Introduction to Population Ecology

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.
 Fig. 1. Cycles in the
population dynamics of the snowshoe hare and its predator the Canadian lynx
(redrawn from MacLulich 1937). Note that percent mortality is an elusive measure,
it may, or may not, be useful since mortality varies with environment and time.
Fig. 2A. Logistic growth (Nt = Nt-1 + Rm(1-Nt-1/K)Nt-1). N = equal population
density at a given time (t). This so-called "logistic equation" was first proposed by
the mathematician Verhulst (1839). In ecology texts this equation is more often
written as DN/dt = rm(1-N/K)N, where D is density at any given time (t). K =
carrying capacity of environment. Rm = loge(Rm + 1).

Royal N. Chapman, Univ. Minnesota, in the 1930s proposed the concept of a balance
between biotic potential and environmental resistance. Chapman`s model was a
mathematical representation of the Malthusian concept, illustrated here by the
logistic growth of a laboratory population of yeast cells. Population growth
trajectory (N1 = N0 + (Rm -sN0)N0 ); Rm = maximum rate of increase, here = 1, s =
interaction coefficient, here = 0,0001, and carrying capacity of environment = 1000.
 Fig. 2B. Population growth
(N1 = N0 + RN0), where N1 = 10 and R=0.5 (blue), R=0 (black), and R=-0.5 (red).

Human populations represent another example of exponential growth. Magnitude of

the problems posed by human population growth can be seen from the fact that it
took more than 1 million years for the human population to first reach 200,000 (the
current daily rate of increase (See: US Census, Historical Estimates of World
Population). The human population is estimated to have first reached 1 billion
persons in 1830, and 2 billion in 1930, a doubling time of 100 years. In 1960, thirty
years later, the population edged past 3 billion, and a mere 15 years later, 4 billion.
In 1986, we exceeded 5 billion for the first time. Despite a slowing of the growth rate,
it is expected that the human population will exceed 6 billion in early 1999 (see:
Univ. North Carolina, Chapel Hill's World Population Counter) . To feed this
population, only as well as we presently do, it will be necessary to increase food
production 20% over the next 10-15 years.

Table 1. UN estimates of historical human

populations and projections for 21st century
(UNDP Press Release, 10/28/1998)

1 billion
1804
2 billion
1927
3 billion
1960
4 billion
1974
5 billion
1987
6 billion
1999
7 billion
2013
8 billion
2028
9 billion
2054
(range of estimates: 10.4-17
2100
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 Lotka-
Volterra 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).
 Fig 4A (above). Steady-state
population model (N1 = N0 + Rm(1 -sN0 /K)N0, where Rm = 2, K = 1000, and intitial
displacement from equilibrium x = -10.

Fig. 4B. Steady-state population model

(N1 = N0 + Rm(1 -sN0 /K)N0 ), when K = 1000, Rm = 3 and initial displacement
from equilibrium x = -10.

Fig. 4C (bottom). Steady-state population model

(N1 = N0 + Rm(1 -sN0 /K)N0 ) , when K = 1000, Rm = 1.5 and initial displacement
from equilibrium x = -10.
A. J. Nicholson (Australian entomologist) was a leading proponent of concept of
density-dependent mortality factors. He maintained that density-dependent
mortality played the key role in regulating prey populations. This is the essence of
the so-called "Balance of Nature" theory. This theory implied a static balance about
a mean (characteristic) equilibrium density with reciprocal (feedback) oscillations in
density about these means.

Nicholson and V. A. Bailey (1935) proposed a population model that incorporated a

"lag effect". This is particularly appropriate to parasitoids were population effects
of attack (oviposition) may not be evident until the parasitoid has completed its
immature development and emerges as a adult (killing the host).

Leading proponents of this view of population dynamics included the early

California biological control workers. The theory and practice of biological control
can be said to revolve about this assumption.

However, a contrary view of the nature of population regulating mortality factors

was argued by others, especially W. R. Thompson (Dominion Parasite Laboratory).
The Canadian workers held that assumptions of Nicholson and like thinkers were
unrealistic, and did not occur in nature, i.e., that the regulating role of a so- called
density-dependent mortality factors was largely myth. These workers argued that it
was unnecessary to postulate such a mechanism of population regulation. They
observed that the environment never remains continually favorable or unfavorable
for any species. If it did so that population would inevitably become either infinite or
decline to extinction. They maintained it was more accurate to say that populations
were (in reality) always in a state of "dynamic equilibrium" with their environment.

H. G. Andrewartha and L. C. Birch were leading proponents of the concept that

populations could be, and often are, regulated by abiotic factors.

These arguments became heated in the mid-1950s and early 1960s. Much of the
confusion and controversy regarding mechanisms of population regulation stemmed
from an inadequate data base, but an even more confining limitation was the
essential impracticality of making the extremely complex calculations required to
manipulate such data. Accordingly rapid advances in theoretical ecology, especially
in the area of population ecology, only occurred with the introduction of high speed
computers and the refinement of statistical theory.
Much of early population theory was developed deductively from controlled
laboratory experiments or fragmented field observations. This resulted in a
tendency to oversimplification and the development of models often not reflecting
biological reality.

One of the most useful starting points for a population ecologist is the development
of a life table. A life table is a schedule of mortality for each cohort (age group) of
individuals in the population. The methods were developed originally for actuarial
or demographic studies. Multifactor studies which incorporated the life table
technique were once largely the province of forest entomologists, but are now widely
used in agriculture.

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 density-

independent 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
categories include: mutualism (++), commensualism (+0), predator/prey (+-),
competition (- -), and amensalism (rare) (-0).

Insects and plants of two types: 1) good colonizers, e.g., weed species, with high
reproductive potential (capacity), adaptable, invaders, readily dispersed, "r-
strategists"; 2) good competitors, high survival, tend to stable population (K =
equilibrium), exploit stable environments, win out in competition, "K-strategists"
(R. H. MacArthur and E. O. Wilson 1967).

Most crop pests are r-strategists, e.g., aphids and phytophagous insects in general.
Natural enemies, i.e., parasites and predators, are mostly K strategists. This is said
to be one reason for the high failure rate associated with introductions of exotic
natural enemies.

Most crop plants are early succession plants, i.e., they are weedy species and
accordingly they also are r-strategists. The r-selected species are particularly suited
to exploiting the ecological patchiness and instability of the agroecosystem.

Fig. 5A + 5B. Island

Biogeography:
(MacArthur and
Wilson 1967)
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.
 Fig. 6 A + 6B. Carl B. Huffaker
et al. 1968 graphically presented four postulated types of functional (behavioral)
responses of predators including entomophagous insects to prey density. (A) number
killed, (B) percent killed. After Huffaker et al. 1968.
Nicholson curve: number of attacks determined by searching capacity (i.e., density-
dependent). 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.

Fig. 8A (above). A population trajectory on its reproduction plane showing growth

over three time increments (N0 to N3) in a consistent envrionment and also
following environmental degradation (broken line) at the end of the second time
period. The critical density (Nc, here shown as red line) is where the population
utilizes resources at the same rate that they are renewed (replaced), for a given
degree of environmental favorability.
Fig. 8B (above). A population trajectory on its reproduction plane showing growth
over three time increments (N0 to N3) in a consistent environment and also
following environmental degradation (broken line) at the end of the second time
period.

Fig. 9A-C. Predator/prey interactions (figures redrawn from Berryman, 1981)

Fig. 9A. Reproduction plane for a prey

species (A), where Ka is the carrying capacity in the absence of predation, Wb is the
marginal cost of predation, Pb is the predator density which drives the prey to
extinction.
 Fig 9B. Reproduction plane for a
predatorspecies (B), where Pa is the minimal prey density needed to sustain a
predator population, and Qa is the marginal benefit of the prey.

Fig 9C. Reproduction plane for a prey

species (A) and superimposed with a particular dynamic tragectory. An infinite
number of curves could be drawn and the relationships need not be straight lines.
The stability of the relationship would depend upon the characteristics of the
curves.
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:

1. Natality (birthrate) is low at low population densities because of problems
associated with low densities, e.g., finding mates, increases as population increases,
peaks and finally declines as intraspecific competition increases, e.g., competition
for food.

2. Predation increases with increasing host (herbivore) density, then declines as host
populations overwhelm (and escape) their regulating influence. This occurs because
the characteristic pattern is for overall predator response to be sigmoid, based on a)
functional response of the individual predator and b) numerical response of the
population.

3. Intraspecific competition increases with increased population (prey) density. This

produces additional mortality as natality shows a down-turn. Other density-
dependent mortality and stress also comes into play producing a marked increase in
combined mortality.

The net result of these interacting factors is that changes in the population are
subjected to dramatic changes in cumulative mortality rates. These are illustrated in
the following figures.

Fig. 10. Generalized relationship between

population density of a herbivore species, natality, mortality caused by enemies
(with a peak at moderatly low low population density) and intraspecific competition
(with a peak at relatively high population density) (after Southwood 1975).
 Fig. 11. Generalized relationship between
herbivore population density in one generation and herbivore population density in
the next generation. The line at 45 degrees is the line of no growth, i.e., population
density stable from one generation to the next. Points above the line indicate
population growth, those below the line population decline. X = extinction point, S =
stable equilibrium point, R = point of release from natural enemies, O = oscillations
equilibrium point, and C = crash point.

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 r-
selected habitats, while natural enemies tend to dominance in more stable
ecosystems.
Common feature of epidemics (epizootics) is that outward migration (emigration)
occurs advancing in waves. Knowledge of spatial distributions and population
dynamics can be used to manage populations over large areas.

Simulation games have been used to study and test ecological theory. One such
model is the so-called "Game of Life". This is a game invented by a mathematician
to illustrate (mimic) environmental feedback loops, a number of individuals
(checkers) are positioned at random on board. Rules are that any individual
adjacent to one or more neighbors dies of isolation and any adjacent to four or more
individuals dies of overcrowding (two negative feedbacks) and that whenever an
empty square exists adjacent to exactly three individuals, a new individual is born (a
parody of real-life).

The population dynamics of pests occurring over large areas can be investigated by
gridding the area and monitoring population trends, studies of dispersal (migration)
reproduction, competition, weather, environmental conditions, etc.

Tactics and Strategies:

Pest management is as applied science with no unique principles. Focus of all pest
management programs is to "erode the homeostatic capability" ("homeostasis") of
pest populations i.e., to reduce the equilibrium position of populations (K) spatially
and temporally so that the frequency and duration of fluctuation above economic
thresholds are reduced or eliminated.

The tactics (ecological manipulations) and strategies (pest management decision-

making processes) distinguish pest management (IPM) from unilateral approaches.

Pest management is the selection, integration, and implementation of pest control

strategies on the basis of predicted economic, ecological, and sociological
consequences (Rabb 1972).

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

Populus 3.4, Simulations of Population Biology by Don Alstad, University of
Minnesota. The Populus software contains a set of simulation models used in
teaching population biology and evolutionary ecology at the University of
Minnesota. The program is distributed without charge. (To download software visit
http://www.cbs.umn.edu/populus/installer.html)

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.
In previous chapters/units we have concentrated on the biology of the
individual cell, tissue, and organism. There are levels of organization above
the individual organism that will be the subject of this unit. Individual
organisms are grouped into populations, which in turn form communities,
which form ecosystems. Ecosystems make up the biosphere, which includes
all life on Earth. If there is life on other planets, will we need another level
of organization?

Biosphere: The sum of all living things taken in conjunction with their
environment. In essence, where life occurs, from the upper reaches of the
atmosphere to the top few meters of soil, to the bottoms of the oceans. We
divide the earth into atmosphere (air), lithosphere (earth), hydrosphere
(water), and biosphere (life).

Ecosystem: The relationships of a smaller groups of organisms with each

other and their environment. Scientists often speak of the interrelatedness
of living things. Since, according to Darwin's theory, organisms adapt to
their environment, they must also adapt to other organisms in that
environment. We can discuss the flow of energy through an ecosystem
from photosynthetic autotrophs to herbivores to carnivores.

Community: The relationships between groups of different species. For

example, the desert communities consist of rabbits, coyotes, snakes, birds,
mice and such plants as sahuaro cactus (Carnegia gigantea), Ocotillo,
creosote bush, etc. Community structure can be disturbed by such things
as fire, human activity, and over-population.

Species: Groups of similar individuals who tend to mate and produce

viable, fertile offspring. We often find species described not by their
reproduction (a biological species) but rather by their form (anatomical or
form species).

Populations: Groups of similar individuals who tend to mate with each

other in a limited geographic area. This can be as simple as a field of
flowers, which is separated from another field by a hill or other area where
none of these flowers occur.

Individuals: One or more cells characterized by a unique arrangement of

DNA "information". These can be unicellular or multicellular. The
multicellular individual exhibits specialization of cell types and division of
labor into tissues, organs, and organ systems.
Organ System: (in multicellular organisms). A group of cells, tissues, and
organs that perform a specific major function. For example: the
cardiovascular system functions in circulation of blood.

Organ: (in multicellular organisms). A group of cells or tissues performing

an overall function. For example: the heart is an organ that pumps blood
within the cardiovascular system.

Tissue: (in multicellular organisms). A group of cells performing a specific

function. For example heart muscle tissue is found in the heart and its
unique contraction properties aid the heart's functioning as a pump. .

Cell: The fundamental unit of living things. Each cell has some sort of
hereditary material (either DNA or more rarely RNA), energy acquiring
chemicals, structures, etc. Living things, by definition, must have the
metabolic chemicals plus a nucleic acid hereditary information molecule.

Organelle: A subunit of a cell, an organelle is involved in a specific

subcellular function, for example the ribosome (the site of protein
synthesis) or mitochondrion (the site of ATP generation in eukaryotes).

Molecules, atoms, and subatomic particles: The fundamental functional

levels of biochemistry.
Organization levels of life, in a graphic format. 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.

It is thus possible to study biology at many levels, from collections of

organisms (communities), to the inner workings of a cell (organelle).

Ecology is the study how organisms interact with each other and their
physical environment. These interactions are often quite complex. Human
activity frequently disturbs living systems and affects these interactions.
Ecological predictions are, of a consequence, often more general than we
would like.

Population Growth | Back to Top

A population is a group of individuals of the same species living in the

same geographic area. The study of factors that affect growth, stability,
and decline of populations is population dynamics. All populations
undergo three distinct phases of their life cycle:

1. growth
2. stability
3. decline
Population growth occurs when available resources exceed the number of
individuals able to exploit them. Reproduction is rapid, and death rates are
low, producing a net increase in the population size.

Population stability is often proceeded by a "crash" since the growing

population eventually outstrips its available resources. Stability is usually
the longest phase of a population's life cycle.

Decline is the decrease in the number of individuals in a population, and

eventually leads to population extinction.

Factors Influencing Population Growth

Nearly all populations will tend to grow exponentially as long as there are
resources available. Most populations have the potential to expand at an
exponential rate, since reproduction is generally a multiplicative process.
Two of the most basic factors that affect the rate of population growth are
the birth rate, and the death rate. The intrinsic rate of increase is the birth
rate minus the death rate.

Two modes of population growth. The Exponential curve (also known as a

J-curve) occurs when there is no limit to population size. The Logistic
curve (also known as an S-curve) shows the effect of a limiting factor (in
this case the carrying capacity of the environment). 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.
Population Growth Potential Is Related to Life History

The age within it's individual life cycle at which an organism reproduces
affects the rate of population increase. Life history refers to the age of
sexual maturity, age of death, and other events in that individual's lifetime
that influence reproductive traits. Some organisms grow fast, reproduce
quickly, and have abundant offspring each reproductive cycle. Other
organisms grow slowly, reproduce at a late age, and have few offspring per
cycle. Most organisms are intermediate to these two extremes.
Population curves. a) three hypothetical populations (labelled I, II, and
III); b, c, and d) three real populations. Note that the real curves
approximate one of the three hypotheticals. 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.

Age structure refers to the relative proportion of individuals in each age

group of a population. Populations with more individuals aged at or before
reproductive age have a pyramid-shaped age structure graph, and can
expand rapidly as the young mature and breed. Stable populations have
relatively the same numbers in each of the age classes.

Comparison of the population age structuire in the United States and

Mexico. Note the deographic bulge in the Mexican population. The effects
of this buldge will be felt for generations. 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.
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 populations are in a growth phase. Since evolving about 200,000

years ago, our species has proliferated and spread over the Earth.
Beginning in 1650, the slow population increases of our species
exponentially increased. New technologies for hunting and farming have
enabled this expansion. It took 1800 years to reach a total population of 1
billion, but only 130 years to reach 2 billion, and a mere 45 years to reach 4
billion.

Despite technological advances, factors influencing population growth will

eventually limit expansion of human population. These will involve
limitation of physical and biological resources as world population
increased to over six billion in 1999. The 1987 population was estimated at
a puny 5 billion.
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.

Populations Transition Between Growth and Stability

Limits on population growth can include food supply, space, and complex
interactions with other physical and biological factors (including other
species). After an initial period of exponential growth, a population will
encounter a limiting factor that will cause the exponential growth to stop.
The population enters a slower growth phase and may eventually stabilize
at a fairly constant population size within some range of fluctuation. This
model fits the logistic growth model. The carrying capacity is the point
where population size levels off.
Relationship between carrying capacity (K) and the population density
over time. 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.

Several Basic Controls Govern Population Size

The environment is the ultimate cause of population stabilization. Two

categories of factors are commonly used: physical environment and
biological environment. Three subdivisions of the biological environment
are competition, predation, and symbiosis.

Physical environment factors include food, shelter, water supply, space

availability, and (for plants) soil and light. One of these factors may
severely limit population size, even if the others are not as constrained. The
Law of the Minimum states that population growth is limited by the
resource in the shortest supply.

The biological role played by a species in the environment is called a niche.

Organisms/populations in competition have a niche overlap of a scarce
resource for which they compete. Competitive exclusion occurs between
two species when competition is so intense that one species completely
eliminates the second species from an area. In nature this is rather rare.
While owls and foxes may compete for a common food source, there are
alternate sources of food available. Niche overlap is said to be minimal.
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.
Graphs showing competition between two species of Paramecium. Since
each population alone prospers (yop two graphs), when they are in a
competition situation one species will win, the other will lose (bottom
graph). 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..

Predators kill and consume other organisms. Carnivores prey on animals,

herbivores consume plants. Predators usually limit the prey population,
although in extreme cases they can drive the prey to extinction. There are
three major reasons why predators rarely kill and eat all the prey:

1. Prey species often evolve protective mechanisms such as camouflage,

poisons, spines, or large size to deter predation.
2. Prey species often have refuges where the predators cannot reach
them.
3. Often the predator will switch its prey as the prey species becomes
lower in abundance: prey switching.
Fluctuations in predator (wolf) and prey (moose) populations over a 40-
year 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.

Symbiosis has come to include all species interactions besides predation

and competition. Mutualism is a symbiosis where both parties benefit, for
example algae (zooxanthellae) inside reef-building coral. Parasitism is a
symbiosis where one species benefits while harming the other. Parasites act
more slowly than predators and often do not kill their host. Commensalism
is a symbiosis where one species benefits and the other is neither harmed
nor gains a benefit: Spanish moss on trees, barnacles on crab shells.
Amensalism is a symbiosis where members of one population inhibit the
growth of another while being unaffected themselves.

The Real World Has a Complex Interaction of Population Controls

Natural populations are not governed by a single control, but rather have
the combined effects of many controls simultaneously playing roles in
determining population size. If two beetle species interact in the
laboratory, one result occurs; if a third species is introduced, a different
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:

 population growth as previous limits are removed

 population decline as new limits are imposed

Agriculture and animal domestication are examples of population increase

of favored organisms. In England alone more than 300,000 cats are put to
sleep per year, yet before their domestication, the wild cat ancestors were
rare and probably occupied only a small area in the Middle East.
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-to-
pesticide 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.

Altering Population Growth | Back to Top

Humans can remove or alter the constraints on population sizes, with both
good and bad consequences. On the negative side, about 17% of the 1500
introduced insect species require the use of pesticides to control them. For
example, African killer bees are expanding their population and migrating
from northward from South America. These killer bees are much more
agressive than the natives, and destroy native honeybee populations.

On a positive note, human-induced population explosions can provide

needed resources for growing human populations. Agriculture now
produces more food per acre, allowing and sustaining increased human
population size.

Human action is causing the extinction of species at thousands of times the

natural rate. Extinction is caused by alteration of a population's
environment in a harmful way. Habitat disruption is the disturbance of the
physical environment of a species, for example cutting a forest or draining
wetlands. Habitat disruption in currently the leading cause of extinction.

Changes in the biological environment occur in three ways.

1. Species introduction: An exotic species is introduced into an area

where it may have no predfators to control its population size, or
where it can gratly out compete native organisms. Examples include
zebra mussels introduced into Lake Erie, and lake trout released
into Yellowstone Lake where they are threatening the native
cutthroat trout populations.
 2. Overhunting: When a predator population increases or becomes
more efficient at killing the prey, the prey population may decline or
go extinct. Examples today include big game hunting, which has in
many places reduced the predator (or in this case prey) population.
In human prehistory we may have caused the extinction of the
mammoths and mastodons due to increased human hunting skill.
3. Secondary extinction: Loss of food species can cause migration or
extinction of any species that depends largely or solely on that
species as a food source.

Overkill is the shooting, trapping, or hunting of a species usually for sport

or economic reasons. Unfortunately, this cannot eliminate "pest" species
like cockroaches and mice due to their large population sizes and capacity
to reproduce more rapidly than we can eliminate them. However, many
large animals have been eliminated or had their populations drastically
reduced (such as tigers, elephants, and leopards).

The death of one species or population can cause the decline or elimination
of others, a process known as secondary extinction. Destruction of bamboo
forests in China, the food for the giant panda, may cause the extinction of
the panda. The extinction of the dodo bird has caused the Calviera tree to
become unable to reproduce since the dodo ate the fruit and processed the
seeds of that tree.

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.

Populations Have a Minimum Viable Size

Even if a number of individuals survive, the population size may become

too small for the species to continue. Small populations may have breeding
problems. They are susceptible to random environmental fluctuations and
genetic drift to a greater degree than are larger populations. The chance of
extinction increases exponentially with decreasing population size.

The minimum viable population (MVP) is the smallest population size that
can avoid extinction by the two reasons listed above. If no severe
environmental fluxes develop for a long enough time, a small population
will recover. The MVP depends heavily on reproductive rates of the
species.

Range and Density | Back to Top

Populations tend to have a maximum density near the center of their

geographic range. Geographic range is the total area occupied by the
species. Outlying zones, where conditions are less optimal, include the zone
of physiological stress (where individuals are rare), and eventually the zone
of intolerance (where individuals are not found).

The environment is usually never uniform enough to support uniform

distribution of a species. Species thus have a dispersion pattern. Three
patterns found include uniform, clumped, and random.

Geographic ranges of species are dynamic, over time they can contract or
expand due to environmental change or human activity. Often a species
will require another species' presence, for example Drosophila in Hawaii.
Species ranges can also expand due to human actions: brown trout are now
found worldwide because of the sprerad of trout fishing.
How to Determine Population Density
Studying population growth and density offers interesting insights for the
geographic region. For instance, a population-density report may yield demographic
shifts within that region. Comparing and contrasting different lengths of time, such
as the previous decade compared to the current population density, can show the
increases or decreases in residents. The population-density calculation also helps
governments and social scientists know where the most people live in a region,
giving weight to the cultural or political influence of a city or region. To find the
population density of any given region, a researcher needs only a few variables to
make an accurate calculation. Variables depend on the unit of measurement,
however, as some countries utilize miles or kilometers to measure land space.
Researchers need to be cognizant of such measurement differences.

Related Searches:
 Ecosystem Ecology
 Population Data

Difficulty:
Easy

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.

o 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
 Appalachian Mountains and then subtract the number of people who
do not live in the Appalachian Mountains.

o 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.

o 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.