KH4119_Unit 06 ES_650-E661 03/16/05 9:27 AM Page 650

UNIT 6 ESSAYS
Ecology: Interaction and Interdependence
in Living Systems

Interdependence Involves
Limiting Factors and
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Carrying Capacity
Newspaper articles worldwide report that conditions change. The limiting factors that

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famines continue to occur in Africa and that influence the carrying capacity may be biotic
new animals are being added to the endangered (living), or abiotic (nonliving), or both. Biotic
species list. What are the causes of these events? factors include food supply and other

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Are they caused by human mismanagement, or organisms. Abiotic factors include space, raw
do some happen naturally? materials, and climate.
Scientists agree that we can find some Climate is the prevailing weather

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of the answers by studying the concepts of conditions in a given area through long periods
limiting factors and carrying capacity. of time. Weather conditions result from abiotic
A limiting factor limiting factors that affect all organisms. These
is anything that weather factors include temperature, sunlight

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can slow, or limit,
the growth
of a population.
The combination
intensity, precipitation (rainfall, snowfall, and
fog), humidity (amount of moisture in the air),
and wind. Although you can measure each
factor alone, each affects the others. Together

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of limiting factors they affect the population size of organisms
in a given habitat living in the area. Climate and particular
influences the weather conditions can affect the populations

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carrying capacity
of that habitat.
Carrying capacity
is the maximum
population of a
particular species
that the habitat
can support.
Figure E15.1 The snowy owl (Nyctea
The carrying
scandiaca) inhabits Arctic tundra regions capacity of an
and in winter is always searching for food in environment is not
this sparse environment. When the winter necessarily fixed.
food supply is especially limited, the snowy It changes as
owl can be seen as far south as Colorado. environmental
size directly. They do this by presenting
optimal conditions for various plants to grow
and reproduce or by presenting conditions that
are adverse to growth and reproduction. The
abundance or lack of certain plants translates
into an abundant or limited food supply for a
chain of other organisms. In this way, climate
and vegetation help determine the carrying
capacity of a particular environment for
various organisms. (See Figure E15.1 for
an example.)
You might be asking yourself, How can
temperature be a factor that limits the size of
a population? In the northern part of the

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United States, the first heavy frost of autumn
kills almost all adult mosquitoes. When the
population of insects drops, the population of
organisms that feed on mosquitoes is affected.
In this example, the limiting factor of
temperature affects the mosquito population
directly by killing most of them. By limiting
other organisms food supply, however,
temperature also affects the carrying capacity
of the winter environment for those organisms.
The same type of interactions take place
between various wildflower populations
and the insects that pollinate them. As
temperatures drop in the autumn, the
wildflowers die. The insects then either die,
become dormant, or migrate. In both cases,
temperature has caused limited population
growth for a period of time.
Water is a requirement of all living
organisms. Thus, it is a limiting factor in
every ecosystem in the biosphere. Almost all
chemical reactions that organisms carry out to
stay alive take place in water. Water molecules
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themselves are a part of many chemical
reactions. A few organisms can survive by
becoming inactive when there is no water. But
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most organisms die. Because water is essential
for all living organisms, it is a limiting factor
that influences the carrying capacity of
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Figure E15.2 Water is a limiting factor
in the savanna in East Africa. Here only
grasses and scattered trees can grow. These
plants, in turn, support grazing animals.
The grazing animals move on when the
rainfall in particular locations decreases.
ESSAY: Interdependence Involves Limiting Factors and Carrying Capacity
virtually every ecosystem. Look at Figure
E15.2. Water limits the carrying capacity of
a savanna in East Africa and consequently
limits the distribution of the organisms.
Because of other weather conditions, rain
and snow alone may not ensure that an
environment has enough water. Wind
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increases the rate of water evaporation. Low
humidity in the desert also results in an
increased evaporation rate. Thus, high rates of
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evaporation can affect the survival of certain
organisms, even when the precipitation is
adequate. In some cases, a high rate of
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evaporation together with low rainfall may
permit only plants with special features to
survive. This situation severely limits the
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carrying capacity of the desert. For example,
desert plants often have thick, succulent stems
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Figure E15.3 Desert organisms have
adaptations that enable them to survive
despite the scarcity of water. The fleshy stems
of this saguaro cactus (Carnegiea gigantea)
store water. The plants extensive shallow
root system increases its access to rainwater.
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KH4119_Unit 06 ES_650-E661 03/16/05 9:27 AM Page 652

that can store water. They also have extensive
root systems that can absorb large quantities
of water during infrequent rainfalls (see
Figure E15.3). Some animals, such as the
kangaroo rat, can survive without drinking
water. They use the water that is released
during cellular respiration and produce a
on the energy pyramid, the fewer of them the
environment can support. Therefore, the
amount of space needed by all organisms at
trophic levels above the producers is linked
primarily to a biotic factorthe availability
of food energy.
The available space is affected by

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nearly solid urine. population density. Population density is the
Light also can be a limiting factor. The number of individuals in relation to the space
penetration of light into the ocean determines the population occupies. For example,

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the depth to which photosynthetic organisms consider an experiment at the University of
can grow. This is generally not beyond the Wisconsin, which is illustrated in Figure
depth of 180 meters (590 feet). If silt or algae E15.4. Researchers gave mice in cages more

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growth reduces the water clarity in a pond or food than they needed each day. As the mice
lake, plant growth is limited. In dense rain reproduced, the density of the population
forests, the tallest trees spread their leaves and increased, and the cages became very crowded.

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take most of the light. The ground below Some female mice stopped taking care of their
them is shaded. This prevents other plants nests and young. Mice continued to be born,
from reaching great heights. Light affects but many newborn mice died from neglect.

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animals indirectly. The amount of light
influences the number of plants that can grow.
This then influences the carrying capacity of
the environment.
Eventually, the death rate of the young mice
reached 100 percent. This kept the population
density from increasing further.
In a similar experiment, conducted in

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Space is another abiotic limiting factor for England, the death rate of young mice was
populations. Every individual needs living not affected. Instead, the birthrate declined
space. Some organisms, however, need more almost to zero. In this experiment, the

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space than others. For example, individual corn
plants grow well when they are planted close
together. A mountain lion, on the other hand,
usually requires many square kilometers to find
extremely low birthrate kept the population
density from increasing. Space, as a limiting
factor, affects all living populations and the
carrying capacity of particular ecosystems for

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enough food to sustain itself. You may those populations.
remember that the higher a given organism is

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a b

Figure E15.4 (a) In one population experiment, researchers provided mice with more
than enough food. (b) As a result, the population grew dramatically. Space then became
a limiting factor that resulted in a high death rate among young mice.

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Systems Analysis
Some situations arise that are so complex that
predictions seem at best unreliable and at worst
impossible. When that happens, scientists turn
to a method of analysis called systems analysis.
amount of shading the pond receives (for
instance, is it surrounded by large trees?) and
the stillness of the water. All of these
components influence and define the system.

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In systems analysis, scientists try to understand Once the components of a system are
enough about the interactions of complex identified, most scientists try to make the

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situations so that they can reliably predict system more manageable by limiting the
the effect that certain changes will have on number of components that are included in
the system. the analysis. For instance, our pond ecologist

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Weather provides an example of a complex might organize her analysis to include only
system that is relatively easy to understand in the effects of several different nutrients on
the short term. For example, dark black clouds the number and size of perch in the pond.

usually lead to rain. The interactions that
produce weather are so complex, however, that
it is nearly impossible to understand or predict
weather across large distances or times (see
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The scientist who sets such a structure knows
that outside factors, such as pH and sunlight,
also influence the fish. However, she might
choose not to take those into consideration

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Figure E16.1). Ecosystems are examples of because they further complicate an already
complex systems. Ecosystems are made up of complex study.
living organisms and nonliving elements in an Limiting the number of components is

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environment. necessary because it focuses the scientists
The first step in analyzing any complex
phenomenon is to identify the components

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of the system under study. The components,
or collection of things, that make up an
ecological system include living (biotic) as well
as nonliving (abiotic) things. For example, an

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ecologist might describe a pond in terms of
its biotic components. These would be its
microscopic organisms, plant life, and fish. She

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also might describe the pond in terms of its
abiotic components. These would be the pH of
the pond water, the nutrient and oxygen levels,
and the depth of sunlight penetration. These
components also include elements outside of
the pond. Thus, the pond system is not
isolated from the world around it. In this case,
the ponds pH is influenced by the pH of
rainwater and the pH of anything that might
wash into the pond. The nutrients are
determined by the amount and type of food
that washes or falls into the pond and the types Figure E16.1 Predicting the weather
of organisms that decay there. And the can be difficult because it involves a
penetration of sunlight is affected by the complex system.

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thinking on a particular area of interest.
Developing and testing hypotheses is much
easier when the focus of the investigation is
narrow. Unfortunately, using a limited
structure somewhat reduces the accuracy of
the analysis. Most systems analysts try to
organize their system in a way that is simple
know as much as possible about the behavior
of a system before attempting to analyze the
effect that changes in particular components
might have on it. For this reason, ecologists
collect large amounts of data in an attempt
to understand how one structured set of
interactions affects another. Despite this

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enough to be manageable and complex effort, ecologists realize that they never will
enough to be accurate. have all of the information necessary to make
The last essential step before beginning an error-free prediction. To compensate for

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any analysis of a complex system is that the this limitation, many ecologists choose to
scientist understands as much as possible be conservative in their prediction. They
about the initial behavior of the system. If generally assume that intervening in

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the initial behavior of the system is not ecosystems will result in some unpredictable
understood, then it is difficult to interpret consequences.
how the system will react when one of its

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components is altered. In the pond example,
say the ecologist does not already know that
perch populations decline each fall. (This

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happens because a particular nutrient that
they require becomes scarce.) In that case, she
might mistakenly attribute the decline to a
recent change in pH. (In fact, the pH change

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might influence the perch population. But
this effect may be small compared with the
annual fluctuations caused by varying

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nutrient levels.)
Once a limited analysis is complete,
scientists may try to combine the results of
several analyses. This way, they can construct

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a more complex, and more accurate,
understanding of the whole system. Figure E16.2 A lake or pond is a good
From this brief description of systems example of a complex system that interacts

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analysis, you can see that it is important to with the world outside of it.

Environmental Ethics and You

Environmental ethics is a branch of ethical
study that considers how people ought to relate
to the natural world. The different ethical
perspectives people have about humanitys
relationship to nature are based on different
values, beliefs, and attitudes. Peoples values,
beliefs, and attitudes about nature are formed
by a variety of influences including the
overarching values and attitudes of their
culture. Peoples attitudes about nature also are
influenced by their personal experiences, the
people they talk to, and the books they read.
These values taken together influence how
people behave toward the environment. This
has a major impact on the state of the natural
world around us.

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Following are brief descriptions of two intrinsic value

rather distinctive ethics. How similar or regardless of their
different is your personal environmental ethic usefulness to
from the two views described here? What humankind. From this
factors have influenced your environmental perspective, humans
ethic? What other environmental ethical have no right to reduce
models are you aware of? the earths biological

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The human-centered environmental richness and diversity
Human-centered ethic
ethic. People who support a human-centered except to satisfy
environmental ethic take the position that fundamental needs.

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humans should dominate the natural world People who support
because they are fundamentally different from this position think that
all other life-forms and are somehow superior the earths resources are

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to or separate from nature. From this in limited supply and
perspective, nonhuman life-forms are that these resources are
commodities that have no value other than for all life-forms, not

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their usefulness to humankind. People holding just for humans. The
this ethic tend to view the earth primarily as a deep ecology ethic
collection of natural resources that humans can takes the position that

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use to promote economic growth and both human and
prosperity. People who support this position nonhuman life will
also think that the earth is vast and has many continue to flourish Deep ecology ethic
resources in unlimited or abundant supply. only if humans greatly

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They point out that human history has been reduce their rate of
characterized by continual progress and that, population growth. People who hold this
through technology, people find solutions to all position think that human interference in the

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problems (including those of resource depletion
and pollution), and progress continues.
The deep ecology environmental
ethic. The deep ecology ethic is a life-
nonhuman world has been excessive and
human impact must be minimized. From this
perspective, people must make economic,
technological, and ideological changes that

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centered ethic. People who support this promote a sustainable lifestyle rather than one
position think that all life-forms on earth have that seeks the highest standard of living.

D Figure E16.3 The Florida Everglades

was established as a national park in
1934 to preserve the plants and animals
that are part of this unique wetland
environment. The influence of humans,
however, has caused disruptions in the
ecosystem. People who have different
viewpoints about the environment have
different ideas about how a unique
environment such as the Everglades
should be managed. What is your
point of view? Do you view this with
a human-centered or a deep ecology
environmental ethic?

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Growing, Growing, Grown

Just look around. You can observe plenty of The J-Shaped Curve
evidence that populations grow. The guppies
in your fish tank have more guppies, the
songbirds in your yard have young each

number of organisms
spring, and the stray cats in the neighborhood
seem to have a litter of kittens every couple of

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months. But have you ever thought about how
populations grow?
In 1798, after years of thinking about

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this question, Thomas Malthus (pictured in
Figure E16.4) reported that in ideal conditions,
populations tend to grow exponentially. In time

exponential growth, both the total population
size and the rate of increase rise steadily across
time. Charles Darwin later wrote in his book
On the Origin of Species (1859) that, There is
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Figure E16.5 The J-shaped curve is
characteristic of populations that are
growing exponentially.

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no exception to the rule that every organic 5.5 trillion flies in several generations or just
being naturally increases at so high a rate, 1 year (refer to Figure E16.6). Even the rather
that, if not destroyed, the earth would soon slow-breeding elephant has the capacity for

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be covered by the progeny of a single pair. explosive population growth. Charles Darwin
Biologists discovered that any population calculated that an elephant population of just
growing exponentially can potentially two elephants (a single breeding pair) could

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approach an infinite size in a relatively short
amount of time if provided with necessary
resources such as food, water, space, and
protection from other organisms.
grow to 19 million elephants in only 750 years.
PREDICTED POPULATION GROWTH OF
THE COMMON HOUSEFLY IN ONE YEAR

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When populations grow
exponentially, the population
growth curve takes on a J-shaped
Generation Population size
1 120
2 7,200
3 432,000

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Figure E16.4 Thomas
Malthus was an English
economist who lived from were ideal, a housefly population
1766 until 1834. He gave
serious thought to how
populations grow.
appearance. This is due to the
continuous increase of a larger
and larger population size (see
Figure E16.5). You can observe this
principle of population growth
when you examine a fast-breeding
species such as the common
housefly or a slow-breeding species
such as the elephant. Biologist
L. O. Howard discovered that, if
the environmental conditions
beginning with just one adult
female could increase to more than
4 25,920,000
5 1,555,200,000
6 93,312,000,000
7 5,598,720,000,000
Source: Data from Kormondy, E. J. (1984). Concepts of Ecology,
third edition. Englewood Cliffs, NJ: Prentice Hall Publishers.
Figure E16.6 Predicted population
growth of the common housefly in
1 year. This prediction is based on the
following observations and assumptions.
An average female fly lays 120 eggs at a
time. About half of the eggs develop into
females. There are seven generations in
1 year. Individual flies live for one
generation.

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Under ideal conditions, populations tend to

increase in size exponentially (that is, 1, 2, 4, 8, The S-Shaped Curve
16, 32, 64, 128). In addition to realizing that,
Thomas Malthus realized that the food supply
tends to increase only arithmetically (for carrying capacity of environment
example, 1, 2, 3, 4, 5, 6, 7). Most of the worlds

number of organisms
organisms serve as the food supply for other

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organisms higher on the food chain. Because of
this and other limiting factors such as space,
populations generally do not continue to grow

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exponentially. When a food supply is limited,
the environments carrying capacity for the
populations that feed on that food supply is

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reduced.
Technological advances in agriculture a time
have helped food production generally keep

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pace with human population growth. Still,
many scientists are concerned that a time will
come when food production will not meet the

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requirements of the worlds rapidly increasing
human population. In other words, we will Population Growth of Paramecium
reach a point when the human population carrying capacity = 375
400
number of organisms

will exceed the carrying capacity of the earth.

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In theory, populations could continue to
grow exponentially if they are provided with 300
ideal conditions and unlimited resources. In

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the real world, conditions are not ideal and
200
resources are in limited supply. So we do not
observe ongoing exponential growth. Logistic
growth represents a pattern of population 100

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growth that scientists have observed in real
populations in natural conditions. The logistic 0
growth curve has the shape of a flattened S 1 2 3 4 5 6

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(see Figure E16.7), rather than the J-shape b days
that is characteristic of exponential growth. In Source: From Gause, G. F. (1934). The Struggle
exponential growth, the population continues for Existence. Baltimore: Williams and Wilkins.

to increase across time. But in logistic growth,

the population levels off at the carrying Figure E16.7 (a) An idealized S-shaped growth
capacity of the environment. curve. (b) Population growth curve for the single-
celled organism Paramecium. Limiting factors force
this population of organisms to level off.

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an ancient king. The king was so delighted

with the game that he asked the
mathematician to name his own reward.
He told the king that he was a humble man
who wished a humble reward. He asked
that a single grain of wheat be placed on the
first square of the chessboard, two grains of

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wheat on the second square, four grains on
the third square and so on, doubling the
number of grains on each square until all

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64 squares on the chessboard were filled.
The king granted his request and ordered
the Master of the Royal Granary to begin

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counting out the grains. But before the
task was anywhere close to being finished,
Legend has it that the game of chess was the king handed his kingdom over to the

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invented by a mathematician who worked for mathematician.

Endless Interactions
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You have just finished a delicious bowl of One type of direct interaction is a
chicken soup for lunch. As you clean up, you predator/prey interaction. In this, one type

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wash out the bowl and throw away the can. of organism (the predator) eats the other
A garbage truck will haul the can to the dump (the prey). The predator benefits from the
where it will lie among the trash. A female fly
might lay her eggs in the can. Her offspring

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might fly from the dump, feed on some decayed
food, and then rest on top of someones peanut
butter sandwich. Each of these interactions

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depends on the one before it. Most of the time,
these interactions go unnoticed.
All populations, including human
populations, interact with one another in a
complex web of relationships. The set of
interacting populations present at one time
in one place is called a community. In your
community, there may be dogs, cats, trees,
weeds, and humans that interact. When you
mow your lawn or your dog bites the mail
Figure E16.8 For food, this red-tailed
carrier, for example, the interaction is very hawk (Buteo jamaicensis) depends directly
direct. Much of the time, however, and indirectly on a number of organisms in
interactions are indirect. its habitat. Note the rodent in its talons.

658 Unit 6 ESSAY: Endless Interactions

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relationship, but the individual prey does not.
An animal can spend its life as a predator and
then abruptly become the prey. A snake, for
example, may prey on ground squirrels and
then become prey for a hawk. The snake
depends on a large population of ground
squirrels. The hawk, then, depends directly
on the snake, and in this case, indirectly on
the quantity of ground squirrels. The hawk,
however, also may prey directly on the
ground squirrels. The limiting factor in this
case is the food supply. Although the
individual ground squirrel does not benefit,
the population of ground squirrels may
benefit by being held in check by the snake
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Figure E16.9 The melaleuca tree (Melaleuca quinqueneruia) was introduced to Florida
from Australia. It has been overwhelming the cypress in the everglades.
population. There also might be a selective
advantage to the ground squirrel population
because the snakes would tend to prey on the
slower, weaker squirrels. Consequently, the
best-adapted ground squirrels would be
more likely to escape becoming prey and
thus would survive to reproduce.
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Competition is another type of direct
interaction. This type of interaction between
organisms benefits neither one. Organisms
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compete for limiting factors, such as space,
food, sunlight, nutrients, and water (see
Figure E16.9). Competition among
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organisms may increase as a particular
resource becomes scarce.
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When tadpoles live in densely packed
areas, the competition increases for available
space and food resources. As a result, these
tadpoles remain tadpoles longer, they suffer
higher death rates, and they develop into
smaller frogs. In communities of wolves,
competition leads to the establishment of
social hierarchies. A high position in the pack
gives the individual an advantage in terms of
obtaining food, mates, shelter, and other
resources. Individuals of any species that
compete successfully can survive to reproduce
and pass on their genetic material to future
generations. In this way, populations adapt
through time to changes in the environment.
Competition also can take place between
different species. If the species needs are
similar and resources are scarce, then more
competition will exist. In New Guinea, four
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species of pigeons rely on a particular fruit
tree for their primary food source. The four
types of pigeons differ in size, and the smaller
pigeons have adapted to feed on the fruit of
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the lower branches.
An experiment involving paramecium
demonstrates another example of this form
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of adaptation. Researchers placed two species
of paramecium in the same test tube, which
was filled with liquid food. The researchers
formulated a hypothesis that one of the species
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would die out as a result of competition. In
fact, both species survived and thrived. One
species fed on the food that settled at the
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bottom of the tube. The other fed on the food
suspended in the liquid. The two species of
paramecium, as well as the four species of
pigeons, occupied separate niches. The niche
of an organism refers to its role in the
community. This is what it eats, what
organisms eat it, where it lives, and what
indirect relationships it has with other
organisms. As a result of occupying different
niches, the two species could feed and
660 Unit 6 ESSAY: Endless Interactions
reproduce without interfering with each
other even though they lived in the same
place, or habitat.
Competition is not the only relationship
that exists among populations of different
species that live in the same habitat. Some
different species of organisms live in direct,
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physical contact with one another. This
relationship is called symbiosis. For example,
certain species of fungi live on the roots of
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many plants. The fungi absorb nutrients from
the soil and secrete an acid that makes the
nutrients available to the plant. At the same
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time, the fungi are nourished by photosynthetic
products from the plant. The fungi also absorb
water and protect the plant against various
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pathogens in the soil. This type of symbiosis
that benefits both organisms is called
mutualism. Another example of mutualism is
the interaction between a cow and a certain
type of bacteria that live in its intestines. The
cow benefits because the bacteria digest the
cellulose in the plants that the cow eats. The
bacteria benefit because the cow provides a
steady supply of food.
A common form of symbiosis is
parasitism. In parasitism, one organism
(the parasite) lives on or in another organism
(the host) and uses it as a food source. The host
usually remains alive during the interaction.
The parasites, however, may weaken a host to
the extent that it becomes susceptible to disease
or becomes an easier prey for predators. A
leech, for example, clings to a turtles skin and
sucks its blood. Other parasitic microorganisms
in the turtle absorb food directly from its
blood. Interactions such as these eventually
may weaken the turtle. In humans, tapeworms
absorb food directly from the intestines, where
they live.
Plants also may have parasites. Large
microorganisms may have smaller parasitic
microorganisms in them. Parasites may be
KH4119_Unit 06 ES_650-E661 03/16/05 9:28 AM Page 661
Figure E16.10 The mistletoe growing in this pine tree is a parasitic plant.
molds, microorganisms, and even other plants
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such as the mistletoe shown in Figure E16.10.
All viruses are parasitic because they require a
host to reproduce and acquire energy.
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These examples illustrate a few of the
many different types of interactions that take
place among organisms. Some of these
interactions, such as mutualistic relationships,
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are beneficial for both organisms. Other
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interactions are not as positive. These
interactions result in some degree of harm
to one member of the interacting pair, as
in parasitic relationships. Each of these
interactions is influenced by limiting factors
such as light, temperature, food supply, and
space. Because of these interactions, there
is a web of interdependence among living
things.
ESSAY: Endless Interactions Unit 6 661