
Environmental Encyclopedia 3
Population growth
Populations increase through reproduction, but they
do not increase infinitely because of environmental limita-
tions, including disease, predation,
competition
for space
and nutrients, and
nutrient
shortages. These limits to popu-
lation growth are collectively known as environmental
resis-
tance
.
An important concept in environmental resistance is
the idea of
carrying capacity
. Carrying capacity is the
maximum number of individuals a habitat can support. In
any finite system, there is a limited availability of food, water,
nesting space, and other essentials, which limits that system’s
carrying capacity. When a population exceeds its environ-
ment’s carrying capacity, shortages of nutrients or other ne-
cessities usually weaken individuals, reduce successful repro-
duction, and raise death rates from disease, until the
population once again falls below its maximum size. A popu-
lation that grows very quickly and exceeds its environment’s
carrying capacity is said to overshoot its environment’s capac-
ity. A catastrophic
dieback
, when the population plummets
to well below its maximum, usually follows an overshoot. In
many cases populations undergo repeated overshoot-dieback
cycles. Sometimes these cycles gradually decrease in severity
until a stable population, in equilibrium with carrying capac-
ity, is reached. In other cases, overshoot-dieback cycles go
on continually, as in the well known case of lemmings.
Prolific breeding among these small arctic rodents leads to
a population explosion every four to six years. In overshoot
years, depletion of the vegetation on which they feed causes
widespread undernourishment, which results in starvation,
weakness, and vulnerability to predators and disease. The
lemming population collapses, only to begin rebuilding,
gradually approaching overshoot and another dieback. At
the same time, related populations fluctuate in response to
cycles in the lemming population: populations of owls and
foxes surge when lemmings become plentiful and fall when
lemmings are few. The grasses and forbs on which lemmings
feed likewise prosper and diminish in response to the lem-
ming population.
As a population grows, the number of breeding adults
increases so that growth accelerates. Increase at a constant
or accelerating rate of change is known as
exponential
growth
. If each female lemming can produce four young
females, and each of those produces another four, then the
population is multiplied by four in each generation. After
two generations there are 16 (or 24) lemmings; after three
generations there are 64 (34); after four generations there
are 256 (44) lemmings, if all survive. When environmental
resistance (predation, nutrient limitations, and so on) causes
a population to reach a stable level, without significant in-
creases or decreases over time, population equilibrium is
achieved. Generally we might consider stable (equilibrium)
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populations the more desirable situation because repeated
diebacks involve extensive suffering and death.
These principles of population biology have strongly
informed our understanding of human population changes.
Over the centuries the world’s human population has tended
to expand to the maximum allowed by available food, water,
and space. When humans exceeded their environment’s car-
rying capacity, catastrophic diebacks (usually involving dis-
ease,
famine
, or war) sometimes resulted. However, in many
cases, diebacks have been avoided through emigration, as in
European migrations to the Americas, or through technolog-
ical innovation, including such inventions as agriculture,
irri-
gation
, and mechanization, each of which effectively ex-
panded environmental carrying capacities. For tens of
thousands of years the human population climbed very grad-
ually, until about the year 1000, when it began to grow
exponentially. Where we once needed a thousand years (200
to 1200
A.D.
) to double our population from 200 million to
400 million, at current rates of growth we would require
only 40 years to double our population. Since the eighteenth
century, population theorists have increasingly warned that
our current pattern of growth is leading us toward a serious
overshoot, perhaps one that will permanently damage our
environment
and result in a consequent dieback. The only
course to avoid such a catastrophe, argue population theo-
rists, is to stabilize our population somewhere below our
environment’s carrying capacity.
Popular awareness of population issues and agreement
with the principle of population reduction have spread in
recent years with the publication of such volumes as The
Population Bomb by Paul Ehrlich, and The Limits to Growth
by Donella Meadows and others. The current population
debate, however has older roots, especially in the work of
Thomas Malthus, an English cleric who in 1798 published
An Essay on the Principle of Population as It Affects the Future
Improvement of Society. Malthus argued that, while un-
checked human population growth increases at an exponen-
tial rate, food supplies increase only arithmetically (a constant
amount being added each year, instead of multiplying by a
constant amount). The consequence of such a disparity in
growth rates is starvation and death. The remedy is to reduce
our reproductive rates, where possible by “moral restraint,”
but where necessary by force. Malthus’ work has remained
well-known principally because of its conclusion about social
policy: because providing food and shelter to the poor only
allows them to increase their rates of “breeding,” assistance
should be withheld. If the poorer classes should starve as a
result, argued Malthus, at least greater rates of starvation at
a later date would be avoided. This conclusion continues to
be promoted today by neo-Malthusians, who protest the
principle of aiding developing countries. Poorer countries,
neo-Malthusians point out, have especially dangerous