modification of work by “Tanapon”/Flickr; credit c: modification of work by Matt Howry)

Woolly mammoths began to go extinct about 10,000 years ago, soon after paleontologists believe humans able to hunt them began to colonize North America and northern Eurasia (Figure 19.2.4). A mammoth population survived on Wrangel Island, in the East Siberian Sea, and was isolated from human contact until as recently as 1700 BC. We know a lot about these animals from carcasses found frozen in the ice of Siberia and other northern regions.

It is commonly thought that climate change and human hunting led to their extinction. A 2008 study estimated that climate change reduced the mammoth’s range from 3,000,000 square miles 42,000 years ago to 310,000 square miles 6,000 years ago.2 Through archaeological evidence of kill sites, it is also well documented that humans hunted these animals. A 2012 study concluded that no single factor was exclusively responsible for the extinction of these magnificent creatures.3 In addition to climate change and reduction of habitat, scientists demonstrated another important factor in the mammoth’s extinction was the migration of human hunters across the Bering Strait to North America during the last ice age 20,000 years ago.

The maintenance of stable populations was and is very complex, with many interacting factors determining the outcome. It is important to remember that humans are also part of nature. Once we contributed to a species’ decline using primitive hunting technology only.

Demographic-Based Population Models

Population ecologists have hypothesized that suites of characteristics may evolve in species that lead to particular adaptations to their environments. These adaptations impact the kind of population growth their species experience. Life history characteristics such as birth rates, age at first reproduction, the numbers of offspring, and even death rates evolve just like anatomy or behavior, leading to adaptations that affect population growth. Population ecologists have described a continuum of life-history “strategies” with K-selected species on one end and r-selected species on the other. K-selected species are adapted to stable, predictable environments. Populations of K-selected species tend to exist close to their carrying capacity. These species tend to have larger, but fewer, offspring and contribute large amounts of resources to each offspring. Elephants would be an example of a K-selected species. r-selected species are adapted to unstable and unpredictable environments. They have large numbers of small offspring. Animals that are r-selected do not provide a lot of resources or parental care to offspring, and the offspring are relatively self-sufficient at birth. Examples of r-selected species are marine invertebrates such as jellyfish and plants such as the dandelion. The two extreme strategies are at two ends of a continuum on which real species life histories will exist. In addition, life history strategies do not need to evolve as suites, but can evolve independently of each other, so each species may have some characteristics that trend toward one extreme or the other.

Section Summary

Populations with unlimited resources grow exponentially—with an accelerating growth rate. When resources become limiting, populations follow a logistic growth curve in which population size will level off at the carrying capacity.

Populations are regulated by a variety of density-dependent and density-independent factors. Life-history characteristics, such as age at first reproduction or numbers of offspring, are characteristics that evolve in populations just as anatomy or behavior can evolve over time. The model of r- and K-selection suggests that characters, and possibly suites of characters, may evolve adaptations to population stability near the carrying capacity (K-selection) or rapid population growth and collapse (r-selection). Species will exhibit adaptations somewhere on a continuum between these two extremes.

Art Exercise

Figure 19.2.2 If the major food source of seals declines due to pollution or overfishing, which of the following would likely occur?

  1. The carrying capacity of seals would decrease, as would the seal population.
  2. The carrying capacity of seals would decrease, but the seal population would remain the same.
  3. The number of seal deaths would increase, but the number of births would also increase, so the population size would remain the same.
  4. The carrying capacity of seals would remain the same, but the population of seals would decrease.

A: The carrying capacity of seals would decrease, as would the seal population.

Multiple Choice

Species with limited resources usually exhibit a(n) ________ growth curve.

A. logistic
B. logical
C. experimental
D. exponential

A

The maximum growth rate characteristic of a species is called its ________.

A. limit
B. carrying capacity
C. biotic potential
D. exponential growth pattern

C

The population size of a species capable of being supported by the environment is called its ________.

A. limit
B. carrying capacity
C. biotic potential
D. logistic growth pattern

B

Species that have many offspring at one time are usually:

A. r-selected
B. K-selected
C. both r- and K-selected
D. not selected

A

A forest fire is an example of ________ regulation.

A. density-dependent
B. density-independent
C. r-selected
D. K-selected

B

Free Response

Describe the growth at various parts of the S-shaped curve of logistic growth.

In the first part of the curve, when few individuals of the species are present and resources are plentiful, growth is exponential, similar to a J-shaped curve. Later, growth slows due to the species using up resources. Finally, the population levels off at the carrying capacity of the environment, and it is relatively stable over time.

Give an example of how density-dependent and density-independent factors might interact.

If a natural disaster such as a fire happened in the winter, when populations are low, it would have a greater effect on the overall population and its recovery than if the same disaster occurred during the summer, when population levels are high.

Footnotes

  1. 1 David Choquenot, “Density-Dependent Growth, Body Condition, and Demography in Feral Donkeys: Testing the Food Hypothesis,” Ecology 72, no. 3 (June 1991):805–813.
  2. 2 David Nogués-Bravo et al., “Climate Change, Humans, and the Extinction of the Woolly Mammoth.” PLoS Biol 6 (April 2008): e79, doi:10.1371/journal.pbio.0060079.
  3. 3 G.M. MacDonald et al., “Pattern of Extinction of the Woolly Mammoth in Beringia.” Nature Communications 3, no. 893 (June 2012), doi:10.1038/ncomms1881.

Glossary

birth rate
the number of births within a population at a specific point in time
carrying capacity
the maximum number of individuals of a population that can be supported by the limited resources of a habitat
death rate
the number of deaths within a population at a specific point in time
density-dependent regulation
the regulation of population in which birth and death rates are dependent on population size
density-independent regulation
the regulation of population in which the death rate is independent of the population size
exponential growth
an accelerating growth pattern seen in populations where resources are not limiting
intraspecific competition
the competition among members of the same species
J-shaped growth curve
the shape of an exponential growth curve
K-selected species
a species suited to stable environments that produce a few, relatively large offspring and provide parental care
logistic growth
the leveling off of exponential growth due to limiting resources
r-selected species
a species suited to changing environments that produce many offspring and provide little or no parental care
S-shaped growth curve
the shape of a logistic growth curve

Contributors

zero population growth
the steady population size where birth rates and death rates are equal
 
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