population ecology

Introduction

      study of the processes that affect the distribution and abundance of animal and plant populations.

      A population is a subset of individuals of one species that occupies a particular geographic area and, in sexually reproducing species, interbreeds. The geographic boundaries of a population are easy to establish for some species but more difficult for others. For example, plants or animals occupying islands have a geographic range defined by the perimeter of the island. In contrast, some species are dispersed across vast expanses, and the boundaries of local populations are more difficult to determine. A continuum exists from closed populations that are geographically isolated from other populations of the same species to open populations that show varying degrees of connectedness.

Genetic variation within local populations

      In sexually reproducing species, each local population contains a distinct combination of genes. As a result, a species is a collection of populations that differ genetically from one another to a greater or lesser degree. These genetic differences manifest themselves as differences among populations in morphology, physiology, behaviour, and life histories; in other words, genetic characteristics ( genotype) affect expressed characteristics ( phenotype). natural selection initially operates on a phenotypic level, favouring or discriminating against expressed characteristics. The gene pool (total aggregate of genes in a population at a certain time) is affected as organisms with phenotypes that are compatible with the environment are more likely to survive for longer periods, during which time they can reproduce more often and pass on more of their genes.

      The amount of genetic variation within local populations varies tremendously, and much of modern conservation biology is concerned with the maintenance of genetic variation within populations of plants and animals. Some small, isolated populations of asexual species often have little genetic variation among individuals, whereas large, sexual populations often have great variation. Two major factors are responsible for this variety: mode of reproduction and population size.

Effects of mode of reproduction: sexual and asexual

      In sexual populations, genes are recombined in each generation, resulting in new genotypes. Offspring in most sexual species inherit half their genes from their mother and half from their father, and their genetic makeup is therefore different from either parent or any other individual in the population. New, favourable mutations that initially appear in separate individuals can be recombined in many ways over time within a sexual population. In contrast, the offspring of an asexual individual are genetically identical to their parent. The only source of new gene combinations in asexual populations is mutation.

      Asexual populations accumulate genetic variation only at the rate at which their genes mutate. Favourable mutations arising in different asexual individuals have no way of recombining and eventually appearing together in any one individual, as they do in sexual populations. A much wider range of favourable gene combinations, therefore, is likely to be found in sexual than asexual populations.

Effects of population size

      Over long periods of time, genetic variation is more easily sustained in large populations than in small populations. Through the effects of random genetic drift, a genetic trait can be lost from a small population relatively quickly (see biosphere: The organism and the environment: The diversity of life: The organization of the biosphere: Processes of evolution (biosphere)). For example, many populations have two or more forms of a gene, which are called alleles (allele). Depending on which allele an individual has inherited, a certain phenotype will be produced. If populations remain small for many generations, they may lose all but one form of each gene by chance alone.

      This loss of alleles happens from sampling error. As individuals mate, they exchange genes. Imagine that initially half of the population has one form of a particular gene, and the other half of the population has another form of the gene. By chance, in a small population the exchange of genes could result in all individuals of the next generation having the same allele. The only way for this population to contain a variation of this gene again is through mutation of the gene or immigration of individuals from another population (see evolution: Genetic variation in populations (evolution)).

      Minimizing the loss of genetic variation in small populations is one of the major problems faced by conservation biologists. Environments are constantly changing, and natural selection continually sorts through the genetic variation found within each population, favouring those individuals with phenotypes best suited for the current environment. Without the genetic variation that allows populations to respond evolutionarily to changes in the physical environment, diseases, predators, and competitors, populations risk extinction.

Population density and growth

Life histories and the structure of populations

      A life history is the sequence and timing of events that occur between birth and death. Populations from different parts of the geographic range that a species inhabits may exhibit marked variations in their life histories (see above Genetic variation within local populations (population ecology)). The patterns of variation seen within and among populations are referred to as the structure of populations. These variations include breeding frequency, age at which reproduction begins, the number of times an individual reproduces during its lifetime, the number of offspring produced at each reproductive episode (clutch size), the ratio of male to female offspring produced, and whether reproduction is sexual or asexual. These differences in life history characteristics can have profound effects on the dynamics, ecology, and evolution of populations.

      Of the many differences in life history that occur among populations, age at the time of first reproduction is one of the most important for understanding the dynamics and evolution of a population. All else being equal, natural selection will favour individuals that reproduce earlier than other individuals within a population, because by reproducing earlier an individual's genes enter the gene pool sooner than those of other individuals that were born at the same time but have not reproduced. The genes of the early reproducers then begin to spread throughout the population. Individuals whose genetic makeup allows them to reproduce earlier in life will come to dominate a population if there is no counterbalancing advantage to those individuals that delay reproduction until later in life.

      Not all populations, however, are made up of individuals that reproduce very early in life. In the course of a lifetime, an individual must devote energy and resources to physiological demands other than reproduction. To reproduce successfully, a plant first may have to grow to a certain height and outcompete its neighbours, and an animal may have to devote energy to growth so that it can reach a size at which it can fend off predators and successfully compete for mates. In many populations, individuals that delay reproduction have a better chance of surviving and leaving offspring than those that attempt to reproduce early. The opposing demands of growth, defense, and reproduction are balanced within the constraints of different environments to produce populations that have a diverse range of life-history strategies.

      Populations often can be divided into one of two extreme types, based on their life-history strategy. Some populations, called r-selected, are considered opportunistic because their reproductive behaviour involves a high intrinsic rate of growth (r)—individuals give birth once at an early age to many offspring. Populations that exhibit this strategy often have been shaped by an extremely variable and uncertain environment. Because mortality occurs randomly in this setting, quantity of progeny rather than quality of care serves the species better. In another strategy, called K-selected, populations tend to remain near the carrying capacity (K), the maximum number of individuals that the environment can sustain. Individuals in a K-selected population give birth at a later age to fewer offspring. This equilibrial life history is exhibited in more stable environments where reproductive success depends more on the fitness of the offspring rather than on their numbers.

Life tables and the rate of population growth

 Differences in life-history strategies greatly affect population dynamics. As stated above, populations in which individuals reproduce at an early age have the potential to grow much faster than populations in which individuals reproduce later. The effect of the age of first reproduction on population growth can be seen in the life tables for a particular species. Life tables were originally developed by insurance companies to provide a means of determining how long a person of a particular age could be expected to live. They are used not only by demographers of human populations but also by plant, animal, and microbial ecologists to make projections about the life expectancies of nonhuman populations. The number of individuals in a closed population (a population in which neither immigration nor emigration occurs) is governed by the rates of birth (natality), growth, reproduction, and death (mortality). Life tables are designed to evaluate how these rates influence the overall growth rate of a population (Table—>).

Survivorship curves (survivorship curve)

    Life tables follow the fate of a group of individuals all born in the same year. Of this group, or cohort, only a certain number of individuals will reach each age, and there is an age above which no individuals ever survive. Plotting the number of those members of the group that are still alive at each age results in a survivorship curve for the population (Figure 1—>). Survivorship curves are usually displayed on a semilogarithmic rather than an arithmetic scale. There are three general types of survivorship curves. Species such as humans and large mammals, which have fewer numbers of offspring but invest much time and energy in caring for their young (K-selected species), usually have a Type I curve (Figure 1—>). This relatively flat curve reflects low juvenile mortality, with most individuals living to old age. A constant probability of dying at any age, shown by the Type II curve (Figure 1—>), is evident as a straight line decreasing over time toward zero. Certain lizards, perching birds, and rodents exhibit this type of survivorship curve. In some species that produce many offspring but provide little care for them (r-selected species), mortality is greatest among the youngest individuals. The Type III survivorship curve indicative of this life history is initially very steep but flattens out as those individuals who reach maturity survive for a relatively longer time (Figure 1—>); it is exhibited by animals such as many insects or shellfish. Many populations have survivorship patterns that are more complex than, or fall in between, these three idealized curves. For example, passerine birds (perching birds such as finches) commonly suffer high mortality during the first year of life and a lower, more constant rate of death in subsequent years.

Calculating population growth

      Life tables also are used to study population growth. The average number of offspring left by a female at each age and the proportion of individuals surviving to each age can be used to evaluate the rate at which the size of the population changes over time. These rates are used by demographers and population ecologists to estimate human population growth and to evaluate the effect that conservation efforts have on endangered species.

 The average number of offspring that a female leaves during her lifetime is called the net reproductive rate (R₀). If all females survived to the oldest possible age for that population, the net reproductive rate would simply be the sum of the average number of offspring left by females at each age. In real populations, however, some females die at every age. The net reproductive rate for a set cohort is obtained by multiplying the proportion of females surviving to each age (l_x) by the average number of offspring produced at each age (m_x) and then adding the products from all the age groups: R₀ = Σl_xm_x. A net reproductive rate of 1.0 indicates that a population is neither increasing nor decreasing but replacing its numbers exactly. Any number below 1.0 indicates a decrease in population, any number above indicates an increase. In the example provided in the Table—>, the net reproductive rate is 2.101, which means that the population of the Galapagos cactus finch (Geospiza scandens) can double its size each generation.

      The other value needed to calculate the rate at which the population can grow is the mean generation time (T). Generation time is the average interval between the birth of an individual and the birth of its offspring. To determine the mean generation time of a population, the age of the individuals (x) is multiplied by the proportion of females surviving to that age (l_x) and the average number of offspring left by females at that age (m_x). This calculation is performed for each age group, and the values are added together and divided by the net reproductive rate (R₀) to yield the result:

 The mean generation time of the Galapagos cactus finch is 6.08 years (Table—>).

      The value that is used by population biologists to calculate the rate of increase of populations is the intrinsic rate of natural increase (r), or the Malthusian parameter. Very simply, this rate can be understood as number of births minus number of deaths per generation time—in other words, the reproduction rate less the death rate. To derive this value using a life table, the natural logarithm of the net reproductive rate is divided by the mean generation time:

       Intrinsic rate of increase (r) calculated for populations of species that differ greatly in their potential for the rate of population growthValues above zero indicate that the population is increasing; the higher the value, the faster the growth rate (Table (Intrinsic rate of increase (r) calculated for populations of species that differ greatly in their potential for the rate of population growth)). The Malthusian parameter can be used to compare growth rates of populations of a species that have different generation times. Some human populations have higher intrinsic rates of natural increase partially because individuals in those groups begin reproducing earlier than those in other groups. Mice have higher intrinsic rates of natural increase than elephants because they reproduce at a much earlier age and have a much shorter mean generation time.

      If a population has an intrinsic rate of natural increase of 0, then it is said to have stable age distribution and is neither growing nor declining in numbers. A growing population has more individuals in the lower age classes than does a stable population, and a declining population has more individuals in the older age classes than does a stable population (see population: Population composition (population)). Many human populations are currently undergoing population increase, far exceeding a stable age distribution. Although the human population has increased almost continuously throughout history, it has skyrocketed since the Industrial Revolution, primarily because of a drop in death rates. No other population has shown such steady growth.

Regulation of populations

Limits to population growth

Exponential and geometric population growth

      In an ideal environment, one that has no limiting factors, populations grow at a geometric rate or exponential rate. Human populations, in which individuals live and reproduce for many years and in which reproduction is distributed throughout the year, grow exponentially. Exponential population growth can be determined by dividing the change in population size (ΔN) by the time interval (Δt) for a certain population size (N):

      The growth curve of these populations is smooth and becomes increasingly steeper over time. The steepness of the curve depends on the intrinsic rate of natural increase for the population. Human population growth has been exponential during the 20th century. Much concern exists about the impact this growth will have, not only on the environment, but on humans as well. The World Bank projection for human population growth predicts that the human population will grow from 4.5 billion in 1995 to 10.2 billion in 2100. This estimate could be offset by three population-control measures: (1) lower the rate of unwanted births, (2) lower the desired family size, and (3) raise the average age at which women begin to bear children or reduce the number of births below the level that would replace current human populations (e.g., one child per woman).

      Insects and plants that live for a single year and reproduce once before dying are examples of organisms whose growth is geometric. In these species a population grows as a series of increasingly steep steps rather than as a smooth curve.

Logistic population growth

      The geometric or exponential growth of all populations is eventually curtailed by limitation of food, competition for other resources, disease, or some other ecological factor. If growth is limited by resources such as food, the exponential growth of the population begins to slow as competition for those resources increases. The growth of the population eventually slows nearly to zero as the population reaches the carrying capacity (K) for the environment. The result is an S-shaped curve of population growth known as the logistic curve. It is determined by the equation:

Population fluctuation

 As stated above, populations rarely grow smoothly up to the carrying capacity and then remain there. Instead, fluctuations in population numbers are the norm. In a few species, such as snowshoe hares, lemmings, lynx, and Arctic foxes, populations show regular cycles of increase and decrease spanning a number of years (Figure 2—>). The causes of these fluctuations are still under debate by population ecologists, and no single cause may provide an explanation for every species. Most major hypotheses link regular fluctuations in population size to factors that are dependent on the density of the population, such as the availability of food.

Factors affecting population fluctuation

      Population ecologists commonly divide the factors that regulate the size of populations into density-dependent and density-independent factors. Density-independent factors, such as weather and climate, affect the same proportion of individuals in a population regardless of population density. In contrast, the effects of density-dependent factors intensify as the population increases in size. For example, some diseases spread faster in dense populations than in sparse populations. Similarly, competition for food and other resources rises with density and affects an increasing proportion of the population. The dynamics of most populations are influenced by both density-dependent and density-independent factors, and the relative effects of the factors vary among populations.

Population cycles

      Because many factors influence population size, erratic variations in number are more common than regular cycles of fluctuation. Some populations undergo unpredictable irruptions in numbers, sometimes temporarily increasing by 10 or 100 times over a few years, only to follow with a similarly rapid crash. Migratory grasshoppers in the arid parts of Africa multiply to such a level that their numbers can blacken the sky overhead; similar upsurges occurred in North America before the 20th century. Some forest insects, such as the gypsy moths that were introduced to North America, are highly irruptive. As with species that fluctuate more regularly, the causes behind population irruptions are not fully known and are unlikely to have a single explanation that applies to all species.

      The size of other populations varies within tighter limits. Some fluctuate close to their carrying capacity, others fluctuate below this level, held in check by various ecological factors, including predators and parasites. The tremendous expansion of many populations of weeds and pests (pest) that have been released into new environments in which their enemies are absent suggests that predators, grazers, and parasites all contribute to maintaining the small sizes of many populations. To control the explosive proliferation of these species, biological control programs have been instituted. With varying degrees of success, parasites or pathogens inimical to the foreign species have been introduced into the environment. The European rabbit (Oryctolagus cuniculus) was introduced into Australia in the 1800s, and its population grew unchecked, wreaking havoc on agricultural and pasture lands. The myxoma virus subsequently was released among the rabbit populations and greatly reduced them (see below The coevolutionary “arms race” versus reduced antagonism (community ecology)). Populations of the prickly pear cactus (Opuntia) in Australia and Africa grew unbounded until the moth borer (Cactoblastis cactorum) was introduced (). Many other similar attempts at biological control have failed, illustrating the difficulty in pinpointing the factors involved in population regulation.

Metapopulations

      Although the dynamics and evolution of a single closed population are governed by its life history, populations of many species are not completely isolated and are connected by the movement of individuals (immigration and emigration). Consequently, the dynamics and evolution of many populations are determined by both the population's life history and the patterns of movement of individuals between populations. Regional groups of interconnected populations are called metapopulations. These metapopulations are, in turn, connected to one another over broader geographic ranges. The mapped distribution of the perennial herb Clematis fremontii variety Riehlii in Missouri shows the metapopulation structure for this plant over an area of 1,129 square kilometres (436 square miles; Figure 4). There is, therefore, a hierarchy of population structure from local populations to metapopulations to broader geographic groups of populations and eventually up to the worldwide collection of populations that constitute a species.

      As local populations within a metapopulation fluctuate in size, they become vulnerable to extinction during periods when their numbers are low. Extinction of local populations is common in some species, and the regional persistence of such species is dependent on the existence of a metapopulation. Hence, elimination of much of the metapopulation structure of some species can increase the chance of regional extinction of species.

      The structure of metapopulations varies among species. In some species one population may be particularly stable over time and act as the source of recruits into other, less stable populations. For example, populations of the checkerspot butterfly (Euphydryas editha) in California have a metapopulation structure consisting of a number of small satellite populations that surround a large source population on which they rely for new recruits. The satellite populations are too small and fluctuate too much to maintain themselves indefinitely. Elimination of the source population from this metapopulation would probably result in the eventual extinction of the smaller satellite populations.

      In other species, metapopulations may have a shifting source. Any one local population may temporarily be the stable source population that provides recruits to the more unstable surrounding populations. As conditions change, the source population may become unstable, as when disease increases locally or the physical environment deteriorates. Meanwhile, conditions in another population that had previously been unstable might improve, allowing this population to provide recruits.

      Overall, the population ecology and dynamics of all species is a complex result of their genetic structure, the life histories of the individuals, fluctuations in the carrying capacity of the environment, the relative influences of all the different kinds of density-dependent and density-independent factors that limit population growth, the spatial distribution of individuals, and the pattern of movement between populations that determines metapopulation structure. It is therefore not surprising that there are often great fluctuations in the numbers of individuals in local populations and that the long-term persistence of species may often require the conservation of many, rather than a few, populations.

John N. Thompson

Additional Reading

Eric R. Pianka, Evolutionary Ecology, 5th ed. (1994), provides a clear treatment of the basics of population ecology. Peter W. Price, Insect Ecology, 2nd ed. (1984), is a comprehensive text on the ecology of the most diverse group of organisms on Earth. Two books that examine the dynamics of plant populations and the evolution and ecology of plant life histories are Michael J. Crawley (ed.), Plant Ecology (1986); and Jonathan W. Silvertown and Jonathan Lovett Doust, Introduction to Plant Population Biology, 3rd ed. (1993). Michael Begon and Martin Mortimer, Population Ecology: A Unified Study of Animals and Plants (1986), examines the range of ecological and genetic factors that influence the structure, growth, and dynamics of populations. Thorough treatments of the theory of how genetic variation arises within populations and changes over time can be found in Daniel L. Hartl and Andrew G. Clark, Principles of Population Genetics, 2nd ed. (1989); and John Maynard Smith, Evolutionary Genetics (1989). One of the most detailed studies ever made of the population ecology and genetics of a vertebrate species is reported in B. Rosemary Grant and Peter R. Grant, Evolutionary Dynamics of a Natural Population: The Large Cactus Finch of the Galápagos (1989). Leslie A. Real (ed.), Ecological Genetics (1994), discusses the ways in which ecological conditions influence the genetic and spatial structure of populations and the life histories of organisms. Ian Newton (ed.), Lifetime Reproduction in Birds (1989), is a collection of long-term studies on the life histories and population ecology of birds. Richard B. Primack, Essentials of Conservation Biology (1993); and Michael E. Soulé (ed.), Conservation Biology: The Science of Scarcity and Diversity (1986), apply the principles of population genetics, population ecology, and evolutionary biology to the conservation of populations.J.A. Drake et al. (eds.), Biological Invasions: A Global Perspective (1989), contains articles on the growth and dynamics of populations of species introduced into areas in which they are not native. Paul R. Ehrlich and Anne H. Ehrlich, The Population Explosion (1990), examines the problems of overpopulation and also offers solutions. Michael Gilpin and Ilkka Hanski (eds.), Metapopulation Dynamics: Empirical and Theoretical Investigations (1991), explores the importance of metapopulation structure to population dynamics and to interactions between species.John N. Thompson

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