
Environmental Encyclopedia 3
Ecosystem
few years by Evelyn Hutchinson, Raymond Lindeman, and
the Odum brothers, Eugene and Howard.
The boundaries of an ecosystem can be somewhat
arbitrary, reflecting the interest of a particular ecologist in
studying a certain portion of the landscape. However, such
a choice may often represent a recognizable landscape unit
such as a woodlot, a wetland, a stream or lake, or—in the
most logical case—a
watershed
within a sealed geological
basin, whose exchanges with the
atmosphere
and outputs
via stream flow can be measured quite precisely. Inputs and
outputs imply an
open system
, which is true of all but the
planetary or global ecosystem, open to
energy flow
but
effectively closed in terms of materials except in the case of
large-scale asteroid impact.
Ecosystems exhibit a great deal of structure, as may
be seen in the vertical partitioning of a forest into tree, shrub,
herb, and moss layers, underlain by a series of distinctive
soil
horizons. Horizontal structure is often visible as a mosaic
of patches, as in forests with gaps where trees have died and
herbs and shrubs now flourish, or in bogs with hummocks
and hollows supporting different kinds of plants. Often the
horizontal structure is distinctly zoned, for instance around
the shallow margin of a lake; and sometimes it is beautifully
patterned, as in the vast
peatlands
of North America that
reflect a very complicated
hydrology
.
Ecosystems exhibit an interesting functional organiza-
tion in their processing of energy and matter. Green plants,
the primary producers of organic matter, are consumed by
herbivores, which in turn are eaten by carnivores that may
in turn be the prey of other carnivores. Moreover, all these
animals may have
parasites
as another set of consumers.
Such sequences of producers and successive consumers con-
stitute a food chain, which is always part of a complicated,
inter-linked food web along which energy and materials
pass. At each step along the food chain some of the energy
is egested or passed through the organisms as feces. Much
more is used for metabolic processes and—in the case of
animals—for seeking food or escaping predators; such energy
is released as heat. As a consequence only a small fraction
(often of the order of 10%) of the energy captured at a given
step in the food chain is passed along to the next step.
There are two main types of food chains. One is made
up of plant producers and animal consumers of living organ-
isms, which constitute a grazing food chain. The other con-
sists of organisms that break down and metabolize dead
organic matter, such as earthworms,
fungi
, and bacteria.
These constitute the
detritus
food chain. Humans rely
chiefly on grazing food chains based on
grasslands
, whereas
in a forest it is usual for more than 90% of the energy trapped
by
photosynthesis
to pass along the detritus food chain.
Whereas energy flows one way through ecosystems
and is dispersed finally to the atmosphere as heat, materials
427
are partially and often largely recycled. For example,
nitro-
gen
in rain and snow may be taken up from the soil by
roots, built into leaf protein that falls with the leaves in
autumn, there to be broken down by soil
microbes
to ammo-
nia and nitrate and taken up once again by roots. A given
molecule of nitrogen may go through this
nutrient
cycle
again and again before finally leaving the system in stream
outflow. Other nutrients, and
toxins
such as
lead
and
mer-
cury
, follow the same pathway, each with a different
resi-
dence time
in the forest ecosystem.
Mature ecosystems exhibit a substantial degree of
sta-
bility
, or dynamic equilibrium, as the endpoint of what is
often a rather orderly
succession
of
species
determined
by the
nature
of the
habitat
. Sometimes this successional
process is a result of the differing life spans of the colonizing
species, at other times it comes about because the colonizing
species alter the habitat in ways that are more favorable to
their competitors, as in an
acid
moss bog that succeeds a
circumneutral sedge fen that has in its turn colonized a pond
as a floating mat. Equilibrium may sometimes be represented
on a large scale by a relatively stable mosaic of small-scale
patches in various stages of succession, for instance in fire-
dominated pine forests. On the millennial time scale, of
course, ecosystems are not stable, changing very gradually
owing to immigration and emigration of species and to
evolutionary changes in the species themselves.
The structure, function, and development of ecosys-
tems are controlled by a series of partially independent envi-
ronmental factors:
climate
, soil parent material,
topogra-
phy
, the plants and animals available to colonize a given
site, and disturbances such as fire and windthrow. Each
factor is, of course, divisible into a variety of components,
as in the case of temperature and precipitation under the
general heading of climate.
There are many ways to study ecosystems. Evelyn
Hutchinson divided them into two main categories, holistic
and meristic. The former treats an ecosystem as a “black
box” and examines inputs, storages, and outputs, for example
in the construction of a lake’s heat budget or a watershed’s
chemical budget. This is the physicist’s or engineer’s ap-
proach to how ecosystems work. The meristic point of view
emphasizes analysis of the different parts of the system and
how they fit together in their structure and function, for
example the various zones of a wetland or a
soil profile
,or
the diverse components of food webs. This is the biologist’s
approach to how ecosystems work.
Ecosystem studies can also be viewed as a series of
elements. The first is, necessarily, a description of the system,
its location, boundaries, plant and animal communities, envi-
ronmental characteristics, etc. Description may be followed
by any or all of a series of additional elements, including:
1) a study of how a given ecosystem compares with others