ECOSYSTEM,

CONCEPT OF
Eugene P. Odum
University of Georgia

I. Introduction
II. The Ecosystem Concept
III. The Human-Dominated Techno-ecosystem

GLOSSARY
autotroph Literally, a self-feeder; an organism that is
able to utilize inorganic carbon (carbon dioxide) as
the sole carbon source for growth; for example, green
plants and certain bacteria.
black box Entity that can be examined at the system
level without specifying its internal contents.
heterotroph Literally, a feeder on others; an organism
that is dependent on organic material from an external source to provide carbon for growth; for example, vertebrates.
industrialized agriculture Modern form of agriculture
that differs from traditional agriculture in the use of
elaborate and expensive machinery, the control of
pests with toxic chemicals rather than biocontrols,
fertilization by synthetic rather than organic products, excessive consumption of water, and farm ownership and management by corporations rather
than individuals.
input environment Collective term for all energy and
materials moving into a given system.
mega-city Modern city with a large, expanding population, characterized by high consumption levels of
energy, water, and food from sources outside the city.

output environment Collective term for all energy and

materials moving out of a given system.
techno-ecosystem Technology-based ecosystem in the
contemporary world that is fundamentally distinct
from natural ecosystems in the use of energy sources
other than sunlight (fossil fuels, nuclear power), an
urbanized concentration of human population, and
the generation of substantial amounts of air and water pollutants and waste materials.

LIVING ORGANISMS AND THEIR NONLIVING (ABIOTIC) ENVIRONMENT are inseparably interrelated
and interact with each other. An ecological system, or
ecosystem, is any unit (a biosystem) that includes all
the organisms (the biotic community) in a given area
interacting with the physical environment so that a ow
of energy leads to clearly dened biotic structures and
cycling of materials between living and nonliving parts.
An ecosystem is more than a geographical unit (or
ecoregion); it is a functional system unit with inputs
and outputs, and with boundaries that can be either
natural or arbitrary.

I. INTRODUCTION
The ecosystem is the rst unit in the molecule to ecosphere hierarchy (as shown in Fig. 1) that is complete,

Encyclopedia of Biodiversity, Volume 2

Copyright 2001 by Academic Press. All rights of reproduction in any form reserved.

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FIGURE 2 This ecosystem model emphasizes the external environment, which must be considered an integral part of the ecosystem
concept. See Fig. 3 for a model emphasizing internal structures
and processes.

FIGURE 1 Hierarchical organization of living systems. The ecosystem is the rst level that is complete.

that is, it has all the components, biological and physical, necessary for survival. Accordingly, it is the basic
unit around which to organize both theory and practice
in ecology. Furthermore, as the shortcomings of the
piecemeal short-term technologic and economic approaches to dealing with complex problems become
ever more evident with each passing year, management
at this level, that is, ecosystem management, emerges as
the challenge for the future.
Since ecosystems are functionally open systems, consideration of both inputs and outputs is an important
part of the concept, as shown in Fig. 2. A diversity of
species and genetic forms, together with a variety of
functions and niches, are essential properties of natural
ecosystems. Ecosystem diversity provides redundancy
in times of environmental uncertainty.

II. THE ECOSYSTEM CONCEPT

The term ecosystem was rst proposed in 1935 by
the British ecologist A. G. Tansley, but of course the

concept is by no means so recent. Allusions to the idea

of the unity of organisms and the environment (as well
as the oneness of humans and nature) can be found as
far back in written history as one might care to look.
Not until the late 1800s did formal statements begin
to appear, interestingly enough, in a parallel manner
in the American, European, and Russian ecological literature. Thus, in 1877 Karl Mobius wrote (in German)
about the community of organisms in an oyster reef as
a biocoenosis, and in 1887 S. A. Forbes, an American,
wrote his classic essay on the lake as a microcosm.
The pioneering Russian V. V. Dokuchaev and his chief
disciple, G. F. Morozov (who specialized in forest ecology), emphasized the concept of the biocoenosis, a
term later expanded by Russian ecologists to geobiocoenosis (Sukachev, 1959).
In addition to biologists, physical scientists and social scientists began to consider the idea that both nature and human societies function as systems. In 1925,
the physical chemist A. J. Lotka wrote in a book entitled
Elements of Physical Biology that the organic and inorganic worlds function as a single system to such an
extent that it is impossible to understand either part
without understanding the whole. It is signicant that
a biologist (Tansley) and a physical scientist (Lotka)
independently and at about the same time came up
with the idea of the ecological system. Because Tansley
coined the word ecosystem and it caught on, he gets
most of the credit, which should be shared with Lotka.
In the 1930s, social scientists developed the holistic
concept of regionalism, especially Howard W. Odum,

ECOSYSTEM, CONCEPT OF

who used social indicators to compare the southern

region of the United States with other regions (Odum,
1936; Odum and Moore, 1938). More recently, Machlis
et al. (1997) and Force and Maddie (1997) have promoted the idea of the human ecosystem that combines
biological ecology and social theories as a basis for
practical ecosystem management. Accordingly, the concept of the ecosystem now brings together organisms,
the physical environment, and humans.
As shown in Fig. 2, a graphic model of an ecosystem
can consist of a box that we can label the system, which
represents the area we are interested in, and two large
funnels that we can label input environment and output
environment. The boundary for the system can be arbitrary (whatever is convenient or of interest), delineating
an area such as a block of forest or a section of beach,
or it can be natural, such as the shore of a lake where the
whole lake is to be the system, or ridges as boundaries of
a watershed.
Energy is a necessary input. The sun is the ultimate
energy source for the biosphere and directly supports
most natural ecosystems within the biosphere. But there

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are other energy sources that may be important for

many ecosystems, for example, wind, rain, water ow,
or fuel (the major source for urban-industrial society).
Energy also ows out of the system in the form of heat
and in other transformed or processed forms such as
organic matter (e.g., food and waste products) and pollutants. Water, air, and nutrients necessary for life,
along with all kinds of other materials, constantly enter
and leave the ecosystem. And, of course, organisms and
their propagules (seeds and other reproductive stages)
enter (immigrate) or leave (emigrate).
In Fig. 2 the system part of the ecosystem is shown
as a black box, which is dened by modelers as a unit
whose general role or function can be evaluated without
specifying its internal contents. Figure 3 is a graphic
model of the solar-powered natural ecosystem showing
internal system components and functions. The interactions of the three basic components, namely, (1) the
community, (2) the ow of energy, and (3) the cycling
of materials, are diagrammed in this simplied compartment model. Energy ow is one-way; some of the incoming solar energy is transformed and upgraded in

FIGURE 3 A functional diagram of a natural ecosystem, with emphasis on internal dynamics involving energy
ow, material cycles, and storage (S), as well as food webs of autotrophs (A) and heterotrophs (H).

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quality (i.e., converted into organic matter, a more upgraded form of energy than sunlight) by the community,
but most of it is degraded and passes through and out
of the system as low-quality heat energy (heat sink).
Energy can be stored, then fed back, or exported, as
shown in the diagram, but it cannot be reused. In contrast with energy, materials, including the nutrients
necessary for life (carbon, nitrogen, phosphorus, and
so on), and water can be used over and over again. The
efciency of recycling and the magnitude of imports
and exports of nutrients vary widely with the type of
ecosystem.
Each box in Fig. 3 is given a distinctive shape that
indicates its general function. Circles are renewable
energy sources, bullets are autotrophs, hexagons are
heterotrophs, and the round-bottomed shapes are storages (in this case of nutrients and organics). The community is depicted as a food web of autotrophs and
heterotrophs linked together with appropriate energy
ows, nutrient cycles, and storages.
Both graphic models (Figs. 2 and 3) emphasize
that a conceptually complete ecosystem includes inputs and outputs along with the system as delimited,
that is, an ecosystem IE S OE (input environment system output environment). This scheme
solves the problem of where to draw lines around
an entity that one wishes to consider, because it does
not matter very much how the box portion of the
ecosystem is delimited. Often, natural boundaries,
such as a lakeshore or forest edge, or political ones,
such as city limits, make convenient boundaries, but
limits can just as well be arbitrary so long as they
can be accurately designated in a geometric sense.
The box is not all there is to the ecosystem, because
if the box were an impervious container, its living
contents (lake or city) would die.
It is important to emphasize that it is the diversity
of ecosystem functions including microbial recycling, inputs and outputs as well as habitats and human land
uses that need to be maintained, not just the diversity
of species or biodiversity in the narrow sense.

III. THE HUMAN-DOMINATED

TECHNO-ECOSYSTEM
Current urban-industrial society not only impacts
natural life-support ecosystems in negative and sometimes positive ways, but has created entirely new
arrangements that we can call techno-ecosystems that

are competitive with and parasitic on natural ecosystems. These human-made systems involve new, powerful energy sources, technology, money, and fuelpowered cities that have little or no parallel in nature.
It is imperative that techno-ecosystems interface with
natural life-support ecosystems in a more positive or
mutualistic manner than is now the case if our rapidly
growing urban-industrial society is to survive in a
nite world.
Before the industrial revolution, humans were a part
ofrather than apart fromnatural ecosystems. In the
ecosystem model of Fig. 3, humans functioned as top
predators and omnivores (the terminal H box in the
food web). Early agriculture, as is the case with traditional or preindustrial agriculture as still widely practiced in many parts of the world, was compatible with
natural systems and often enriched the landscape in
addition to providing food. But with the increasing use
of fossil fuels and atomic ssionenergy sources many
times more powerful than sunlighttogether with the
mushrooming growth of cities and increasing use of
money-based market economics as the basis for decision
making, the model of Fig. 3 is no longer adequate. We
need to create a new model for this techno-ecosystem,
a term suggested by pioneer landscape ecologist Zev
Naveh (1982).
Figure 4 is our graphic model for these new (in terms
of human history) fuel-powered systems. It includes
the four components listed earlier: powerful energy
sources, technology, money, and cities. The model
shows the inputs of the new fuel energy sources and
natural resources, and the increasing outputs of air,
water, and solid waste pollution that are very much
larger and more toxic than anything that comes out of
natural ecosystems. In Fig. 5 we add to the technoecosystem model some natural ecosystems that provide
life-supporting goods and services (breathing, drinking,
and eating!) and that maintain homeorhetic (i.e., pulsing) global balances in the atmosphere, soils, freshwater, and oceans. Note that money circulates as a twoway ow between society and human-made systems,
but not natural systems, thereby creating a vast market
failure when society fails to pay for ecosystem services.
A modern city,1 of course, is the major component
of the fabricated techno-ecosystem. It is a very energetic
hot spot that requires a large area of low-energy natural

1
The term city is used synonymously with the geographers
term standard metropolitan district (SMD), which includes industrial areas and residential suburbs that often extend far beyond ofcial
city limits.

ECOSYSTEM, CONCEPT OF

FIGURE 4 A human-dominated techno-ecosystem.

FIGURE 5 A human-dominated techno-ecosystem and natural ecosystem.

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and seminatural countryside to maintain it. Current

cities clean and recycle no air and or water (to the
point of being redrinkable), grow little or no food, and
generate a huge waste stream that impacts wide areas
of downstream rural landscapes and oceans. The city
does export money that pays for some natural resources,
and the city provides many desirable cultural institutions, such as museums and symphonies, that are not
available in rural areas.
In summary, cities are essentially parasites on the
low-energy countryside. To call a city a parasite is not
to belittle it, but to be realistic. In undisturbed nature,
parasites and hosts tend to coevolve for coexistence;
otherwise, if the parasite takes too much from its host,
both die if the parasite has only one host. Currently
humans have only one habitable hostthe earth.
Especially threatening to the global life-support ecosystems is the explosive growth of mega-cities in the
less-developed nations, caused in part by the increasing
dominance of another techno-ecosystem, that of industrialized agriculture, with its often excessive consumption of water and use of toxic and enriching chemicals.
These systems produce more food products per unit of
space, but in turn are prodigious polluters and by their
economic might drive small farmers out of business
worldwide, forcing them into cities that are unable to
assimilate them. This current situation illustrates what
engineer and former president of MIT Paul Gray (1989)
has written: A paradox of our time is the mixed blessing
of almost every technological development. In other

words, technology has its destructive as well as benecial side. To bring the natural and technical ecosystems
into a mutualist relationship will be societys greatest
challenge in the twenty-rst century.

See Also the Following Articles

ECOSYSTEM SERVICES, CONCEPT OF ENERGY FLOW AND
ECOSYSTEMS HUMAN EFFECTS ON ECOSYSTEMS,
OVERVIEW

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