Fundamentals of Ecology

FI FTH E DITIO N

Eugene P. Odum, Ph.D.

Late of Universit y of Georgia lnsntute of Ecology

Gary W. Barrett, Ph.D.

Odum ProJessor oJ Ecolof;y, University of Georgia Institute oJ Ecology

BR CKMW Australia • Canada • Mexico • Singapore • Spain • United Kingdom • United Stateg
Ecoephere

Biome

Landscape

Ecosystem

Population

Organism

'gan sy${0

Organ

Type TheS
cope of Eealogy

Ecology: History and Relevance to

Humankind
2 LeveIs•of•Organization Hierarchy
The Emergent Property Principle
Transcending Functions and Control
Processes
Ecological Interfacing
6 About Models
7 Disciplinary Reductionism to Transdisciplinary

1
1 Ecology: Hlstory anM Relevance to
The word ecology is derived from the Greek oiitos, meaning “household,” and logos,
meaning “study.” Thus, the study of the environmental house includes all the
organ- isms in it and all the functional processes that make the house habitable.
Literally, then, ecology is the study of “lite at home” with emphasis on “the totality
or pat- tern of relations between organisms and their environment,” to cite a
standard dic- tionary definition of the word tMerriAm-\Vebsters Collegiate Dictionary,
10th edition,
ecology”)
The word economics is also derived from the Greek root oihos. As nomics
means ’management," economics translates as ’the management of the household”
and, ac- cordingly, ecology and economics should be companion disciplines.
Unfortunately, many people view ecologists and economists as adversaries with
antithetical visions. Table 1-1 attempts to illustrate perceived differences between
economics and ecol ogy. Later, this book will consider the confrontation that results
because each disci- pline takes a narrow view of its subject and, more important,
the rapid development of a new interface discipline, ecological economics, that is
beginning to bridge the gap between ecology and economics (Costanza,
Cumberland, ct a1. 1997; Barrett and Fa- rina 2000; L. R. Brown 2001).
Ecology was of practical interest early in human history. In primitive society, all
individuals needed to know their environment—that is, to understand the forces of
nature and the plants and animals around them—to survive. The beginning of civi-
lization, in fact, coincided with the use of fire and other tools to modify the
environ ment. Because of technological achievements, humans seem to depend less
on the natural environment for their daily needs; many of us forget our continuing
depen- dence on nature for air, water, and indirectly, food, not to mention
waste assimila- tion, recreation, and many other services supplied by nature. Also,
economic systems, of whatever political ideology, value things made by human
beings that primarily benefit the individual, but they place little monetary value
on the goods and services of nature that benefit us as a society. Until there is a
crisis, humans tend to take nat

School of Cornucopian Neo-Malthusian

thought
Energy
S-shaped
Gromnh{om
K-selected
Technological approach High technology Appropriate technology
System Services provided Services provided
by econom c by natural capital
capit
Resource use Linear (disposal) expansion
em al expan
Futuristic goal Exploration and
Circular (recycling)
g capa
Sustainability and
 SECTION 1 Ecology: History and Relevant to Humankind 3

Figure 1•1. Earthscage as viewed from Apollo 17 travel-

ing toward the Moon. View of the ecosphere from “outside
the box?
;0s
n-
Ily,

.ic-

‘i’‘ ’
OI—

;ap ural goods and services for granted; we assume they are unlimiied or somehow re-
'a- placeable by technological innovations, even though we know that life necessities
such as oxygen and water may be recyclable but not replaceable. As long as the
life- support services are considered free, they have no value in current market
systems (see H. T. Odum and E. P. Odum 2000).
Like all phases of leasing, the science of ecology has had a gradual if
›n-
spasmodic development during recorded history, The mitings of Hippocrates,
he
Aristotle, and other philosophers of anciens Greece clearly contain references to
:n-
ecological topics. However, the Greeks did not have a word for ecology. The word
la-
ecology is of recent origin, having been first proposed by the German biologist
ns,
Ernst Haeckel in 1869. Haeckel defined ecology as “the study of the natural
ily
environment including the re- lations of organisms to one another and to their
surroundings” (Haeckel 1869). Be- fore this, during a biological renaissance in the
at-
eighteenth and nineteenth centuries, many scholars had contributed to the subject,
even though the word ecology was not in use. For example, in the early 1700s,
Antoni van Leeuwenhoek, best known as a premier microscopist, also pioneered
the study of food chains and population regu- lation, and the wiitings of the English
botanist Richard Bradley revealed his under- standing of biological productivity.
All three of thèse subjects are important areas of modem ecology.
As a recognized, distinct field of science, ecology dates from about 1900, but
only in the past few decades has the word become part of the general vocabulary.
At first, the field was rather sharply divided along taxonomic lines (such as plant
ecology and animal ecology), but the biotic community concept of Frederick E.
Clements and Victor E. Shelford, the food chain and material cycling concepts of
Raymond Linde- man and G. Evelyn Hutchinson, and the whole lake studies of
Edward A. Birge and Chauncy Juday, among others, helped establish basic theory
for a unified field of gen- eral ecology. The work of thèse pioneers will be cited
often in subsequent chapters. What can best be described as a worldwide
environmental awareness movement burst upon the scene during two years,
1968 to 1970, as astronauts took the first photographs of Earth as seen from
outer space. For the first time in human history, we were able to see Earth as a
whole and to realize how alone and fragile Earth hov- ers in space (Fig. 1-1).
Suddenly, during the 1970s, almost everyone became con-
4 CHAPTER 1 ne Scope of Ecology

cemed about pollution, natural areas, population growth, food and energy con-
sumption, and biotic diversity, as indicated by the wide coverage of environmental
concerns in the popular press. The 1970s were frequently referred to as the
“decade of the environment,” initiated by the first “Earth Day” on 22 April 1970.
Then, in the 1980s and 1990s, environmental issues were pushed into the pofitical
background by concerns for human relations—problems such as crime, the
cold war, govern- ment budgets, and welfare. As we enter the early stages of the
twenty-first century, environmental concerns are again coming to the forefront
because human abuse of Earth continues to escalate. We hope that this time, to use
a medical analogy, our em- phasis will be on prevention rather than on treatment,
and ecology as outlined in this book, can contribute a great deal to prevention
technology and ecosystem health (Barrett 2001).
The increase in public attention had a profound effect on academic ecology. Be-
fore the 1970s, ecology was viewed largely as a subdiscipline of biology. Ecologists
were staffed in biology departments, and ecology courses were generally found
only in the biological science curricula. Although ecology remains strongly rooted
in bi- ology, it has emerged from biology as an essentially new, integrative
discipline that links physical and biological processes and forms a bridge between
the natural sci- ences and the social sciences (E. P. Odum 1977). Most colleges
now ofer campus- wide courses and have separate majors, departments, schools,
centers, or institutes of ecology. While the scope of ecology is expanding, the
study of how individual or- ganisms and species interface and use resources
intensifies. The multilevel approach, as outlined in the next section, brings together
“evolutionary” and “systems” think- ing, two approaches that have tended to divide
the field in recent years.

Levels-of-Organization Hierarchy

Perhaps the best way to delimit modem ecology is to consider the concept of levels
of organization, visualized as an ecological spectrum (Fig. 1-2) and as an
extended ecological hierarchy (Fig. 1-3). Hierarchy means “an arrangement
into a graded series” (Merrinm-Webster’s Collegiate Dictionary, 10th edition, s.v.
“hierarchy”). Inter- action with the physical environment (energy and matter) at
each level produces characteristic functional systems. A system, according to a
standard definition, con- sists of “regularly interacting and interdependent
components forming a unified

BIOTIC COMPONENTS Genes Cells Organs Organisms Populations Communities

plus
ABIOTIC COMPONENTS Matter Energy
equals ]

BIO6YSTEMS Genetic CeII Organ Organismic Population Ecosystems

systems systems systems systems sy$tems

Figure 1•2. Ecological levels-of-organization spectrum emphasizing the interaction of living

(biotic) and nonliving (abiotic) components.
 SECTION 2 Levels-of•Organization Hierarchy 5

Figure 1-3. Ecological levels- Energetics

of organization hierarchy; seven Evolution Behavior
transcending processes or func-
Development
tions are depicted as vertical
components of eleven integra-
Reguatioo Integration
Ecosphere
ti\ie levels of organization
(after Barrett et a). 199’7).
Biome

Landscape

Ecosystem

Community

Population

'Oan system

O,gan

whole" (Merriam-\Vebstcr’s Collegiate Dictionnry, 10th edition, s.v. “system”).

Systems containing living (biotic) and nonliving (abiotic) components constitute
biosJstems, ranging from genetic systems to ecological systems (Fig. 1-2). This
spectrum may be conceived of or studied at any level, as illustrated in Figure 1-2,
or at any inter- mediate position convenient or practical for analysis. For example,
host-parasite sys- tems or a two species system of mutually linked organisms (such
as the fungi-algae partnership that constitutes the lichen) are intermediate levels
between population and community.
Ecology is largely, but not entirely, concerned with the system levels beyond
that of the organism (Figs. 1 3 and 1 4). In ecology, the term population, originally
coined to denote a group of people, is broadened £o include groups of individuals
of any one find of organism. Likewise, community, in the ecological sense
(sometimes designated as “biotic community"), includes all the populations
occupying a given area. The community and the nonliving environment function
together as an eco- logical system or ecosystem. Biocoenosis and hiogeocoenosis
(literally, “life and Earth
6 CHAPTER. 1 The.Scope of Ecology

Figure 1•4. Compared with the strong aet-point Eçosphere

controls at the organism level and below, organization
and function at the population level and .above are much
Jess .tigh’tIy regu- lated, with more pulsing.and dha?tic
behavior, but ihay are controlled neyertheleae, by No éet-point controls
Landccapes
alternating positive and negative Teeoaacs-in omer WofiO feedback (+ and -)
,.tne exni@*t nomeornes/e ae op- pond to Romeo fasis. I maintaining
Failure to ra gñiz» thie differ nce in cybernetica has Ecosystems pulsing states
reeulted in much collusion about the bal- ance. of
nature. Communities HOMEORHESI6

Pogulations

Organ syetems
Set-point controls
Organs feedback (+ and
-)
Tissues steady states

Cells HOMEOSTA6I9

Moleculee

AtOfTÏS

functioning together"), terms frequently used in European and Russian literature,

are rouglily eqü ivalent to community and ecosystem, respectively. Referring again
to Figure 1-3, 'the next level in the ecological hierarchy is the landscape, a term
origi- nally referring to a painting and defined as an.expanse. of scenery seen by
the eye as one view" tMerriam-Websfer’sCollegiate Dictionnry, l0th edition, s.v.
“landscape”). In ecology, landscape is defined as a “heterogenous area
composed of a cluster of in- teracting ecosystems that are repeated in a similar
manner throughout” tForman and Godron 1986). A watetsíied is a convenient
landscape-level unit for large-scale study and mariagement because it usually has
identifiable natural boundaries. Biome is a term in wide use for a large regional or
subcontïnentaI'system characterized by a ma- jor vegetation type or other
identifying.'landscape ,aspect, as, for exemple, the 'Tem- perate Deciduous Forest
biome or the Conó nental Shelf Ocean biome. A region is a large geological or
political area that may contain more than one biome—for ex- emple, the
regions. of the Midwest, the Appalachian Mountains, or the Pacific Coast. The
largest and ríiost nearly self-suíficient biologicalsystem is. often desigpated as the
ecosphere, which includes all the living organisms of Earth interacting with
the physical environment as a whole to maintain a self-adjusting, loosely
controlled puls- ing°state (more about the concept of “pulsing state” later in this
chapter).
Hierarchical.theory provides,a convenient framework for subdividing and
exam- ining compten situations or extensive gradients, btit,it is more than just a
useful rank- order classification. It is a holistic approach to understanding and
dealing with'com-
 SECTION 3 The Emergent Property Principle 7

plen situations, and is.an alternative to.the reductionist approach of seeking

answers by reducing problems to lower-level analysis (Ahl and Allen 1996).
More than 50 years ago, N’ovlkoff (l9i5) pointed out that there is both
contimi- its and discontinuity in the evolution of the universe. Development
ma7 lie viewed
,as continuous because it involves never-ending change, but it is. also discontinuous
b.ecause it passes through a series of dinerent leveb of organization. As we shall
dis- cuss in,Chapter 3, the organized.stare'of life is maintained by a continuous
but.step- wise Oow of energy. Thus, dividing.a graded series, or hierarchyi into
components is in many cases arbitrary, but sometimes subdivisions can be based
on natural discon- tinuities. Because each level in the levels-of-organizetion
spectnim is “integrated” or interdependent with other.levels, there can be no.sharp
lines or breaks in a functional sense, not even between organism and population.
The individual organism, for ex- ample, .cannot survive for long without .its
population, any more than the organ would be able to survive. for long as a
self-perpetuating unit without its organism. Similarly, the .community
cannot.exist without the cycling of materials and the flow
.of energy in the ecosystem. This argument is applicable to the previously discussed
mistaken notion that human civilization can exist.separately from the natural
world. It is very important to emphasize.that hierarchies in nature are nertrd—that
is, each level is made up of groups of lower-level units (popula'tions are composed
of groups of organisms, ,for example). In sharp contrast, human-organized
hierarchies in governments, cooperations, universitiñ , or the. military are
nortnestrd (sergeants are npt composed of groups of privates, for example).
Accordingly, human-organized hierarchies tend to be more rigid and more sharply
separated as compared to natu- rallevels of organization. For more on hierarchical
theory, see T. F. H. Allen and Statr
(1982), O’Neill ct a1. (1986), and Ahl and Allen (1996).

3
An imponant consequence of hierarchical organization is that as c0mponents, or
.subsets, are combined to produce larger functional, wholes, new properties emerge
that were rtot present at .the level below. Accordingly, an emergent property of
an.
.ecological level or unit cannot be predicted from the stud7 ° *• components.of
that level or unit. Another way to express.the same concept is nonreducible
property—
i- that.is, a property of the.whole not reducible to, the.sum of the propenies of the
parts.
a Though findings at any one level aid.in the.smdy of the next Ievel„ they
ngvgr com- pletely explain.the phenomena occurring at the next level', which must
itself be stud-
t. ied to complete the picture.
ie Two examples, one. from the physical realm and gne from the ecological realm,
ie wik.sulfice to illustrate emergent properties. When hydrogen and oxygen are com-
bined in a certain molecular configuration, ,water is formed—a liqliid wi'th proper-
ties utterly di8erent from tho.se of its gaseous components. When certain algae and
coelenterate. anirñ als ,evolve together to produce a coral, ari e&c'ient nutrient
cycling mechanism is created that enables the combined system to maintain
a.high rate of productivity in waters with a very low nutrient content. Thus,
the.fabu1ous produc-
8 CHAPTER 1 The Scope of Ecology

tivity and diversity of coral reefs are emergent properties only at the level of the
reef community.
Salt (1979) suggested that a distinction be made between emergent properties,
as defined previously, and collective properties, which are summations of the
behav- ior of components. Both are properties of the whole, but the collective
properties do not involve new or unique characteristics resulting from the
functioning of the whole unit. Birth rate is an example of a population level
collective property, as it is merely a sum of the individual births in a designated
time period, expressed as a fraction or percent of the total number of individuals in
the population. New properties emerge because the components interact, not
because the basic nature of the components is changed. Parts are not “melted
down,” as it were, but integrated to produce unique new properties. It can be
demonstrated mathematically that integrative hierarchies evolve more rapidly from
their constituents than nonhierarchical systems with the same number of elements;
they are also more resilient in response to disturbance. Theoretically, when
hierarchies are decomposed to their various levels of subsystems, the latter can still
interact and reorganize to achieve a higher level of complexity.
Some attributes, obviously, become more complex and variable as one proceeds
to higher levels of organization, but often other attributes become less complex and
less variable as one goes from the smaller to the larger unit. Because feedback
mech- anisms (checks and balances, forces and counterforces) operate throughout,
the am- plitude of oscillations tends to be reduced as smaller units function within
larger units. Statistically, the variance of the whole-system level property is less
than the sum of the variance of the parts. For example, the rate of photosynthesis of
a forest community is less variable than that of individual leaves or trees within the
commu- nity, because when one component slows down, another component may
speed up to compensate. When one considers both the emergent properties and the
increasing homeostasis that develop at each level, not all component parts must be
known be- fore the whole can be understood. This is an important point, because
some contend that it is useless to try to work on complex populations and
communities when the smaller units are not yet fully understood. Quite the
contrary, one may begin study at any point in the spectrum, provided that adjacent
levels, as well as the level in question, are considered, because, as already noted,
some attributes are predictable from parts (collective properties), but others are not
(emergent properties). Ideally, a system-level study is itself a threefold hierarchy:
system, subsystem (next level below), and suprasystem(next level above). for more
on emergent properties, see T. F. H. Allen and Starr (1982), T. F. H. Allen and
Hoekstra (1992), and Ahl and Allen (1996).
Each biosystem level has emergent properties and reduced variance as well as a
summation of attributes of its subsystem components. The folk wisdom about the
forest being more than just a collection of trees is, indeed, a first working principle
of ecology. Although the philosophy of science has always been holistic in seeking
to understand phenomena as a whole, in recent years the practice of science has
become increasingly reductionist in seeking to understand phenomena by detailed
study of smaller and smaller components. Laszlo and Margenau (1972) described
within the history of science an alternation of reductionist and holistic thinking
(reduciionism- consiructionism and atomism-holism are other pairs of words used
to contrast these philosophical approaches). The law of diminishing returns may
very well be involved here, as excessive efort in any one direction eventually
necessitates taking the other (or another) direction.
The reductionist approach that has dominated science and technology since Isaac
 SECTION 4 T¥ansoending Functlona and Control Procescas 9

Newton has.made major contribuöons. For example,-research at the cellular and mo-
lecular levels has established a firm basis for the future cure and prevention of
can-
as cers at the level of the organism. However, cell-level science will contribute very little
tö the well-being or survival of human civilisation if we understand the higher leviels
ÄO of organisation so inadequately that we ,can find no solptioris to population. over-
le growth, pollution, and other forms,of soeietal and environmental.disorders. Both ho-
ly lism and reductionü m must be accorded' equal value—and simultaneously, not al-
or ternatively (E. P. Odum 1977; Barrett 1994). Ecology seeks.synthesis, not separation.
ge The revival of the holistic disciplines may be due at least partly to cîtizen dissatisfac-
. is tion with 'the specialized scientist who cannot respond to. the large-scale problems
ue that need urgent attention. (Historian Lynn White’s 1980 essay “The Ecology of Our
Science" is tecommended reading on this viewpoint.) Accordin@ty, we shall discuss
he ecolÔ gical principles at the ecosystëro level, with appiopriate attention to organism,
e. population, and community sub.sets and to landscape, biome, and ecosphere supra-
sets. This is the philosophieal basis for the organisation of the chapters in this
book. Fortunately, in the past 10 years, technological advances have allowed
humans to
ds deal qü antita'tively with large, complex systems such as ecosystems and landscapes.
nd Tracer methodology, mass chemiscy (spectrometry, colorimetry, chromatography).
ii- remote sensing, automatic monitoring, mathematic modeling, geographical informa-
tion systems (GIS), and computer technology are providing the tools. Technology is,
of coursei a double-edged sword; it can be the means of understanding the
he whole- ness of humans.and nature or of destroying it.

ng 4
ad
he Whereas each level in the ecological hierarchy can be expected to have unique
dy
.emer- gent and eollecü ve properties, there are basic functions that operate at all
levels. Ex- amples of such transcending functions are behavior, development,
diversity, ener- getics, evolution, integration, and regulation tsee Fig. 1-3 for
details). Sonne of these (energetics. for example) operate the same throughout the
,),
hierarchy, but others dif- fer in mddus operondi at difeient lexels. Natural selection
en evoluü on, for ,example, in- volves mutations and other direct.genetic.interactions at
the organism level but indi- rect. coevoluü onary and.group selection
› processes at higher levels:
a lt is especially important to emphasize that although p.ositive and negative
he Feed- back controls are universal, from the organism.down,. control..is set point,. in
Ge that it in- volves very. exacting genetic, hormonal, and neural controls on growth
and develop-
ne .ment. 'leading to what is often called homeostasis. As noted on the right-hand
of side of Figure 1-4, there aTe no.set-point controls above the orgariism level
ne (no.chemostats or thermostats.in nature)..According1y, feedback control is much
looser, resulting in pulsing rather than steady states.. The .term homeorhesis,. from
the Greek meaning “maintaining the ßow,‘ has been suggesied for this pulsing
control. lu other words, there are no equilibriums at the ecosysterri and ecosphere
levels, but there are pulsing 'balances, such as.between.productipn and respü ation
nT between oxygen and caibon dioxide in the,atmosphere. Failure to recognize this
dißerence in ‹:ybernetics (the sci- enee.dealing with mechanisms of control or
regul
ation
) has
resul
ted
in
muc
h
conf
u-
sion.
abo
ut
the.r
ealit
ies
ofth
e so-
calle
d
“bal
ance
of
natu
re.’
JO CHAPTER 1 The Scope of Ecology

5 Ecological Interfacing
Because ecology is a broad, multilevel discipline, it interfaces well with traditional
disciplines that tend to have more narrow focus. During the past decade, there has
been a rapid rise of interface fields of study accompanied by new societies,
journals, symposium volumes, books—and new careers. Ecological economics,
one of the most important, was mentioned in the first section in this chapter.
Others that are re- ceiving a great deal of attention, especially in resource
management, are agroecology, biodiversity, conservation ecology, ecological
engineering, ecosystem health, ecotox- icology, environmental ethics, and
restoration ecology.
In the beginning, an interface effort enriches the disciplines being
interfaced. Lines of communication are established, and the expertise of narrowly
trained “ex- perts” in each field is expanded. However, for an interface field to
become a new dis- cipline, something new has to emerge, such as a new concept or
technology. The con- cept of nonmarket goods and services, for example, was a
new concept that emerged in ecological economics, but that initially neither
traditional ecologists nor econo- mists would put in their textbooks (Daily 1997;
Mooney and Ehrlich 1997).
Throughout this book, we will refer to natural capital and economic capital.
Nat- ural capital is defined as the benefits and services supplied to human
societies by natural ecosystems, or provided “free of cost” by unmanaged natural
systems. These benefits and services include purification of water and air by
natural processes, de- composition of wastes, maintenance of biodiveisii y, control
of insect pests, pollina- tion of crops, mitigation of floods, and provision of
natural beauty and recreation, among others (Daily 1997),
Economic capital is defined as the goods and services provided by
humankind, or the human workforce, typically expressed as the gross national
product (GNP). Gross national product is the total monetary value of all goods
and services pro- vided in a country during one year. Natural capital is
typically quantified and ex- pressed in units of energy, whereas economic capital
is expressed in monetary units (Table 1-1). Only in recent years has there been
an attempt to value the world's ecosystem services and natural capital in
monetary terms. Costanza, d'Arge, et al. (1997) estimated this value to be in the
range of 16 to 54 trillion U.S. dollars per year for the entire biosphere, with an
average of 33 trillion U.S. dollars per year. Thus it is wise to protect natural
ecosystems, both ecologically and economically, because of the benefits and
services they provide to human societies, as will be illustrated in the chapters that
follow.

6 About Models
11 ecology is to be discussed at the ecosystem level, for reasons already indicated,
how can this complex and formidable system level be dealt with? We begin by
describing simplified versions that encompass only the most important, or basic,
properties and functions. Because, in science, simplified versions of the real world
are called modeis, it is appropriate now to introduce this concept.
A model (by definition) is a formulation that mimics a real-world phenomenon
 SECTION 6 About Models 11

and by which predictions can be made. In their simplest form, models may be
verbal or graphic (inJormal). Ultimately, however, models must be statistical and
mathe- matical (Jorrnal) if their quantitative predictions are to be reasonably
good. For ex-
al ample, a mathematical formulation that mimics numerical changes in a population
35
of insects and that predicts the numbers in the population at some time would be
considered a biologically useful model. lf the insect population in question is a pest
ie species, the model could have an economically imporiant application.
e- Computer-simulated models permit one to predict probable outcomes as param-
eters in the model are changed, as new parameters are added, or as old ones
are re- moved. Thus, a mathematical formulation can often be “tuned” or
refined by com- puter operations to improve the “fit” to the real-world
phenomenon. Above all,
d. models summarize what is understood about the situation modeled and thereby de-
limit aspects needing new or better data, or new principles. When a model does not work—
when it poorly mimics the real world—computer operations can often pro-
n- vide clues to the re hnements or changes needed. Once a model proves to be a useful
td mimic, opportunities for experimentation are unlimited, because one can introduce
o- new factors or perturbations and see how they would alTeci the system. Even
when a model inadequately mimics the real world, which is often the case in its
early stages
it- of development, it remains an exceedingly useful teaching and research tool if it re-
by veals key components and interactions that merit special attention.
Contrary to the feeling of many who are skeptical about modeling the complex-
Ie- ity of nature, information about only a relatively small number o[ variables is often a
sufficient basis for effective models because key factors, or emergent and other inte-
›n grative properties, as discussed in Sections 2 and 3, often dominate or control a large
,
percentage of the action. Watt (1963), for example, stated, “We do not need a
id, tremen- dous amount of information about a great many variables to build
PJ revealing mathe- matical models.” Though the mathematical aspects of modeling
’o- are a subject for ad- vanced texts, we should review the first steps in model
building.
d's Modeling usually begins with the construction of a diagram, or “graphic
al. model,” which is often a box or compartment diagram, as illustrated in Figure 1-5.
*ar Shown are two properties, and Pj, that interact, 1, to produce or affect a third
property , 3, when the system is driven by an energy source, E. Five Flow
: is
of pathways, F, are shown,
.he with Ft representing the input and F the output for the system as a whole.
Thus, at a minimum, there are five ingredients or components for a working model
of an eco- logical situation, namely, (1) an energy source or other outside forcing
function, E;
(2) properties called state variables, P , Pt, . . . P ; (3) ßow pathways, F l , F y, ..
. F„ showing where energy flows or material transfers connect properties with
each other and with forces; (4) interaction functions, I, where forces and
properties internet to modify, amplify, or control fiows or create new “emergent”
properties; and (5) feed- back loops, L.
ow
Figure 1-5 could serve as a model for the producrion of photochemical
ing
smog in the air over Los Angeles. In this case, Pt could represent hydrocarbons and
.nü P nitro- gen oxides, two producu of automobile exhaust emission. Under the
ers,
driving force of sunlight energy, E, these interact to produce photochemical
smog, Pj. In this case, the interaction function, I, is a synergistic or
ton
a humans than is P or P acting alone.
u Alternatively, Figure 1-5 could depict a grassland ecosystem in which
g repre- sents the green plants that convert the energy of the Sun, E, to food. P
m might repre- sent a herbivorous animal that eats plants, and Pj an omnivorous
e animal that can eat
n
t
a
t
i
v
e

o
n
e
,

i
n

t
h
a
t

i
s

m
o
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e

s
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 12 CHAPTER The Scope of Ecology

6i@U@ 1-5. Compartment diagram showing the five basic components of primary interest
in modeling ecological sy8terns. E = energy souce (foming function)› P„ P„ P# = 8tate
variables; F -Ft = flow pathwayei I interaction function; L - feedback loop.

either the herbivores or the plants. In this case, the interaction function, 1, could
rep- resent several possibilities. It could be a no-preference switch if observation in
the real world showed that the omnivore P eats either Pt or P , according to
availability. Or I could be specified to be a constant percentage value if it was
found that the diet of P was composed of, say, 80 percent plant and 20 percent
animal matter, irre- spective of the state of r P . Or 1 could be a seasonal switch
if P feeds on plants during one part of the year and on animals during another
season. Or I could be a threshold switch if P greatly prefers animal food and
switches to plants only when
* reduced to a low level.
Feedboch loops are imponant features of ecological models because they repre-
sent control mechanisms. Figure 1-6 is a simplified diagram of a system that
features a feedback loop in which “downstream” output, or some pan of it, is
ted back or re- cycled to a8eci or perhaps control "upstream” components. For
example, the feed- back loop could represent predation by “downstream"
organisms, C, that reduce and thereby tend to control the growth of "upstream"
herbivores or plants B and A in the food chain. Often, such feedback actually
promotes the growth or survival of a downstream component, such as a grazer
enluncing the growth of plants (a “reward feedback," as it were).

gure 1 Compartment model with a feedback or con-

trol loop that ransforms a linear system into a partiaI|y cycli-

Feedback loop
 SECTION 6 About Models 13

Figure t•7. Interaction of positive

and negative feedbacks in the Atmospheric COC Climatic
relation- ships of atmospheric CO , concentration warming
climate warming, soil respiration, and
carbon se- questration (modified
after Luo et at. 2001).
Carbon Photosynthesis Acclimatization
sequestration of soil espi ation

Vegetation Nutrient
groMh availability

"””“" -“““’*
” Respiration —+ Positive
” - - + Negative

Figure 1-6 could also represent a desirable economic system in which

resources, A, are converted into useful goods and services, B, with the production
of wastes, C, that are recycled and used again in the conversion process (A —+ B),
thus reducing the waste output of the system. By and large, natural ecosystems
have a circular or loop design rather than a linear structure. Feedback and
cybernetics, the science of
p. controls, are discussed in detail in Chapter 2.
he Figure 1-7 illustrates how positive and negative feedback can interact in the rela-
tionship between atmospheric CO concentration and climatic warming. An
increase in CO, has a positive greenhouse effect on global warming and on
plant growth. However, the soil system acclimates to the warming, so soil
respiration does not con-
ZI.S
tinue to increase with warming. This acclimation results in a negative feedback on
carbon sequestration in the soil, thus reducing emission of CO to the atmosphere,
en according to a study by Luo et al. (2001).
Compartment models are greatly enhanced by making the shape of the "boxes™
re- indicate the general function of the unit. In Figure 1-8, some of the symbols from
‘€
5 the H. T. Odum energy language (H. T. Odum and E. P. Odum 1982 ; H. T. Odum
1996) are depicted as used in this book. In Figure 1-9, these symbok are used in a
d- model of a pine forest located in Florida. Also, in this diagram estimates of the
ad amount of energy flow through the units are shown as indicators of the relative im-
he portance of unit functions.
'a In summary, good model definition should include three dimensions: (1) the
rd space to be considered (how the system is bounded); (2) the subsystems (compo-
nents) judged to be important in overall function; and (3) the time interval to be
con-
sidered. Once an ecosystem, ecological situation, or problem has been properly
defined and bounded, a testable hypothesis or series of hypotheses is developed
that can be rejected or accepted, at least tentatively, pending further
experimentation or analysis. For more on ecological modeling, see Patten and
jergensen (1995), H. T. Odum and E. C. Odum (2000), and Gunderson and
Holling (2002).
In the following chapters, the paragraphs headed by the word statement are, in
eifect, “word" models of the ecological principle in question. In many cases, graphic
models are also presented, and in some cases, simplified mathematical
formula- tions are included. Most of all, this book attempts to provide the
principles, concepts,
1ä CHAPTER \ The Scope of Eœlogy

Energy
circuit (A Consumer
pathway or (Usgs producer energy
flow of for self-maintenance)
energy)

Storage
Energy source (A compartment of
(Source of energy energy storage)
from
outside the system)

Interaction
(Two or more
flows
Heat sink of energy to produce
(Degraded energy a high-quality
after use in work) energy)

Producer Capital transaction

(Conver!s and concentrates (Flow of money to
solar energy) pay for flow of energy)

Figure 1-8. The H. T. Odum energy langu ge.symbole used in model diagrame in this book.

Rain

1 Runo#

Soil 1
WBt8F 2
organics
nütrlënts

2
Trees & Animals
other piants 12 2
0.1
1
Litter 2.9
2,000 12,986 1000 11 Micro-
Respiration
organisms
1.9

Figure t•9. Ecoayetem model Laing T. Odum).

energy language symbole arid including
estimated rated of energy fiow'for
a’Nor- ida pine forest (coarteey of H. 1@ joules/m2/day
Heat. sink (used energy)
 SECTION 7 Disóplinary Rediictiqriism.to Tranediściplinary Holism 15

simplifications, and abstractions that one must deduce from the real world before
one can understand and deal with situations and problems or construct
mathema.tical mo.dels of them.

In a paper entitled "The Emergence or Ecology as a New Integrative Discipline,”

E. P. Odum (1977) noted that ecology had become a new holistic discipline,.
having roots in the biological, physical, and spcial sciences, rather than just a
subdisclpliné of biology.. Thus, a goal of ecology is to link the natural and social
sciences. It should be noted that rrtost disciplines and .disciplinary approaches
are based on increased specialization in isolation (Fig. 1-10). The early evolution
and development of ecol-

f°•igure 1•10• Progression

DISCIPLINARY
of relations am.ong specializing in isolation
disciplines from discipli/iary
rédustlonism to
transdisciplinary hpliâm (af-
ter Jantsch 19I2). czi czi cz MULTIDISCIPLINARY
no cooperation

CRO56DTSCIPLINARY
rigid poJarization toward
specific monodisciplinar.y
CCINCOQ(

INTERDISCIPLINARY
coordination by higher-
level concept

TRANSDI6CłPLlNARY
multi-level coordińation of
en1ire education / irinovation
system
16 CHAPTER 1 The Scope of Ecology

ogy was frequently based on multidisciplinary approaches (multi = “many"), espe-

cially during the 1960s and 1970s. Unfortunately, the multidisciplinary approaches
lacked cooperation or focus. To achieve cooperation and define goals, institutes or
centers were established on campuses throughout the world, such as the Institute of
Ecology located on the campus of the University of Georgia. These
crossdisciplinary approaches (cross — “traverse"; Fig. 1-10) frequently resulted in
polarization toward a specific monodisciplinary concept, a poorly funded
administrative unit, or a nar- row mission. A crossdisciplinary approach also
frequently resulted in polarized fac- ulty reward systems. Institutions of higher
learning, traditionally built on disciplinary structures, have difficulties in
administering programs and addressing environmen- tal problems as well as taking
advantage of opportunities at greater temporal and spa- tial scales.
To address the dilemma, interdisciplinary approaches (inter = "among”) were
employed, resulting in cooperation on a higher-level concept, problem, or question.
For example, the process and study of natural ecological succession provided a
higher-level concept resulting in the success of the Savannah River Ecological
Labo- ratory (SREL) during its conception. Researchers theorized that new system
proper- ties emerge during the course of ecosystem development and that it is these
proper- ties that largely account for species and growth form changes that occur (E.
P. Odum 1969, 1977; see Chapter 8 for details). Today, interdisciplinary
approaches are com- mon when addressing problems at ecosystem, landscape, and
global levels.
Much remains to be done, however. There is an increased need to solve
problems, promote environmental literacy, and manage resources in a
transdisciplinary man- ner. This multilevel, large-scale approach involves entire
education and innovation systems (Fig. I -10). This integrative approach to the need
for unlocking cause-and- effect explanations across and among disciplines
(achieving a transdisciplinary un- derstanding) has been termed consilimce (E. O.
Wilson 1998), susiainabilit y science (dates et al. 2001), and integrative science
(Barrett 2001). Actually, the continued de- velopment of the science of ecology (the
“study of the household” or “place where we live”) will likely evolve into that
much-needed integrative science of the future. This book attempts to provide the
knowledge, concepts, principles, and approaches to underpin this educational need
and learning process.