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Background
Thomas Malthus
The first significant contribution to the theory of population ecology was that of
Thomas Malthus, an English clergyman, who in 1798 published his Essay on the
Principle of Population. Malthus introduced the concept that at some point in time
an expanding population must exceed supply of prerequisite natural resources, i.e.,
population increases exponentially resulting in increasing competition for means of
subsistence, food, shelter, etc. This concept has been termed the "Struggle for
Existence".
Charles Darwin
Malthus's theories profoundly influenced Charles Darwin 1859, On the Origin of
Species, e.g., the concept of "Survival of the Fittest". Mortality of this type can be
termed "facultative mortality" (as opposed to catastrophic mortality, e.g., weather,
insecticides).
Harry Smith, pioneering biological control worker with the University of California
(1935), proposed the equivalent and now accepted terms density-dependent and
density-independent. Density-dependent mortality factors are those that are
facultative in effect, density-independent mortality factors are those that are
catastrophic in effect.
A density-dependent mortality factor is one that causes a varying degree of
mortality in subject population,and that the degree of mortality caused in a function
(i.e., related) to the density of the subject (affected) population (density-geared,
feedback regulation, self-regulating or self-limiting) may and typically involves a lag
effect., e.g., most biological control agents.
1 billion
2 billion
3 billion
4 billion
5 billion
6 billion
7 billion
8 billion
9 billion
(range of estimates: 10.4-17
billion)
Exercise: Compute annual population growth rates for each of the time perioids
above:
Equation for population growth model is X=X0ert where the original population is
X0 mathematical constant: natural log (e = 2.718), represents Malthusian
parameter. Population will increase in size to X, over time (t), if rate of increase (r) is
positive. If population not growing, i.e., r = 0.0, then rt = 0.0, and e0.0= l.0, X = X0.
When r has a value greater than 0 population will increase, e.g., rt = 0.693 (e0.693 =
2.0), population will double in time t (X = 2X0). The doubling time of this population
will be t = 0.693/r. If the population is decreasing, r is a negative value. The function
of X = X0ert can be made a straight line by the natural log transformation, i.e., lnX =
lnX0 + rt (the equation for a straight line). Characteristics of this line are: an
intercept at lnX0 and a slope of r. Linear regression analysis can be applied (since
this is a straight line), with the independent variable being time. The linear equation
thus describes the rate of population growth (+) or decline (-). Algebraic
rearrangement of this equation permits one to solve for the rate of population
increase.
In the 1920s, A. J. Lotka (1925) and V. Volterra (1926) devised mathematical models
representing host/prey interaction. This was the first attempt to mathematically
represent a population model as achieving a cyclic balance in mean mean
(characteristic) population density, i.e., to attain a dynamic equilibrium. The LotkaVolterra curve assumes that prey destruction is a function not only of natural enemy
numbers, but also of prey density, i.e., related to the chance of encounter.
Populations of prey and predator were predicted to flucuate in a regular manner
(Volterra termed this "the law of periodic cycle"). Lotka-Volterra model is an
oversimplification of reality, as these curves are derived from infinitesimal calculus
when in nature association is seldom continuous over time (life cycles being finite).
The final test of any population model is its usefulness to predict (generation to
generation) changes in abundance or to explain why changes occur at particular
population densities.
Consequences of recent advances in understanding of population dynamics has lead
to almost universal (among ecologists) acceptance of the proposition that population
growth is geared to population density.
Differences in the relative importance of density-dependent and densityindependent mortality factors varies in different environments, e.g., the role of
biotic components tends to be greater in more stable (benign) environments.
Competitive processes/cooperative processes: we often think of interactions between
individuals even within species as only negative, but interactions can be positive
(even between species), e.g., defense against predators, genetic diversity (concept of
minimum density), mate finding (sustainable population), early mortality may favor
subsequent survival.
Interactions between species can be very complex even when only 2 species are
considered (through impact on environment of other species). Each species can
affect environment of other positively (+), negatively (-), or have no effect (0). Major
Equilibrium models for near and distant islands. Equilibrium (in number of species
present) occurs where curves of rates of immigration and rates of extinction
intersect. I is the initial rate of immigration and P is the total number in the species
pool on the mainland.
The parallel between agroecosystems and island defaunation is obvious. Many
factors affect the various population processes: number of potential invaders,
distance of source of invaders, conditions for invasion and settling, attractiveness
(favorability) of crop.
Effects of various regulating factors on the population dynamics of the host tend to
differ: predators, harmonic (cyclic), parasitoids, intrusive, pathogens, disruptive,
food, catastrophic (W. J. Turnock and J. A. Muldrew 1971).
Numerical response is the key to the success of entomophagous insects; occurs in the
following ways:
1. Concentration, not different from sigmoid curve of functional response.
2. Immediate numerical response, increased survival
3. Delayed numerical response, increased reproduction.
Nicholson curve: number of attacks determined by searching capacity (i.e., densitydependent). C. S. Holling (1965) disk curve: characteristic of invertebrate predators.
Sigmoid curve: characteristic of vertebrate predators. Thompson curve: number of
attacks limited by capacity for consumption or production of eggs, but not by
searching capacity or host density.
Experimental data suggests that Holling's disc curve is characteristic of the
predatory behavior of most entomophagous insects, i.e., the number of prey
attacked increases but at a proportionately slower rate as prey density increases.
Interactions between species, especially between predators and prey are of great
theoretical and practical interest to ecologists.
Environmental feedbacks: Populations degrade their environment, utilizing its
resources. If the environment is highly favorable, resources are not limiting, i.e., the
resources are replaced or recover faster than they are utilized. This favors rapid
(exponential) growth of the population. As the rate of utilization of resources
approaches the rate of replacement, the rate of population increase slows. Finally,
population overshots carrying capacity of environment (resources depleted faster
than can be renewed). This results in severe competition and the population
collapses. When the population drops below the replacement value the environment
recovers.
Fig. 7 (above). A reproduction plane divided into zones of population growth (R > 0)
and decline (R < 0) by the equilibrium diagonal line (R = 0). The density of the
population at equilibrium (K) changes in direction to the favorability of the
environment (F). The equilibrum line is further divided into 3 sections with different
stability properties: 1) a lower section where sK <1 providing asymptomic stability, a
midsection where 1 < sK < 2 providing dampened stability, and an upper section
sK>2 which is unstable.
Interactions (reproduction planes) between predator and prey. There must be some
prey to sustain the predator. Pb is the predator density which drives prey to
extinction. Ka is the carrying capacity in the absence of the predator. Wb is the
marginal cost of a given level of predation. Pa is the minimum prey density required
to sustain the predator population (and avoid extinction). Qa is the marginal benefit
of prey. The superimposed reproduction planes show equilibrium lines for predator,
Eb, and prey, Ea, and a particular dynamic trajectory.
We can draw an infinite variety of these curves, relationships not necessarily
straight line, stability will depend on characteristic of curves (assuming these
represent real life situation).
Community stability: A central tenet of classical ecology is that complex
communities tend to be more stable (largely based on observation) theory recently
challenged, argument that simple systems may be less subject to external
disturbances. What has emerged is that:
1. Competitive interactions between species lead to instability unless dominated by
negative feedbacks within self-loops, i.e., one species setting in motion cyclic
oscillations in numbers of another.
2. The number of competing species increase the competitive interactions must be
proportionately weaker or instability will result.
3. Interactions between tropic levels (predator/prey) tend to stabilize populations
(community).
4. community stability is frequently interrupted by severe environmental
disruptions, leading to a series of successional communities gradually evolving to
climax associations. Frequent or continual disruptions may lead to persistent
nonclimax community. Herbivores play an important role in plant succession by
tending to harvest unthrifty members of a plant community, e.g., overmature trees.
They also recycle nutrients and increase productivity and vigor of community, mutualistic?
Population dynamics/epidemiology theory is a vast and formidable subject.
However, the synoptic model developed by T. R. E. Southwood (1975 and
subsequent papers) is most instructive, and summarizes much of the theoretical
concepts and empirical bases for contemporary model building.
The model has 3 salient features:
Fig. 13 (above). Synoptic model of population growth (after Southwood and Comins
1976, J. Anim. Ecol. 45: 949-965). The synoptic model of Southwood demonstrates
the link between habitat stability (natural ecosystems evolving toward a K-selected
type, agroecosystems representing an r-selected type) and relative favorability of
each for pests and natural control agents. Pests having a relative advantage in rselected habitats, while natural enemies tend to dominance in more stable
ecosystems.
Ecology References
Websites
Population Ecology Reference List, Site maintained by Alexei Sharov, Virginia
Tech.
See on-line course: Quantitative Population Ecology "This site is an absolute must
for all teaching Population Ecology or students learning it, but it should also not be
missed by anybody else." (Plant Pathology Internet Guide Book).
Also visit: Modelling Forest Insect Dynamics
Print
Liss, W. J., L. J. Gut, P. H. Westigard, and C. E. Warren. 1986. Perspectives on
arthropod community structure, organization, and development in agricultural
crops. Annual Review of Entomology 31: 455-478.
Schowalter, T. D., W. W. Hargrove, and D. A. Crossley, Jr. 1986. Herbivory in
forested ecosystems. Annual Review of Entomology 31: 177-196.
Finklestein, L. and E. R. Carson. 1985. Mathematical Modelling of Dynamic
Biological Systems. John Wiley & Sons. 355pp.
Taylor, L. R. 1984. Assessing and interpreting the spatial distributions of insect
populations. Annual Review of Entomology 29: 321-57.
Huffaker, C. B. and R. L. Rabb, editors. 1984. Ecological Entomology. John Wiley &
Sons. 844 pp.
Price, P. W., C. M. Slobdchikoff, and W. S. Gaud, editors. 1984. A New Ecology:
Novel Approaches to Interactive Systems. John Wiley & Sons. 515 pp.
Price, P. W. 1984. Insect Ecology. John Wiley & Sons. 600 pp.
Odum, H. T. 1983. Systems Ecology: an Introduction. John Wiley & Sons. 644 pp.
Stinner, R. E. C. S. Barfield, J. L. Stimac, and L. Dohse. 1983. Dispersal and
movement of insect pests. Annual Review of Entomology 28: 319-335.
Berryman, A. A. 1981. Population Systems: A General Introduction. Plenum Press.
Levins, R. and M. Wilson. 1980. Ecological theory and pest management. Annual
Review of Entomology 25: 287-308.
Furtick, W. R. 1976. Implementing pest management programs: an international
perspective. In: Integrated Pest Management, J. L. Apple and R. F. Smith, editors,
pp. 17-27. Plenum.
Apple, J. L. and R. F. Smith. 1976. Progress, problems, and prospects for integrated
pest management. In: Integrated Pest Management, J. L. Apple and R. F. Smith,
editors, pp. 179-97. Plenum Press.
Huffaker, C. B., F. J. Simmons, and J. E. Laing. 1976. The theoretical and empiracal
basis of biological control. In: Theory and Practice of Biological Control, C. B.
Huffaker and P. S. Messenger, editors, Chapter 3, pp. 41-78. Academic Press.
Price, P. W. and G. P. Waldbauer. 1975. Ecological Aspects of Pest Management. In :
Introduction to Insect Pest Management, R. L. Metcalf and W. Luckmann, editors,
pp. 37-73. Wiley-Interscience.
Dempster, J. P. 1975. Animal Population Ecology. Academic Press.
Pielou, E. C. 1975. Ecological Diversity. John Wiley & Sons, Inc.
Rabb, R. L., R. E. Stinner, and G. A. Carlson. 1974. Ecological principles as a basis
for pest management in the agroecosystem. In: Proceedings of the Summer Institute
on Biological Control of Plant Insects and Diseases, F. G. Maxwell and F. A. Harris,
editors, pp. 19-45. Univ. Press of Mississippi.
Sailer, R. I. 1971. Invertebrate predators. In: Toward Integrated Control, pp. 32-44.
USDA Forest Research Paper NE-194.
Turnock, W. J. and J. A. Muldrew. 1971. Parasites. In: Toward Integrated Control,
pp. 59-87. USDA Forest Research Paper NE-194.
Huffaker, C. B., P. S. Messenger, and P. DeBach. 197l. The natural enemy
component in natural control and the theory of biological control. In: Biological
Control, C. B. Huffaker, editors, pp. 16- 67. Plenum.
Southwood,T. R. E. and M. J. Way. 1970. Ecological Background to Pest
Management. In: Concepts of Pest Management, pp. 6-29. R. L. Rabb and F. E.
Guthrie, editors. N. Carolina State Univ.
Varley, G. C. and G. R. Gradwell. 1970. Recent advances in insect population
dynamics. Annual Review of Entomology 15: 1-24.
Harcourt, D. G. 1969. The development and use of life tables in the study of natural
insect populations. Annual Review of Entomology 14: 175-191.
Huffaker, C. B. and P. S. Messenger. 1964. Population cology- historical
development. In: Biological Control of Insect Pests and Weeds. P. DeBach, editors,
Chapter 3, pp. 45-73. Reinhold.
Huffaker, C. B. and P. S. Messenger. 1964. The concept and significance of natural
control. In: Biological Control of Insect Pests and Weeds, P. DeBach, editor, Chapter
4, pp. 74-117. Reinhold.
The Baby Boomers and Gen X. As the population bulge, the baby Boomers
born after World War II, aged and began to have children of their own this
created a secondary bulge termed Generation X. What happens when the
Generation X members begin to have their own children? Image from
Purves et al., Life: The Science of Biology, 4th Edition, by Sinauer Associates
(www.sinauer.com) and WH Freeman (www.whfreeman.com), used with permission.
Human population growth over the past 10,000 years. Note the effects of
worldwide disease (the Black death) and technological advances on the
populatiuon size. Images from Purves et al., Life: The Science of Biology, 4th
Edition, by Sinauer Associates (www.sinauer.com) and WH Freeman
(www.whfreeman.com), used with permission.
Paramecium aurelia has a population nearly twice as large when it does not
have to share its food source with a competing species. Competitive release
occurs when the competing species is no longer present and its constraint
on the winner's population size is removed.
Fluctuations in predator (wolf) and prey (moose) populations over a 40year span. Note the effects of declines in the wolf population in the late
1960s and again in the early 1980s on the moose population. Image from
Purves et al., Life: The Science of Biology, 4th Edition, by Sinauer Associates
(www.sinauer.com) and WH Freeman (www.whfreeman.com), used with permission.
outcome develops. The latter situation is more like nature, and changes in
one population may have a domino effect on others.
Which factors, if either, is more important in controlling population
growth: physical or biological? Physical factors may play a dominant role,
and are called density independent regulation, since population density is
not a factor The other extreme has biological factors dominant, and is
referred to as density dependent regulation, since population density is a
factor. It seems likely that one or the other extreme may dominate in some
environments, with most environments having a combination control.
Population Decline and Extinction
Extinction is the elimination of all individuals in a group. Local extinction
is the loss of all individuals in a population. Species extinction occurs when
all members of a species and its component populations go extinct.
Scientists estimate that 99% of all species that ever existed are now extinct.
The ultimate cause of decline and extinction is environmental change.
Changes in one of the physical factors of the environment may cause the
decline and extinction; likewise the fossil record indicates that some
extinctions are caused by migration of a competitor.
Dramatic declines in human population happen periodically in response to
an infectious disease. Bubonic plague infections killed half of Europe's
population between 1346 and 1350, later plagues until 1700 killed one
quarter of the European populace. Smallpox and other diseases decimated
indigenous populations in North and South America.
Human Impact
Human populations have continued to increase, due to use of technology
that has disrupted natural populations. Destabilization of populations
leads to possible outcomes:
Pollution
Pollutants generally are (unplanned?) releases of substances into the air
and water. Many lakes often have nitrogen and phosphorous as limiting
nutrients for aquatic and terrestrial plants. Runoff from agricultural
fertilizers increases these nutrients, leading to runaway plant growth, or
eutrophication. Increased plant populations eventually lead to increased
bacterial populations that reduce oxygen levels in the water, causing fish
and other organisms to suffocate.
Pesticides and Competition
Removal of a competing species can cause the ecological release of a
population explosion in that species competitor. Pesticides sprayed on
wheat fields often result in a secondary pest outbreak as more-tolerant-topesticide species expand once less tolerant competitors are removed.
Removal of Predators
Predator release is common where humans hunt, trap, or otherwise reduce
predator populations, allowing the prey population to increase.
Elimination of wolves and panthers have led to increase in their natural
prey: deer. There are more deer estimated in the United States than there
were when Europeans arrived. Large deer populations often cause over
grazing that in turn leads to starvation of the deer.
Introduction of New Species
Introduction of exotic or alien non-native species into new areas is perhaps
the greatest single factor to affect natural populations. More than 1500
exotic insect species and more than 25 families of alien fish have been
introduced into North America; in excess of 3000 plant species have also
been introduced. The majority of accidental introductions may fail,
however, once an introduced species becomes established, its population
growth is explosive. Kudzu, a plant introduced to the American south from
Japan, has taken over large areas of the countryside.
Kudzu covering a building (left) and closeup of the flowers and leaves
(right). Images from http://www.alltel.net/~janthony/kudzu/, photographs by Jack
Anthony, used with permission.
Giant pandas eat an estimated 10,000 pounds of bamboo per panda per
year. Image of a giant panda eating bamboo from
http://www.bonus.com/contour/Save_our_Earth/http@@/library.thinkquest.org/298
8/e-animals.htm#Giant Panda.
Instructions
Things You'll Need
Census Data
1.
o
1
Properly define a region. Since a researcher needs exact numbers, a
researcher has to have the correct definition of a region. Does the
calculation include two or more states or is it covering a land mass
region, such as the Appalachian Mountains? Questions such as these
need to be determined before any calculation can begin.
2
Record the population of a region. Governments and research
organizations measure the population of states, regions, or an entire
country in annual reports. A researcher needs the most recent
population number for an accurate density report. If the region is not
defined, say for the Appalachian Mountains example, the researcher
needs to add the populations of all the states that lie within the
3
Calculate or find the area of the region. The area is defined by the
square miles or kilometers that define the region. Most governments
define the area of a country, state, or municipality. If the measured
region includes two or more countries or states, the researcher needs
to add up the total areas of each state.
4
Divide the area of the region by the population. The researcher's
formula should look something like this: population density =
population of region/area of region. The quotient after the calculation
is the population density.