Population Ecology: Chapter Concepts

687
Population Ecology
Chapter Concepts
33.1 Scope of Ecology
Ecology is the study of the interactions of
organisms with each other and the physical
environment. 688
The scope of ecology encompasses the
individual, the population, the community, the
ecosystem, and the biosphere. 688
33.2 Characteristics of Populations
Population size depends upon births, deaths,
immigration, and emigration. 690
Two patterns of population growth (exponential
and logistic) have been developed. 690
Mortality within a population is often illustrated
by a survivorship curve. 691
33.3 Regulation of Population Size
Factors that affect population size are classied
as density-independent and density-dependent.
692
33.4 Life History Patterns
Life history patterns range from one in which
many young receive little care to one in which
few young receive much care. 694
33.5 Human Population Growth
The human population is still growing
exponentially, and how long this can continue is
not known. 695
These poppies are members of a population whose size is
determined by the carrying capacity of the environment.
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how the shes feed (Fig. 33.1). Most organisms do not exist
singly; rather, they are part of a population, a functional unit
that interacts with the environment. Apopulation is dened
as all the organisms within an area belonging to the same
species. At this level of study, ecologists are interested in fac-
tors that affect the growth and regulation of population size.
Acommunity consists of all the various populations in-
teracting at a locale. In a coral reef, there are numerous pop-
ulations of shes, crustaceans, corals, and so forth. At this
level ecologists want to know how interactions like preda-
tion and competition affect the organization of a community.
An ecosystem contains a community of populations and
also the abiotic (nonliving) environment. Energy ow and
chemical cycling are signicant aspects of understanding
how an ecosystem functions. The biosphere is that portion
of the entire earths surface where living things exist.
Modern ecology is not just descriptive, it also develops
hypotheses that can be tested. Acentral goal of modern ecol-
ogy is to develop models which explain and predict the dis-
tribution and abundance of organisms. Ultimately, ecology
considers not one particular area, but the distribution and
abundance of populations in the biosphere.
Ecology is the study of the interactions of
organisms with other organisms and with the
physical environment. These interactions determine
the distribution and abundance of organisms at a
particular locale and over the earths surface.
Ecology is the study of the interactions of organisms with
each other and with the physical environment. Ecology, like
so many biological disciplines, is wide-ranging. At one of its
lowest levels, ecologists study how the individual organism
is adapted to its environment. For example, they study why
shes in a coral reef live only in warm tropical waters and
688 Part 7 Behavior and Ecology 33-2
I
magine a watering hole that can accommodate 100 ze-
bras. If, at rst, there are only two zebras, and each pair
of zebras produces only four zebras, how many more
generations could there be without overtaxing the water
hole? Youre correct if you say four and incorrect if you say
ve. The problem is that you cant just consider the newly ar-
rived zebrasyou have to add the number of zebras already
there.
2 4 8 16 32 64 128
Also, notice that when its time to stop, there are only 62 ze-
bras (30 32). Thats one of the unusual things about pop-
ulation growthat one point it seems as if there is plenty of
room, and then all of a sudden, theres not enough room.
Modern ecologists now recognize that knowledge of
population growth has almost unlimited application possibil-
ities, including the management of wildlife to prevent extinc-
tion and the maintenance of food (organic nutrients) sources
for the human population.
Organism Population Community Ecosystem
Figure 33.1 Ecological levels.
The study of ecology encompasses various levels, from the individual organism to the population, community, and ecosystem.
Density and Distribution of Populations
Population density is the number of individuals per unit area
or volume. If we calculated the density of the human popu-
lation, we would know how many individuals there are per
square mile, for example. From this we might get the im-
pression that humans are uniformly distributed, but we
know full well that most people live in cities. Even within a
city more people live in particular neighborhoods than oth-
ers. Population distribution is the pattern of dispersal of indi-
viduals within the area of interest.
There are three patterns of distribution: uniform, ran-
dom, and clumped (Fig. 33.2). Human beings have the
clumped pattern of distribution, which is the most common
pattern. Today, ecologists want to discover what causes the
spatial distribution of organisms. With reference to human
beings, we know that many cities sprung up at the junction
of rivers or near inlets that make good harbors for ships. In a
study of the distribution of hard clams in a bay on the south
shore of Long Island, New York, it was found, as discussed
on page 693, that clams are apt to occur where the sediment
contains oyster shells. Hopefully, this information can be
used to transform areas of low abundance to areas of high
abundance of clams.
As with clams, the distribution of organisms can be due
to abiotic factors. Physical factors like a particular inorganic
nutrient (the oyster shells provide calcium carbonate for the
formation of clam shells) can determine where organisms
occur. Also important are precipitation and temperature,
which can be limiting factors for the distribution of an or-
ganism. Limiting factors are those factors that particularly
determine whether an organism lives in an area. Trout live
only in cool mountain streams where there is a high oxygen
content, but carp and catsh are found in rivers near the
coast because they can tolerate warm waters, which have a
low concentration of oxygen. The timberline is the limit of
tree growth in mountainous regions or in high latitudes.
Trees cannot grow above the high timberline because of low
temperature and the fact that water remains frozen most of
the year.
The distribution of organisms can also be due to biotic
(living) factors. In Australia, the red kangaroo does not live
outside inland areas because it is adapted to feeding on the
arid grasses that grow there. And there are more humans
where the soil is suitable for growing crops than where the
soil is rocky and poor in inorganic nutrients.
Ecology as a science includes a study of the
distribution of organisms: where and why
organisms are located in a particular place at a
particular time.
Chapter 33 Population Ecology 689 33-3
a.
b.
c.
food
heat
fuel
materials waste
Figure 33.2 Patterns of distribution within a population.
Members of a population may be distributed uniformly, randomly, or
usually in clumps. a. Golden eagle pair distribution is uniform over a
suitable habitat area due to the territoriality of the birds. b. The
distribution of female moose with calves is random over a suitable
habitat. c. Human beings tend to be clumped in cities where many
people take up residence. Cities take resources from and send their
waste to surrounding regions.
Populations have a particular pattern of growth and sur-
vivorship among other possible characteristics.
Patterns of Population Growth
Populations have a certain size and the size can stay the
same from year to year, increase, or decrease, according to a
per capita rate of increase. Suppose, for example, a human
population presently has a size of 1,000 individuals, and the
birthrate is 30 per year, and the death rate is 10 per year. The
per capita rate of increase per year will be:
= 0.02 = 2.0% per year
(Notice that our per capita rate of increase disregarded either
immigration or emigration, which for the purpose of our dis-
30 10
1,000
cussion can be assumed to be equivalent.) The highest possi-
ble per capita rate of increase for a population is called its bi-
otic potential (Fig. 33.3). Whether the biotic potential is high
or low depends on such factors as the following:
1. usual number of offspring per reproduction
2. chances of survival until age of reproduction
3. how often each individual reproduces
4. age at which reproduction begins.
Suppose we are studying the growth of a population of
insects that are capable of infesting and taking over an area.
Under these circumstances exponential growth is expected.
An exponential pattern of population growth results in a J-
shaped curve (Fig. 33.4a). This pattern of population growth
can be likened to compound interest at the bank: as your
money increases, the more interest you will get. If the insect
population has 2,000 individuals and the per capita rate of
increase is 20% per month, then there will be 2,400 insects af-
ter one month, 2,880 after two months, and 3,456 after three
months, and so forth.
Notice that a J-shaped curve has these phases:
lag phase: during this phase, growth is slow because the
population is small.
exponential growth phase: during this phase, growth is
accelerating and the population is exhibiting its
biotic potential.
Usually, exponential growth cannot continue for long
because of environmental resistance. Environmental resis-
tance is all those environmental conditions such as a limited
supply of food, an accumulation of waste products, in-
creased competition, or predation that prevent populations
from achieving their biotic potential. Due to environmental
resistance, growth levels off and a pattern of population
growth called logistic growth is expected. Logistic growth
results in an S-shaped growth curve (Fig. 33.4b).
Figure 33.3 Biotic potential.
Animal husbandry relies on biotic potential. If a single female pig has
her rst litter at nine months, and produces two litters a year, each of
which contain an average of four females (which in turn reproduced at
the same rate), there would be 2,220 pigs by the end of three years.
Time
N
u
m
b
e
r

o
f

O
r
g
a
n
i
s
m
s
a. Time
N
u
m
b
e
r

o
f

O
r
g
a
n
i
s
m
s
b.
lag
exponential
growth
lag
exponential
growth
deceleration
stable equilibrium
environmental
resistance
carrying
capacity
Figure 33.4 Patterns of population growth.
a. Exponential growth results in a J-shaped growth curve because the growth rate is positive. b. Logistic growth results in an S-shaped growth
curve because environmental resistance causes the population size to level off and be in a steady state.
Notice that an S-shaped curve has these phases:
lag phase: during this phase, growth is slow because the
population is small.
exponential growth phase: during this phase, growth is
accelerating due to biotic potential.
deceleration phase: during this phase, the rate of
population growth slows down.
stable equilibrium phase: during this phase, there is little
if any growth because births and deaths are about
equal.
The stable equilibrium phase is said to occur at the carrying
capacity of the environment. The carrying capacity is the
number of individuals the environment can normally
support.
Our knowledge of logistic growth has practical impli-
cations. The model predicts that exponential growth will
occur only when population size is much lower than the
carrying capacity. So, as a practical matter, if we are using a
sh population as a continuous food source, it would be
best to maintain the population size in the exponential
phase of growth. Biotic potential is having its full effect
and the birth rate is the highest it can be during this phase.
If we oversh, the population will sink into the lag phase,
and it will be years before exponential growth recurs. On
the other hand, if we are trying to limit the growth of a
pest, it is best to reduce the carrying capacity rather than
reduce the population size. Reducing the population size
only encourages exponential growth to begin once again.
Farmers can reduce the carrying capacity for a pest by al-
ternating rows of different crops rather than growing one
type of crop per the entire eld.
Exponential growth produces a J-shaped curve
because population growth accelerates over time.
Logistic growth produces an S-shaped curve
because the population size stabilizes when the
carrying capacity of the environment has been
reached.
Survivorship
Population growth patterns assume that populations are
made up of identical individuals. Actually, the individuals
are in different stages of their life span. Let us consider how
many members of an original group of individuals born at
the same time, called a cohort, are still alive after certain in-
tervals of time. If we plot the number surviving, a survivor-
ship curve is produced.
For the sake of discussion, three types of idealized sur-
vivorship curves are recognized (Fig. 33.5a). The type I
curve is characteristic of a population like humans in which
most individuals survive well past the midpoint, and death
does not come until near the end of the life span. On the
other hand, the type III curve would be typical for a popula-
tion of oysters in which most individuals die very young. In
the type II curve, survivorship decreases at a constant rate
throughout the life span. This has been found typical of a
population of song birds.
Sometimes populations do not t any of these curves ex-
actly. For example, in a cohort of Poa annua plants, most in-
dividuals survive till six to nine months, and then the
chances of survivorship diminish at an increasing rate.
There is much that can be learned about the life history
of a species by studying its survivorship curve. Would you
predict that most or few members of a population with a
type III survivorship curve are contributing offspring to the
next generation? Obviously since death comes early for
most members, only a few are living long enough to repro-
duce. What about the other two types of survivorship
curves?
Populations have a pattern of survivorship that
becomes apparent from studying the survivorship
curve of a cohort.
N
u
m
b
e
r

o
f

S
u
r
v
i
v
o
r
s
Percent of Life Span
1,000
100
10
0 50 100
I
II
III
human
Hydra
oyster
Figure 33.5 Survivorship curves.
Human beings have a type I survivorship curve: the individual usually
lives a normal life span and then death is increasingly expected.
Hydras have a type II curve in which chances of surviving are the
same for any particular age. Oysters have a type III curve: most
deaths occur during the free-swimming larva stage, but those that
survive to adulthood usually live a normal life span.
Ecologists want to determine when and if the two patterns of
population growth just discussed occur in nature. They want
to know if the factors which regulate population growth are al-
ways extrinsic (environmental) or whether there are also in-
trinsic (based on the anatomy, physiology, or behavior of the
organism) factors which regulate population growth.
In one study, 4 male and 21 female reindeer (Rangifer)
were released on St. Paul Island in the Bering Sea off Alaska
in 1911. St. Paul Island had a completely undisturbed envi-
ronment. The reindeer fed on lichens, which grow slowly
and cannot recover quickly from grazing. Normally, rein-
deer migrate seasonally, giving their food source a chance to
regrow, but these reindeer were conned to an island. Also,
there was little hunting pressure, and there were no preda-
tors. The herd grew exponentially to about 2,000 reindeer;
overgrazed the habitat; and then abruptly declined to only 8
animals in 1950. Such a precipitous drop is called a popula-
tion crash (Fig. 33.6). In this instance it does seem as if envi-
ronmental factors, such as food source and possibly disease,
are regulating population size.
For some time ecologists have recognized that the envi-
ronment contains both abiotic and biotic components. They
suggested that abiotic factors like weather and natural disas-
ters were density-independent. By this they meant that the
number of organisms present did not inuence the effect of
the factor. Fires dont necessarily kill a larger percentage of
individuals as the population increases in size (Fig. 33.7). On
the other hand, biotic factors like parasitism, competition,
and predation were designated as density-dependent. Con-
sider, for example, an area in which there are only 100 holes
for crabs to hide in. The greater the number of crabs beyond
100, the better the chance a shorebird will nd one and eat it.
In a population study of the great tit, Parus major, it was
found that the population size uctuates above and below
the carrying capacity. While density-dependent and density-
independent factors are involved, the researchers believe
that territoriality, an intrinsic factor, also plays a role. Terri-
toriality is apparent when members of a population are
spaced out more than would be expected from a random oc-
cupation of the area.
Density-independent and density-dependent
factors can often explain the population dynamics
of natural populations. Both types of factors are
extrinsic to the organism; perhaps intrinsic factors
like territoriality also play a role.
1910 1920 1930 1940 1950
0
500
1,000
1,500
2,000
N
u
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b
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o
f

R
e
i
n
d
e
e
r
b.
a.
decline as a result
of sudden resource
depletion
exponential
growth
Figure 33.6 Density-dependent effect.
a. A reindeer, Rangifer. b. On St. Paul Island, Alaska, reindeer grew
exponentially for several seasons and then underwent a sharp
decline as a result of overgrazing the available range.
Figure 33.7 Density-independent effect.
A re can start and rage out of control regardless of how many
organisms are present.
The hard clam is a commercially harvested bivalve mollusk that
is found throughout the Great South Bay located on the south
shore of Long Island. The clam is eaten raw on the half shell,
and is also baked. Hard clams are a necessary ingredient for
New England and Manhattan clam chowder. The Great South
Bay has often been referred to as a hard clam factory because
in the 1970s, over half of the hard clams harvested in the United
States came from its waters. For the past 20 years, Jeffrey Kass-
ner (Fig. 33A) has been studying the distribution of hard clams
in the eastern third of the bay for the Town of Brookhaven.
Knowing the distribution and abundance of hard clams, as
well as the responsible environmental factors of this distribu-
tion is important to Brookhaven because the hard clam industry
has an annual value of $10 million and employs 300 shermen.
Of particular interest to Brookhaven is the possibility of using
this information to develop projects, such as utilizing aquacul-
ture technology, that will increase hard clam abundance.
Kassner rst took a census of the hard clam population. For
several weeks during the summer, a barge-mounted crane with
a one square meter clamshell bucket was used to take bottom
grabs at 232 stations located throughout the study area. Each
bottom sample was placed in a one square meter wire sieve and
washed with a high pressure water hose to separate the hard
clams from the sediment so that the hard clams could be
counted and measured in order to calculate various demo-
graphic parameters of the hard clam population. The eldwork
was physically hard, and by the end of the day, the crew of stu-
dents and biologists were tired, wet, and covered with mud. The
results, however, have proven to be well worth the effort.
Working with Dr. Robert Cerrato of the
State University of New York, Kassner has
drawn a composite census map showing
the distribution of and abundance of the
clams. He was surprised to nd that hard
clams are not distributed uniformly
throughout the study area, but occur in dis-
tinct patches of high and low abundance. A
dense assemblage of clams is traditionally
referred to as a clam bed and six such
beds were identied.
Kassner and students found that nearly
all of the beds coincided with areas of high
shell content sediment associated with
relict oyster reefs. This observation is of
historical note as well as biological interest
because up until the early years of this cen-
tury, the Great South Bay was a major pro-
ducer of oysters and was the source of the
world famous Blue Point Oyster. Although oysters are no
longer found in the Great South Bay because of environmental
changes, they left behind a legacy in the sediment that now sup-
ports high abundances of hard clams.
The knowledge that hard clam abundance is positively asso-
ciated with relict oyster reefs might make it possible to trans-
form low-abundance areas into high-abundance areas. Kassner
felt he needed a complete sedimentary portrait of high- and
low-abundance areas. To develop one he borrowed techniques
generally associated with other marine science disciplines: from
shipwreck hunting, he used a side-scan sonar to map the topog-
raphy of the bottom; from deep-sea research, he used a ROV
(Remotely Operated Vehicle) to photograph the bottom; from
commercial shing, a fathometer to map the bottom; and from
pollution studies, a sediment prole camera to photograph the
sediment-water interface where hard clams live and feed.
Kassner, with the help of students, is now in the process of
putting all this different information together on a single map.
Because it will take time and perhaps more studies to develop
the portrait, he is concurrently exploring the feasibility of hav-
ing shells placed on low-abundance areas in order to create new
relict oyster reefs. If this strategy works, it would be a tremen-
dous boon to the shellsh industry because nearly three-
quarters of the study area is low abundance.
Kassner feels his work is not only scientically interesting, it
is also personally rewarding. Over the years, he has become
friends with many shermen and knows that if he is successful,
he will be helping them to continue an occupation that in some
cases goes back generations.
693
Distribution of Hard Clams in the Great South Bay
Figure 33A
Jeffrey Kassner, with the help of students, is preparing a sedimentary portrait of high
and low clam abundance areas in Great South Bay, Long Island, New York. This
information will be used to increase the yield of clams for local shermen.
We have already had an opportunity to point out that popu-
lations vary on such particulars as their rate of growth and
life span. Such particulars are a part of a species life history
pattern, which is based on genetically determined variations
that are subject to natural selection.
The logistic growth model has been used to suggest that
one possible pattern is an opportunistic pattern and the other
is an equilibrium pattern (Fig. 33.8). These are also called
r-selected and K-selected because in mathematical formulas
for population growth, r represents the per capita rate of in-
crease and K represents the carrying capacity. Members of
opportunistic populations are small in size, mature early,
and have a short life span. They tend to produce many rela-
tively small offspring and to forego parental care in favor of
a greater number of offspring. The more offspring, the more
likely some of them will survive a population crash. Because
of their short life span and ability to disperse to new locales,
density-dependent mechanisms such as predation and com-
petition are unlikely to play a major role in regulating popu-
lation size and growth rates. Classic examples of such
opportunistic species are many insects and weeds.
In contrast, we know there are populations whose size
remains pretty much at the carrying capacity. Resources
such as food and shelter are relatively scarce for these indi-
viduals, and those who are best able to compete will have
the largest number of offspring. These organisms allocate
energy to their own growth and survival and to the growth
and survival of their offspring. Therefore they are fairly
large, are slow to mature, and have a fairly long life span.
They are specialists rather than colonizers and tend to be-
come extinct when their normal way of life is destroyed. The
best possible examples of equilibrium species are found
among birds and mammals. The Florida panther is the
largest of the animals in the Florida Everglades, requires a
very large range, and produces few offspring, which must
be cared for. Currently, the Florida panther is unable to com-
pensate for a reduction in its range, and is therefore on the
verge of extinction.
Nature is actually more complex than these two possible
life history patterns and most populations lie somewhere in
between these two extremes. For example a cod is a rather
large sh weighing 1025 pounds and measuring up to
3 feet in lengthbut the cod releases gametes in vast num-
bers, the zygotes form in the sea, and the parents make no
further investment in developing offspring. Of the 6 to 7 mil-
lion eggs released by a single female cod, only a few will be-
come adult sh.
Differences in the environment result in different
selection pressures and a range of life history
characteristics.
Figure 33.8 Life history patterns.
Dandelions are an opportunistic species with the characteristics
noted, and bears are equilibrium species with the characteristics
noted. Often the distinctions between these two possible life
history patterns are not as clear cut as they may seem.
Opportunistic Pattern
Small individuals
Short life span
Fast to mature
Many offspring
Little or no care of offspring
Equilibrium Pattern
Large individuals
Long life span
Slow to mature
Few offspring
Much care of offspring
The human population has an exponential pattern of growth
and a J-shaped growth curve (Fig. 33.9c). It is apparent from
the position of 1999 on the growth curve in Figure 33.9a that
growth is still quite rapid. The equivalent of a medium-sized
city (200,000) is added to the worlds population every day,
and 88 million (the equivalent of the combined populations
of the United Kingdom, Norway, Ireland, Iceland, Finland,
and Denmark) are added every year.
The present situation can be appreciated by considering
the doubling time. The doubling timethe length of time it
takes for the population size to doubleis now estimated to
be 47 years. Such an increase in population size will put ex-
treme demands on our ability to produce and distribute re-
sources. In 47 years, the world will need double the amount
of food, jobs, water, energy, and so on just to maintain the
present standard of living.
Many people are gravely concerned that the amount of
time needed to add each additional billion persons to the
world population has taken less and less time. The rst billion
didnt occur until 1800; the second billion arrived in 1930; the
third billion in 1960, and today there are nearly 6 billion. Only
if the per capita rate of increase declines can there be zero
population growth, when the birthrate equals the death rate
and population size remains steady. The worlds population
may level off at 8, 10.5, or 14.2 billion, depending on the speed
with which the per capita rate of increase declines.
More-Developed Versus Less-Developed
Countries
The countries of the world can be divided into two groups.
The more-developed countries (MDCs), typied by coun-
tries in North America and Europe, are those in which pop-
ulation growth is low and the people enjoy a good standard
of living (Fig. 33.9a). The less-developed countries (LDCs),
such as countries in Latin America, Africa, and Asia, are
those in which population growth is expanding rapidly and
the majority of people live in poverty (Fig. 33.9b). (Some-
times the term third-world countries is used to mean the
less-developed countries. This term was introduced by
those who thought of the United States and Europe as the
rst world and the former USSR as the second world.)
The more-developed countries (MDCs) doubled their
populations between 1850 and 1950. This was largely due to
a decline in the death rate, the development of modern med-
icine, and improved socioeconomic conditions. The decline
in the death rate was followed shortly thereafter by a decline
in the birthrate, so that populations in the MDCs experi-
enced only modest growth between 1950 and 1975. This se-
quence of events (i.e., decreased death rate followed by
decreased birthrate) is termed a demographic transition.
Yearly growth of the MDCs as a whole has now stabi-
lized at about 0.1%. The populations of a few of the MDCs
Italy, Denmark, Hungary, Swedenare not growing or are
actually decreasing in size. In contrast, there is no leveling
more-developed countries
less-developed
countries
1999
1750
0
2
4
6
8
10
12
1800 1850 1900 1950 2000 2050 2100 2150
P
o
p
u
l
a
t
i
o
n

(
i
n

b
i
l
l
i
o
n
s
)
Source: Population Reference Bureau.
b.
c.
Figure 33.9 World population growth.
People in the (a) more-developed countries have a high standard of living and will
contribute least, while people in the (b) less-developed countries have a low standard of
living and will contribute most to the world population growth. c. World population growth
to 1998 with estimates to 2150.
a.
off and no end in sight to U.S. population growth, as dis-
cussed in the reading on page 698. Although yearly growth
of the United States is only 0.6%, many people immigrate to
the United States each year. In addition, there was an un-
usually large number of babies born between 1947 and 1964
(called a baby boom). Therefore, a large number of women
are still of reproductive age.
Although the death rate began to decline steeply in
the LDCs following World War II with the importation of
modern medicine from the MDCs, the birthrate remained
high. The yearly growth of the LDCs peaked at 2.5% be-
tween 1960 and 1965. Since that time, a demographic tran-
sition has begun: the decline in the death rate has slowed
and the birthrate has fallen. Ayearly growth of 1.8% is ex-
pected by the end of the century. Still, because of expo-
nential growth, the population of the LDCs may explode
from 4.4 billion today to 10.2 billion in 2100. Most of this
growth will occur in Africa, Asia, and Latin America.
Ways to greatly reduce the expected increase have been
suggested:
1. Establish and/or strengthen family planning programs.
Adecline in growth is seen in countries with good
family planning programs supported by community
leaders. Currently, 25% of women in the sub-Saharan
Africa say they would like to delay or stop
childbearing, yet they are not practicing birth control;
likewise, 15% of women in Asia and Latin American
have an unmet need of birth control.
2. Use social progress to reduce the desire for large
families. Many couples in the LDCs presently desire as
many as four to six children. But providing education,
raising the status of women, and reducing child
mortality are desirable social improvements that could
cause them to think differently.
3. Delay the onset of childbearing. Adelay in the onset of
childbearing and wider spacing of births could cause a
temporary decline in the birthrate and reduce the
present reproductive rate.
Age Distributions
The age-structure diagrams of MDCs and LDCs in Figure
33.10 divide the population into three age groups: depen-
dency, reproductive, and postreproductive. The LDCs are
experiencing a population momentum because they have
more women entering the reproductive years than older
women leaving them.
Laypeople are sometimes under the impression that if
each couple has two children, zero population growth (no
increase in population size) will take place immediately.
However, replacement reproduction, as it is called, will still
cause most countries today to continue growing due to the
age structure of the population. If there are more young
women entering the reproductive years than there are older
women leaving them, then replacement reproduction will
still result in growth of the population.
Many MDCs have a stable age structure, but most
LDCs have a youthful prolea large proportion of the
population is younger than the age of 15. This means that
their populations will still expand greatly, even after
replacement reproduction is attained. The more quickly
replacement reproduction is achieved, however, the
sooner zero population growth will result.
Currently, the less-developed countries are expanding
dramatically because of exponential growth.
80+
7579
7074
6569
6064
5559
5054
4549
4044
3539
3034
2529
2024
1519
1014
59
04
80+
7579
7074
6569
6064
5559
5054
4549
4044
3539
3034
2529
2024
1519
1014
59
04
0
Males Females
Less Developed
More Developed
millions
postreproductive
postreproductive
reproductive
reproductive
dependency
a.
b.
300 300 250 250 200 200 150 150 100 100 50 50
Figure 33.10 Age-structure diagrams (1998).
The diagrams illustrate that (a) the MDCs are approaching
stabilization, whereas (b) the LCDs will expand rapidly due to their
age distributions. Source: United Nations Population Division, 1998.
A Sustainable World
While we are sometimes quick to realize that the
growing populations of the LDCs are putting a
strain on the environment, we should realize that
the excessive resource consumption of the MDCs
also stresses the environment. Environmental im-
pact is measured not only in terms of population
size, but in terms of resource consumption and the
pollution caused by each person in the population.
An average American family, in terms of per capita
resource consumption and waste production, is the
equivalent of 30 people in India (Fig. 33.11).
Before the industrial revolution, people felt
better connected to the plants and animals on
which they depended, and they were better able to
live in a sustainable way. After the industrial revo-
lution, we especially began to think of ourselves as
separate from nature and endowed with the right
to exploit nature as much as possible. But our in-
dustrial society lives on borrowed carrying capacity
our cities not only borrow resources from the
country, our entire population borrows from the
past and future. The forests of the Carboniferous
period have become the fossil fuels that sustain our way of
life today, and the environmental degradation we cause is
going to be paid for by our children.
Ecologists have two favorite sayings: (1) Everything is
connected to everything else, and (2) there is no free lunch.
We have seen that if you affect one part of the carbon cycle,
you affect the entire balance of carbon in the entire world.
Ecological effects know no boundaries. Coal that is burned
in the Midwest releases acids into the atmosphere that af-
fect lakes in the Northeast. And plants and animals arent
the only organisms affected. Humans are dependent on
natural cycles just as much as any organism in the bio-
sphere. What we do to natural ecosystems will eventually
be felt by us also. The second saying means that we have to
pay for what we do. If we build a home on a oodplain, we
can expect that it will be ooded once in a while. When we
burn fossil fuels, we can expect acid rain and global warm-
ing as a consequence. Many times it is difcult to predict
the particular consequences, but we can be assured that
eventually they will become apparent.
Overpopulation and overconsumption account for in-
creased pollution, and also for the present mass extinction
of wildlife. We are expected to lose one-third to two-thirds
of the earths species, any one of which could possibly
have made a significant contribution to agriculture or
medicine. It should never be said, What use is this organ-
ism? Aside from its contribution to the ecosystem in
which it lives, one never knows how a particular organism
might someday be useful to humans. Adult sea urchin
skeletons are now used as molds for the production of
small articial blood vessels, and armadillos are used in
leprosy research.
It is clearly time for a new philosophy. In a sustain-
able world, development will meet economic needs of all
peoples while protecting the environment for future gen-
erations. Various organizations have singled out commu-
nities to serve as models of how to balance ecological and
economic goals. For example, in Clinch Valley of south-
west Virginia, the Nature Conservancy is helping to revive
the traditional method of logging with draft horses. This
technique, which allows the selective cutting of trees, pre-
serves the forest and prevents soil erosion, which is so
damaging to the environment. The United Nations has an
established bioreserve system, a global network of sites
that combine preservation with research on sustainable
management for human welfare. More than 100 countries
are now participants in the program. However, sustain-
ability is more than likely incompatible with the kinds of
consumption/waste patterns currently practiced in devel-
oped countries.
All peoples can benet from a sustainable world
where economic development and environmental
preservation are considered complementary, rather
than opposing, processes.
Population
P
e
r
c
e
n
t
0
20
40
60
80
100
Hazardous
waste
production
Consumption
fossil fuels
other industrialized countries
Source: Natural Resources Defense Council, 1993.
developing countries
United States
metals paper
5
17
78
18
10
72
25
35
40
20 20
33
42
25
60
Figure 33.11 Resource consumption for MDCs and LDCs.
The populations of MDCs are smaller than LDCs. Yet, the MDCs produce most of
the hazardous wastes because their consumption of fossil fuels, metals, and
paper, for example, is much greater than the LDCs.
Summarizing the Concepts
Ecology is the study of the interactions of organisms with other organ-
isms and with the physical environment. Ecology encompasses several
levels of study: organism, population, community, ecosystem, and -
nally the biosphere. Population density is simply the number of indi-
viduals per unit area or volume. Distribution of these individuals can
be uniform, random, or clumped. Most members of a population are
clumped as are the members of a human population. Limiting factors
such as water, temperature, and availability of organic nutrients often
determine a populations distribution.
Future population size is dependent upon the per capita rate of increase.
The per capita rate of increase is calculated by subtracting the number of
deaths from the number of births and dividing by the number of indi-
viduals in the population. (Immigration and emigration are usually con-
sidered to be equal.) Every population has a biotic potential, the greatest
possible per capita rate of increase under ideal circumstances.
Two possible patterns of population growth are considered. Expo-
nential growth results in a J-shaped curve because as the population
increases in size so does the expected increase in new members. Most
environments restrict growth, and exponential growth cannot continue
indenitely. Under these circumstances logistic growth occurs and an
S-shaped growth curve results. When the population reaches carrying
capacity, the population stops growing because environmental resis-
tance opposes biotic potential.
Populations tend to have one of three types of survivorship
curves, depending on whether most individuals live out the normal
life span, die at a constant rate regardless of age, or die early.
Population growth is limited by density-independent (e.g., weather)
and density-dependent factors (predation, competition, and resource
availability). Do some populations have an intrinsic means of regulat-
ing population growth as opposed to density-independent and
density-dependent factors, which are extrinsic means? Territoriality is
given as an example of a possible intrinsic means of regulation.
The logistic growth model has been used to suggest that life history
patterns depend on natural selection and vary from those species that
are opportunists to those that are in equilibrium with the carrying ca-
pacity of the environment. Opportunistic species produce many young
within a short period of time and rely on rapid dispersal to new, unoc-
cupied environments. Population size is regulated by density-
independent factors. Equilibrium species produce a limited number of
young, which they nurture for a long time, and population size is reg-
ulated by density-dependent factors.
The human population is expanding exponentially, and it is unknown
when the population size will level off. Most of the expected increase
will occur in certain LDCs (less-developed countries) of Africa, Asia,
and Latin America. Support for family planning, human development,
and delayed childbearing could help prevent an expected increase.
T
he answer to how to curb the expected
increase in the worlds population lies
in discovering how to curb the rapid pop-
ulation growth of the less-developed
countries. In these countries, population
experts have discovered what they call the
the virtuous cycle. Family planning
leads to healthier women, and healthier
women have healthier children, and the
cycle continues. Women no longer have to
have many babies for a few to survive.
More education is also helpful because
better educated people are more interested
in postponing childbearing and promot-
ing womens rights. Women who have
equal rights with men tend to have fewer
children.
There isnt any place where women
have had the choice that they havent cho-
sen to have fewer children, says Beverly
Winikoff at the Population Council in New
York City. Governments dont need to
resort to force. Bangladesh is a case in
point. Bangladesh is one of the densest
and poorest countries in the world. In 1990
the birthrate was 4.9 children per woman
and now it is 3.3. This achievement was
due in part to the Dhaka-based Grameen
Bank, which loans small amounts of
money mostly to destitute women to start
a business. The bank discovered that when
women start making decisions about their
lives, they also start making decisions
about the size of their families. Family
planning within Grameen families is twice
as common as the national average; in fact,
those women who get a loan promise to
keep their families small! Also helpful has
been the network of village clinics that
counsel women who want to use contra-
ceptives. The expression contraceptives
are the best contraceptives refers to the
fact that you dont have to wait for social
changes to get people to use contracep-
tivesthe two feed back on each other.
Unfortunately, some of the less-
developed countries faced with economic
crisis have cut back on their family plan-
ning programs, and the more-developed
countries have not taken up the slack. In-
deed, some foreign donors have also cut
back on aidthe U.S. by one-third.
Questions
1. Do you think less-developed countries
should simply make contraception
available, or should more persuasive
methods be employed? Explain.
2. Do you think that more-developed
countries should be concerned about
population growth in the less-developed
countries? Why or why not?
3. Are you in favor of foreign aid to help
countries develop family planning
programs? Why or why not?
Go To Student OLC
4. An S-shaped growth curve
a. occurs when there is no environmental resistance.
b. includes an exponential growth phase.
c. occurs if survivorship is short-lived.
d. occurs in natural populations but not laboratory ones.
e. All of these are correct.
5. If a population has a type I survivorship curve (most live the
entire life span), which of these would you also expect?
a. a single reproductive event per adult
b. overlapping generations
c. sporadic reproductive events
d. reproduction occurring near the end of the life span
e. None of these are correct.
6. Apyramid-shaped age distribution means that the
a. prereproductive group is the largest group.
b. population will grow for some time in the future.
c. more young women are entering the reproductive years
than older women leaving theirs.
d. country is more likely an LDC rather than an MDC.
7. Which of these is a population-independent regulating factor?
a. competition d. weather
b. predation e. resource availability
c. size of population
8. Fluctuations in population growth can correlate to changes in
a. predation.
b. weather.
c. resource availability.
d. population regulating factors.
9. An equilibrium life history pattern includes all but
a. large individuals.
b. long life span.
c. individuals slow to mature.
d. few offspring.
e. little or no care of offspring.
10. The human population
a. is undergoing exponential growth.
b. is not subject to environmental resistance.
c. uctuates from year to year.
d. only grows if emigration occurs.
11. Label this S-shaped growth curve.
Time
N
u
m
b
e
r

o
f

O
r
g
a
n
i
s
m
s
a.
b.
c.
d. e.
stable equilibrium
Studying the Concepts
1. What are the various levels of ecological study? 688
2. What are three types of distribution patterns for a popula-
tion? Explain why the human population has a clumped
pattern. 689
3. How do you calculate the per capita rate of increase for a
population? What is biotic potential? 690
4. What type growth curve indicates that exponential growth is
occurring? What are the environmental conditions for expo-
nential growth? 690
5. What type growth curve indicates that biotic potential is
being opposed by environmental resistance? What
environmental conditions are involved in environmental
resistance? 690
6. What is the carrying capacity of an area? 691
7. Describe the three general types of survivorship
curves. 691
8. Give examples of extrinsic density-independent and
density-dependent factors that regulate population
size. 692
9. Give support to the belief that intrinsic factors might regulate
population size in some populations. 692
10. Name and give ve contrasting characteristics for the two
extreme life history patterns. 694
11. Why would you expect the life histories of natural popula-
tions to have a mixture of characteristics from these two
patterns? 694
12. What type of growth curve presently describes the popula-
tion growth of the human population? 695
13. Distinguish between MDCs and LDCs. Include a reference to
age-structure diagrams. 69596
14. Explain why the population of LDCs is expected to increase
tremendously. What steps could be taken to prevent this from
occurring? 696
Testing Yourself
Choose the best answer for each question.
1. Which of these levels of ecological study involves both abiotic
and biotic components?
a. organisms d. ecosystem
b. populations e. All of these are correct.
c. communities
2. When phosphorus is made available to an aquatic commu-
nity, the algal populations suddenly bloom. This indicates
that phosphorus is
a. a density-dependent regulating factor.
b. gaseous.
c. a reproductive factor.
d. a limiting factor.
3. AJ-shaped growth curve should be associated with
a. exponential growth.
b. biotic potential.
c. no environmental resistance.
d. high per capita rate of increase.
Understanding the Terms
age-structure diagram 696
biosphere 688
biotic potential 690
carrying capacity 691
cohort 691
community 688
demographic transition 695
doubling time 695
ecology 688
ecosystem 688
environmental resistance 690
exponential growth 690
less-developed country (LDC)
695
logistic growth 690
more-developed country
(MDC) 695
population 688
replacement reproduction
696
sustainable world 697
zero population growth 696
Match the terms to these denitions:
a. Group of organisms of the same species occupy-
ing a certain area.
b. Maximum population growth rate under ideal
conditions.
c. Growth, particularly of a population, in which the
increase occurs in the same manner as compound interest.
d. Due to industrialization, a decline in the birthrate
following a reduction in the death rate so that the population
growth rate is lowered.
e. Largest number of organisms of a particular
species that can be maintained indenitely by a given
environment.
Thinking Scientically
1. Considering the population of zebras described in the rst
paragraph:
a. Plot the growth curve for this zebra population using
number of zebras versus time.
b. How would you describe the shape of your curve?
c. If the zebra population happens to outstrip the carrying
capacity of the environment, what would happen to the
curve?
2. Consider this denition of overpopulation: ...where there are
more people than can live on the earth in comfort, happiness,
and health, and still leave the world a t place for future
generations.
1
a. Do comfort and happiness mean the typical standard of
living seen in developed countries or in less-developed
countries? Should everyone in the world have the same
standard of living? Why or why not?
b. What standard of health is acceptable for the developed
countries? Whose responsibility is it to achieve this end?
c. Should citizens and private industry work to nd ways to
make the world an ecologically t place for future genera-
tions? Why?
d. When discussing overpopulation, should we think in
terms of the world, the country, or the area?
1
From George Morris, 1973. Overpopulation: Everyones Baby. London: Priory Press
Limited, p. 24.
Using Technology
Your study of population ecology is supported by these
available technologies:
Essential Study Partner CD-ROM
Ecology Populations
Human Impact
Visit the Mader web site for related ESP activities.
Exploring the Internet
The Mader Home Page provides resources and tools as
you study this chapter.
http://www.mhhe.com/biosci/genbio/mader

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