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Principles of Terrestrial

Ecosystem Ecology

F. Stuart Chapin III


Pamela A. Matson
Harold A. Mooney

Springer
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1
The Ecosystem Concept

Ecosystem ecology studies the links between organisms and their physical environ-
ment within an Earth System context. This chapter provides background on the con-
ceptual framework and history of ecosystem ecology.

Introduction The supply of fish from the sea is now declin-


ing because fisheries management depended on
Ecosystem ecology addresses the interactions species-based approaches that did not ade-
between organisms and their environment as an quately consider the resources on which com-
integrated system. The ecosystem approach is mercial fish depend. A more holistic view of
fundamental in managing Earth’s resources managed systems can account for the complex
because it addresses the interactions that link interactions that prevail in even the simplest
biotic systems, of which humans are an integral ecosystems. There is also an increasing appreci-
part, with the physical systems on which they ation that a thorough understanding of eco-
depend. This applies at the scale of Earth as a systems is critical to managing the quality and
whole, a continent, or a farmer’s field. An quantity of our water supplies and in regulating
ecosystem approach is critical to resource man- the composition of the atmosphere that deter-
agement, as we grapple with the sustainable use mines Earth’s climate.
of resources in an era of increasing human
population and consumption and large, rapid
changes in the global environment. Overview of Ecosystem Ecology
Our growing dependence on ecosystem con-
cepts can be seen in many areas. The United The flow of energy and materials through
Nations Convention on Biodiversity of 1992, organisms and the physical environment pro-
for example, promoted an ecosystem approach, vides a framework for understanding the diver-
including humans, to conserving biodiversity sity of form and functioning of Earth’s physical
rather than the more species-based approaches and biological processes. Why do tropical
that predominated previously. There is a grow- forests have large trees but accumulate only a
ing appreciation of the role that individual thin layer of dead leaves on the soil surface,
species, or groups of species, play in the func- whereas tundra supports small plants but an
tioning of ecosystems and how these functions abundance of soil organic matter? Why does
provide services that are vital to human the concentration of carbon dioxide in the
welfare. An important, and belated, shift in atmosphere decrease in summer and increase
thinking has occurred about managing ecosys- in winter? What happens to that portion of the
tems on which we depend for food and fiber. nitrogen that is added to farmers’ fields but is

3
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4 1. The Ecosystem Concept

not harvested with the crop? Why has the intro- tion of plants by herbivores, and the consump-
duction of exotic species so strongly affected tion of herbivores by predators. Most of these
the productivity and fire frequency of grass- fluxes are sensitive to environmental factors,
lands and forests? Why does the number of such as temperature and moisture, and to bio-
people on Earth correlate so strongly with the logical factors that regulate the population
concentration of methane in the Antarctic dynamics and species interactions in communi-
ice cap or with the quantity of nitrogen enter- ties. The unique contribution of ecosystem
ing Earth’s oceans? These are representative ecology is its focus on biotic and abiotic factors
questions addressed by ecosystem ecology. as interacting components of a single integrated
Answers to these questions require an under- system.
standing of the interactions between organisms Ecosystem processes can be studied at many
and their physical environments—both the spatial scales. How big is an ecosystem? The
response of organisms to environment and appropriate scale of study depends on the ques-
the effects of organisms on their environment. tion being asked (Fig. 1.1). The impact of zoo-
Addressing these questions also requires plankton on the algae that they eat might be
that we think of integrated ecological systems studied in the laboratory in small bottles. Other
rather than individual organisms or physical questions such as the controls over productiv-
components. ity might be studied in relatively homogeneous
Ecosystem analysis seeks to understand the patches of a lake, forest, or agricultural field.
factors that regulate the pools (quantities) and Still other questions are best addressed at the
fluxes (flows) of materials and energy through global scale. The concentration of atmospheric
ecological systems. These materials include CO2, for example, depends on global patterns
carbon, water, nitrogen, rock-derived minerals of biotic exchanges of CO2 and the burning of
such as phosphorus, and novel chemicals such fossil fuels, which are spatially variable across
as pesticides or radionuclides that people have the globe. The rapid mixing of CO2 in the
added to the environment. These materials are atmosphere averages across this variability,
found in abiotic (nonbiological) pools such as facilitating estimates of long-term changes in
soils, rocks, water, and the atmosphere and in the total global flux of carbon between Earth
biotic pools such as plants, animals, and soil and the atmosphere.
microorganisms. Some questions require careful measure-
An ecosystem consists of all the organisms ments of lateral transfers of materials. A water-
and the abiotic pools with which they interact. shed is a logical unit in which to study the
Ecosystem processes are the transfers of energy effects of forests on the quantity and quality of
and materials from one pool to another. Energy the water that supplies a town reservoir. A
enters an ecosystem when light energy drives watershed, or catchment, consists of a stream
the reduction of carbon dioxide (CO2) to form and all the terrestrial surfaces that drain into
sugars during photosynthesis. Organic matter it. By studying a watershed we can compare the
and energy are tightly linked as they move quantities of materials that enter from the
through ecosystems. The energy is lost from air and rocks with the amounts that leave in
the ecosystem when organic matter is oxidized stream water, just as you balance your check-
back to CO2 by combustion or by the respira- book. Studies of input–output budgets of water-
tion of plants, animals, and microbes. Materials sheds have improved our understanding of the
move among abiotic components of the system interactions between rock weathering, which
through a variety of processes, including the supplies nutrients, and plant and microbial
weathering of rocks, the evaporation of water, growth, which retains nutrients in ecosystems
and the dissolution of materials in water. (Vitousek and Reiners 1975, Bormann and
Fluxes involving biotic components include the Likens 1979).
absorption of minerals by plants, the death of The upper and lower boundaries of an
plants and animals, the decomposition of dead ecosystem also depend on the question being
organic matter by soil microbes, the consump- asked and the scale that is appropriate to the
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Overview of Ecosystem Ecology 5

Figure 1.1. Examples of a) Global ecosystem


ecosystems that range in size
by 10 orders of magnitude:
an endolithic ecosystem in
the surface layers of rocks, 5,000 km
How does carbon loss
1 ¥ 10-3 m in height (d); a
from plowed soils
forest, 1 ¥ 103 m in diameter influence global climate?
(c); a watershed, 1 ¥ 105 m in
length (b); and Earth, 4 ¥ 107 m
in circumference (a). Also
shown are examples of ques- b) Watershed
tions appropriate to each scale.

10 km
How does
deforestation
influence the
water supply to
neighboring towns?

c) Forest ecosystem

1 km
How does acid rain
influence forest
productivity?

d) Endolithic ecosystem
rock surface

lichen zone What are the biological


1 mm
controls over rock
algal zone weathering?

question. The atmosphere, for example, extends questions that address plant effects on water
from the gases between soil particles all the way and nutrient cycling, the bottom of the ecosys-
to outer space. The exchange of CO2 between a tem might be the maximum depth to which
forest and the atmosphere might be measured roots extend because soil water or nutrients
a few meters above the top of the canopy below this depth are inaccessible to the vegeta-
because, above this height, variations in CO2 tion. Studies of long-term soil development, in
content of the atmosphere are also strongly contrast, must also consider rocks deep in the
influenced by other upwind ecosystems. The soil, which constitute the long-term reservoir of
regional impact of grasslands on the moisture many nutrients that gradually become incorpo-
content of the atmosphere might, however, be rated into surface soils (see Chapter 3).
measured at a height of several kilometers Ecosystem dynamics are a product of many
above the ground surface, where the moisture temporal scales. The rates of ecosystem pro-
released by the ecosystem condenses and cesses are constantly changing due to fluctua-
returns as precipitation (see Chapter 2). For tions in environment and activities of organisms
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6 1. The Ecosystem Concept

on time scales ranging from microseconds to have a pervasive influence. The complications
millions of years (see Chapter 13). Light capture associated with the current nonequilibrium
during photosynthesis responds almost instan- view require a more dynamic and stochastic
taneously to fluctuations in light availability view of controls over ecosystem processes.
to a leaf. At the opposite extreme, the evolution Ecosystems are considered to be at steady
of photosynthesis 2 billion years ago added state if the balance between inputs and outputs
oxygen to the atmosphere over millions of to the system shows no trend with time
years, causing the prevailing geochemistry (Johnson 1971, Bormann and Likens 1979).
of Earth’s surface to change from chemical Steady state assumptions differ from equilib-
reduction to chemical oxidation (Schlesinger rium assumptions because they accept tempo-
1997). Microorganisms in the group Archaea ral and spatial variation as a normal aspect of
evolved in the early reducing atmosphere of ecosystem dynamics. Even at steady state, for
Earth. These microbes are still the only organ- example, plant growth changes from summer to
isms that produce methane. They now function winter and between wet and dry years (see
in anaerobic environments such as wetland soils Chapter 6). At a stand scale, some plants may
and the interiors of soil aggregates or animal die from old age or pathogen attack and be
intestines. Episodes of mountain building and replaced by younger individuals. At a landscape
erosion strongly influence the availability of scale, some patches may be altered by fire or
minerals to support plant growth. Vegetation is other disturbances, and other patches will be
still migrating in response to the retreat of Pleis- in various stages of recovery. These ecosystems
tocene glaciers 10,000 to 20,000 years ago. After or landscapes are in steady state if there is
disturbances such as fire or tree fall, there are no long-term directional trend in their pro-
gradual changes in plant, animal, and microbial perties or in the balance between inputs and
communities over years to centuries. Rates of outputs.
carbon input to an ecosystem through photo- Not all ecosystems and landscapes are in
synthesis change over time scales of seconds to steady state. In fact, directional changes in
decades due to variations in light, temperature, climate and environment caused by human
and leaf area. activities are quite likely to cause directional
Many early studies in ecosystem ecology changes in ecosystem properties. Nonetheless,
made the simplifying assumption that some it is often easier to understand the relationship
ecosystems are in equilibrium with their envi- of ecosystem processes to the current environ-
ronment. In this perspective, relatively undis- ment in situations in which they are not also
turbed ecosystems were thought to have recovering from large recent perturbations.
properties that reflected (1) largely closed Once we understand the behavior of a system
systems dominated by internal recycling of in the absence of recent disturbances, we can
elements, (2) self-regulation and deterministic add the complexities associated with time lags
dynamics, (3) stable end points or cycles, and and rates of ecosystem change.
(4) absence of disturbance and human influ- Ecosystem ecology uses concepts developed
ence (Pickett et al. 1994, Turner et al. 2001). at finer levels of resolution to build an under-
One of the most important conceptual standing of the mechanisms that govern the
advances in ecosystem ecology has been the entire Earth System. The biologically mediated
increasing recognition of the importance of movement of carbon and nitrogen through
past events and external forces in shaping ecosystems depends on the physiological
the functioning of ecosystems. In this non- properties of plants, animals, and soil micro-
equilibrium perspective, we recognize that organisms. The traits of these organisms are
most ecosystems exhibit inputs and losses, their the products of their evolutionary histories
dynamics are influenced by both external and and the competitive interactions that sort
internal factors, they exhibit no single stable species into communities where they success-
equilibrium, disturbance is a natural compo- fully grow, survive, and reproduce (Vrba and
nent of their dynamics, and human activities Gould 1986). Ecosystem fluxes also depend
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History of Ecosystem Ecology 7

Earth system Context ecosystem ecologists about the rates at which


science
Climatology the land or water surface interacts with the
atmosphere, rocks, and waters of the planet
Hydrology (Fig. 1.2). Conversely, the global budgets of
materials that cycle between the atmosphere,
Ecosystem ecology land, and oceans provide a context for under-
standing the broader significance of processes
Community
Soil science studied in a particular ecosystem. Latitudinal
ecology
and seasonal patterns of atmospheric CO2 con-
centration, for example, help define the loca-
Population Geochemistry
ecology
tions where carbon is absorbed or released
from the land and oceans (see Chapter 15).
Physiological
ecology Mechanism

Figure 1.2. Relationships between ecosystem History of Ecosystem Ecology


ecology and other disciplines. Ecosystem ecology
integrates the principles of several biological and Many early discoveries of biology were moti-
physical disciplines and provides the mechanistic vated by questions about the integrated nature
basis for Earth System Science. of ecological systems. In the seventeenth
century, European scientists were still uncertain
about the source of materials found in plants.
on the population processes that govern Plattes, Hooke, and others advanced the novel
plant, animal, and microbial densities and idea that plants derive nourishment from
age structures as well as on community both air and water (Gorham 1991). Priestley
processes, such as competition and predation, extended this idea in the eighteenth century by
that determine which species are present and showing that plants produce a substance that is
their rates of resource consumption. Ecosystem essential to support the breathing of animals.At
ecology therefore depends on information about the same time MacBride and Priestley
and principles developed in physiological, evo- showed that breakdown of organic matter
lutionary, population, and community ecology caused the production of “fixed air” (carbon
(Fig. 1.2). dioxide), which did not support animal life.
The supply of water and minerals from soils De Saussure, Liebig, and others clarified the
to plants depends not only on the activities of explicit roles of carbon dioxide, oxygen,
soil microorganisms but also on physical and and mineral nutrients in these cycles in the
chemical interactions among rocks, soils, and nineteenth century. Much of the biological
the atmosphere. The low availability of phos- research during the nineteenth and twentieth
phorus due to the extensive weathering and centuries went on to explore the detailed
erosional loss of nutrients in the ancient soils of mechanisms of biochemistry, physiology,
western Australia, for example, strongly con- behavior, and evolution that explain how life
strains plant growth and the quantity and types functions. Only in recent decades have we
of plants and animals that can be supported. returned to the question that originally moti-
Principles of ecosystem ecology must therefore vated this research: How are biogeochemical
also incorporate the concepts and understand- processes integrated in the functioning of
ing of disciplines such as geochemistry, hydrol- natural ecosystems?
ogy, and climatology that focus on the physical Many threads of ecological thought have
environment (Fig. 1.2). contributed to the development of ecosystem
Ecosystem ecology provides the mechanistic ecology (Hagen 1992), including ideas relating
basis for understanding processes that occur to trophic interactions (the feeding relation-
at global scales. Study of Earth as a physical ships among organisms) and biogeochemistry
system relies on information provided by (biological influences on the chemical processes
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8 1. The Ecosystem Concept

in ecosystems). Early research on trophic inter- that there were insufficient data to draw such
actions emphasized the transfer of energy broad conclusions and that it was inappropriate
among organisms. Elton (1927), an English to use mathematical models to infer general
zoologist interested in natural history, relationships based on observations from a
described the role that an animal plays in a single lake. Hutchinson, Lindeman’s postdoc-
community (its niche) in terms of what it eats toral adviser, finally (after Lindeman’s death)
and is eaten by. He viewed each animal species persuaded the editor to publish this paper,
as a link in a food chain, which described the which has been the springboard for many of the
movement of matter from one organism to basic concepts in ecosystem theory (Lindeman
another. Elton’s concepts of trophic structure 1942).
provide a framework for understanding the H. T. Odum, also trained by Hutchinson,
flow of materials through ecosystems (see and his brother E. P. Odum further developed
Chapter 11). the systems approach to studying ecosystems,
Hutchinson, an American limnologist, was which emphasizes the general properties of
strongly influenced by the ideas of Elton and ecosystems without documenting all the under-
those of Russian geochemist Vernadsky, who lying mechanisms and interactions. The Odum
described the movement of minerals from soil brothers used radioactive tracers to measure
into vegetation and back to soil. Hutchinson the movement of energy and materials through
suggested that the resources available in a lake a coral reef. These studies enabled them to doc-
must limit the productivity of algae and that ument the patterns of energy flow and metab-
algal productivity, in turn, must limit the abun- olism of whole ecosystems and to suggest
dance of animals that eat algae. Meanwhile, generalizations about how ecosystems function
Tansley (1935), a British terrestrial plant ecolo- (Odum 1969). Ecosystem budgets of energy
gist, was also concerned that ecologists focused and materials have since been developed for
their studies so strongly on organisms that many fresh-water and terrestrial ecosystems
they failed to recognize the importance of (Lindeman 1942, Ovington 1962, Golley 1993),
exchange of materials between organisms and providing information that is essential for gen-
their abiotic environment. He coined the term eralizing about global patterns of processes
ecosystem to emphasize the importance of such as productivity. Some of the questions
interchanges of materials between inorganic addressed by systems ecology include informa-
and organic components as well as among tion transfer (Margalef 1968), the structure of
organisms. food webs (Polis 1991), the hierarchical changes
Lindeman, another limnologist, was strongly in ecosystem controls at different temporal
influenced by all these threads of ecological and spatial scales (O’Neill et al. 1986), and the
theory. He suggested that energy flow through resilience of ecosystem properties after distur-
an ecosystem could be used as a currency to bance (Holling 1986).
quantify the roles of organisms in trophic We now recognize that element cycles inter-
dynamics. Green plants (primary producers) act in important ways and cannot be under-
capture energy and transfer it to animals stood in isolation. The availability of water and
(consumers) and decomposers. At each trans- nitrogen are important determinants of the rate
fer, some energy is lost from the ecosystem at which carbon cycles through the ecosystem.
through respiration. Therefore, the productivity Conversely, the productivity of vegetation
of plants constrains the quantity of consumers strongly influences the cycling rates of nitrogen
that an ecosystem can support. The energy and water.
flow through an ecosystem maps closely to Recent global changes in the environment
carbon flow in the processes of photosynthesis, have made ecologists increasingly aware of the
trophic transfers, and respiratory release of changes in ecosystem processes that occur in
carbon. Lindeman’s dissertation research on response to disturbance or other environmen-
the trophic-dynamic aspect of ecology was ini- tal changes. Succession, the directional change
tially rejected for publication. Reviewers felt in ecosystem structure and functioning result-
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History of Ecosystem Ecology 9

ing from biotically driven changes in resource predictably with climate (Jenny 1941, Rodin
supply, is an important framework for under- and Bazilevich 1967, Lieth 1975). The studies
standing these transient dynamics of ecosys- also showed that, in a given climatic regime, the
tems. Early American ecologists such as Cowles properties of vegetation depended strongly on
and Clements were struck by the relatively pre- soils and vice versa (Dokuchaev 1879, Jenny
dictable patterns of vegetation development 1941, Ellenberg 1978). Process-based studies of
after exposure of unvegetated land surfaces. organisms and soils provided insight into many
Sand dunes on Lake Michigan, for example, are of the mechanisms underlying the distributions
initially colonized by drought-resistant herba- of organisms and soils along these gradients
ceous plants that give way to shrubs, then small (Billings and Mooney 1968, Mooney 1972,
trees, and eventually forests (Cowles 1899). Larcher 1995, Paul and Clark 1996). These
Clements (1916) advanced a theory of commu- studies also formed the basis for extrapolation
nity development, suggesting that this vegeta- of processes across complex landscapes to char-
tion succession is a predictable process that acterize large regions (Matson and Vitousek
eventually leads, in the absence of disturbance, 1987, Turner et al. 2001). These studies often
to a stable community type characteristic of a relied on field or laboratory experiments
particular climate (the climatic climax). He sug- that manipulated some ecosystem property or
gested that a community is like an organism process or on comparative studies across envi-
made of interacting parts (species) and that ronmental gradients. This approach was later
successional development toward a climax expanded to studies of intact ecosystems, using
community is analogous to the development whole-ecosystem manipulations (Likens et al.
of an organism to adulthood. This analogy 1977, Schindler 1985, Chapin et al. 1995) and
between an ecological community and an carefully designed gradient studies (Vitousek et
organism laid the groundwork for concepts of al. 1988).
ecosystem physiology (for example, the net Ecosystem experiments have provided both
ecosystem exchange of CO2 and water vapor basic understanding and information that are
between the ecosystem and the atmosphere). critical in management decisions. The clear-
The measurements of net ecosystem exchange cutting of an experimental watershed at
are still an active area of research in ecosystem Hubbard Brook in the northeastern United
ecology, although they are now motivated by States, for example, caused a fourfold increase
different questions than those posed by in streamflow and stream nitrate concentra-
Clements. His ideas were controversial from tion—to levels exceeding health standards
the outset. Other ecologists, such as Gleason for drinking water (Likens et al. 1977). These
(1926), felt that vegetation change was not as dramatic results demonstrate the key role of
predictable as Clements had implied. Instead, vegetation in regulating the cycling of water
chance dispersal events explained much of and nutrients in forests. The results halted plans
the vegetation patterns on the landscape. This for large-scale deforestation that had been
debate led to a century of research on the planned to increase supplies of drinking water
mechanisms responsible for vegetation change during a long-term drought. Nutrient addition
(see Chapter 13). experiments in the Experimental Lakes Area
Another general approach to ecosystem of southern Canada showed that phosphorus
ecology has emphasized the controls over limits the productivity of many lakes (Schindler
ecosystem processes through comparative 1985) and that pollution was responsible
studies of ecosystem components. This interest for algal blooms and fish kills that were
originated in studies by plant geographers and common in lakes near densely populated areas
soil scientists who described general patterns of in the 1960s. This research provided the basis
variation with respect to climate and geological for regulations that removed phosphorus from
substrate (Schimper 1898). These studies detergents.
showed that many of the global patterns of Changes in the Earth System have led to
plant production and soil development vary studies of the interactions among terrestrial
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10 1. The Ecosystem Concept

ecosystems, the atmosphere, and the oceans. oxidation of hydrogen sulfide (H2S) to produce
The dramatic impact of human activities on the organic matter. Decomposer microorganisms
Earth System (Vitousek 1994a) has led to the (microbes) break down dead organic material,
urgent necessity to understand how terrestrial releasing CO2 to the atmosphere and nutrients
ecosystem processes affect the atmosphere in forms that are available to other microbes
and oceans. The scale at which these ecosystem and plants. If there were no decomposition,
effects are occurring is so large that the large accumulations of dead organic matter
traditional tools of ecologists are insufficient. would sequester the nutrients required to
Satellite-based remote sensing of ecosystem support plant growth. Animals are critical com-
properties, global networks of atmospheric ponents of ecosystems because they transfer
sampling sites, and the development of global energy and materials and strongly influence the
models are important new tools that address quantity and activities of plants and soil
global issues. Information on global patterns of microbes. The essential abiotic components of
CO2 and pollutants in the atmosphere, for an ecosystem are water; the atmosphere, which
example, provide telltale evidence of the major supplies carbon and nitrogen; and soil minerals,
locations and causes of global problems (Tans which supply other nutrients required by
et al. 1990). This gives hints about which ecosys- organisms.
tems and processes have the greatest impact on An ecosystem model describes the major
the Earth System and therefore where research pools and fluxes in an ecosystem and the factors
and management should focus efforts to under- that regulate these fluxes. Nutrients, water, and
stand and solve these problems (Zimov et al. energy differ from one another in the relative
1999). importance of ecosystem inputs and outputs vs.
The intersection of systems approaches, internal recycling (see Chapters 4 to 10). Plants,
process understanding, and global analysis is an for example, acquire carbon primarily from the
exciting frontier of ecosystem ecology. How do atmosphere, and most carbon released by res-
changes in the global environment alter the piration returns to the atmosphere. Carbon
controls over ecosystem processes? What are cycling through ecosystems is therefore quite
the integrated system consequences of these open, with large inputs to, and losses from,
changes? How do these changes in ecosystem the system. There are, however, relatively large
properties influence the Earth System? The pools of carbon stored in ecosystems, so the
rapid changes that are occurring in ecosystems activities of animals and microbes are some-
have blurred any previous distinction between what buffered from variations in carbon up-
basic and applied research. There is an urgent take by plants. The water cycle of ecosystems
need to understand how and why the ecosys- is also relatively open, with water entering
tems of Earth are changing. primarily by precipitation and leaving by evap-
oration, transpiration, and drainage to ground-
water and streams. In contrast to carbon,
Ecosystem Structure most ecosystems have a limited capacity to
store water in plants and soil, so the activity of
Most ecosystems gain energy from the sun and organisms is closely linked to water inputs.
materials from the air or rocks, transfer these In contrast to carbon and water, mineral ele-
among components within the ecosystem, then ments such as nitrogen and phosphorus are
release energy and materials to the environ- recycled rather tightly within ecosystems, with
ment. The essential biological components of annual inputs and losses that are small relative
ecosystems are plants, animals, and decom- to the quantities that annually recycle within
posers. Plants capture solar energy in the the ecosystem. These differences in the “open-
process of bringing carbon into the ecosystem. ness” and “buffering” of the cycles fundamen-
A few ecosystems, such as deep-sea hydro- tally influence the controls over rates and
thermal vents, have no plants but instead patterns of the cycling of materials through
have bacteria that derive energy from the ecosystems.
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Controls over Ecosystem Processes 11

The pool sizes and rates of cycling differ variations in climate explain the distribution of
substantially among ecosystems (see Chapter biomes (types of ecosystems) such as wet trop-
6). Tropical forests have much larger pools ical forests, temperate grasslands, and arctic
of carbon and nutrients in plants than do tundra (see Chapter 2). Within each biome,
deserts or tundra. Peat bogs, in contrast, parent material strongly influences the types of
have large pools of soil carbon rather than soils that develop and explains much of the
plant carbon. Ecosystems also differ substan- regional variation in ecosystem processes (see
tially in annual fluxes of materials among Chapter 3). Topographic relief influences both
pools, for reasons that will be explored in later microclimate and soil development at a local
chapters. scale. The potential biota governs the types and
diversity of organisms that actually occupy
a site. Island ecosystems, for example, are
Controls over frequently less diverse than climatically similar
Ecosystem Processes mainland ecosystems because new species
reach islands less frequently and are more
Ecosystem structure and functioning are gov- likely to go extinct than in mainland locations
erned by at least five independent control (MacArthur and Wilson 1967). Time influences
variables. These state factors, as Jenny and the development of soil and the evolution
co-workers called them, are climate, parent of organisms over long time scales. Time also
material (i.e., the rocks that give rise to soils), incorporates the influences on ecosystem
topography, potential biota (i.e., the organisms processes of past disturbances and environ-
present in the region that could potentially mental changes over a wide range of time
occupy a site), and time (Fig. 1.3) (Jenny 1941, scales. These state factors are described in
Amundson and Jenny 1997). Together these more detail in Chapter 3 in the context of soil
five factors set the bounds for the characteris- development.
tics of an ecosystem. Jenny’s state factor approach was a major
On broad geographic scales, climate is the conceptual contribution to ecosystem ecology.
state factor that most strongly determines First, it emphasized the controls over processes
ecosystem processes and structure. Global rather than simply descriptions of patterns.
Second, it suggested an experimental approach
to test the importance and mode of action of
Climate Time each control. A logical way to study the role of
each state factor is to compare sites that are as
Disturbance
regime
similar as possible with respect to all but one
Human factor. For example, a chronosequence is a
Modulators activities series of sites of different ages with similar
Topo- Ecosystem climate, parent material, topography, and
graphy processes potential to be colonized by the same organ-
isms (see Chapter 13). In a toposequence,
Resources Biotic ecosystems differ mainly in their topographic
community
position (Shaver et al. 1991). Sites that differ
primarily with respect to climate or parent
Parent Potential material allow us to study the impact of these
material biota state factors on ecosystem processes (Vitousek
et al. 1988, Walker et al. 1998). Finally, a com-
Figure 1.3. The relationship between state factors
(outside the circle), interactive controls (inside the parison of ecosystems that differ primarily in
circle), and ecosystem processes. The circle repre- potential biota, such as the mediterranean
sents the boundary of the ecosystem. (Modified with shrublands that have developed on west coasts
permission from American Naturalist, Vol. 148 © of California, Chile, Portugal, South Africa, and
1996 University of Chicago Press, Chapin et al. 1996.) Australia, illustrates the importance of evolu-
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12 1. The Ecosystem Concept

tionary history in shaping ecosystem processes unlike resources, are neither consumed nor
(Mooney and Dunn 1970). depleted by organisms (Field et al. 1992). Mod-
Ecosystem processes both respond to and ulators include temperature, pH, redox state of
control the factors that directly govern their the soil, pollutants, UV radiation, etc. Modula-
activity. For example, plants both respond to tors like temperature are constrained by
and influence their light, temperature, and climate (a state factor) but are sensitive to
moisture environment (Billings 1952). Interac- ecosystem processes, such as shading and evap-
tive controls are factors that both control and oration. Soil pH likewise depends on parent
are controlled by ecosystem characteristics (Fig. material and time but also responds to vegeta-
1.3) (Chapin et al. 1996). Important interactive tion composition.
controls include the supply of resources to Landscape-scale disturbance by fire, wind,
support the growth and maintenance of organ- floods, insect outbreaks, and hurricanes is a crit-
isms, modulators that influence the rates of ical determinant of the natural structure and
ecosystem processes, disturbance regime, the process rates in ecosystems (Pickett and White
biotic community, and human activities. 1985, Sousa 1985). Like other interactive con-
Resources are the energy and materials in the trols, disturbance regime depends on both state
environment that are used by organisms to factors and ecosystem processes. Climate,
support their growth and maintenance (Field for example, directly affects fire probability
et al. 1992). The acquisition of resources by and spread but also influences the types and
organisms depletes their abundance in the quantity of plants present in an ecosystem
environment. In terrestrial ecosystems these and therefore the fuel load and flammability
resources are spatially separated, being avail- of vegetation. Deposition and erosion during
able primarily either aboveground (light and floods shape river channels and influence
CO2) or belowground (water and nutrients). the probability of future floods. Change in
Resource supply is governed by state factors either the intensity or frequency of disturbance
such as climate, parent material, and topogra- can cause long-term ecosystem change. Woody
phy. It is also sensitive to processes occurring plants, for example, often invade grasslands
within the ecosystem. Light availability, for when fire suppression reduces fire frequency.
example, depends on climatic elements such as The nature of the biotic community (i.e., the
cloudiness and on topographic position, but is types of species present, their relative abun-
also sensitive to the quantity of shading by dances, and the nature of their interactions)
vegetation. Similarly, soil fertility depends on can influence ecosystem processes just as
parent material and climate but is also sensitive strongly as do large differences in climate or
to ecosystem processes such as erosional loss of parent material (see Chapter 12). These species
soils after overgrazing and inputs of nitrogen effects can often be generalized at the level of
from invading nitrogen-fixing species. Soil functional types, which are groups of species
water availability strongly influences species that are similar in their role in community or
composition in dry climates. Soil water avail- ecosystem processes. Most evergreen trees, for
ability also depends on other interactive example, produce leaves that have low rates of
controls, such as disturbance regime (e.g., com- photosynthesis and a chemical composition
paction by animals) and the types of organisms that deters herbivores. These species make up a
that are present (e.g., the presence or absence of functional type because of their ecological sim-
deep-rooted trees such as mesquite that tap the ilarity to one another. A gain or loss of key
water table). In aquatic ecosystems, water functional types—for example, through intro-
seldom directly limits the activity of organisms, duction or removal of species with important
but light and nutrients are just as important ecosystem effects—can permanently change
as on land. Oxygen is a particularly critical the character of an ecosystem through changes
resource in aquatic ecosystems because of its in resource supply or disturbance regime.
slow rate of diffusion through water. Introduction of nitrogen-fixing trees onto
Modulators are physical and chemical prop- British mine wastes, for example, substantially
erties that affect the activity of organisms but, increases nitrogen supply and productivity
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Human-Caused Changes in Earth’s Ecosystems 13

and alters patterns of vegetation development vents an uncontrolled growth of a predator’s


(Bradshaw 1983). Invasion by exotic grasses population, thereby stabilizing the population
can alter fire frequency, resource supply, tro- sizes of both predator and prey. There are also
phic interactions, and rates of most ecosystem positive feedbacks in ecosystems in which both
processes (D’Antonio and Vitousek 1992). components of a system have a positive effect
Elimination of predators by hunting can cause on the other, or both have a negative effect on
an outbreak of deer that overbrowse their food one another. Plants, for example, provide their
supply. The types of species present in an mycorrhizal fungi with carbohydrates in return
ecosystem depend strongly on other interactive for nutrients. This exchange of growth-limiting
controls (see Chapter 12), so functional types resources between plants and fungi promotes
respond to and affect most interactive controls the growth of both components of the sym-
and ecosystem processes. biosis until they become constrained by other
Human activities have an increasing impact factors.
on virtually all the processes that govern ecosys- Negative feedbacks are the key to sustaining
tem properties (Vitousek 1994a). Our actions ecosystems because strong negative feedbacks
influence interactive controls such as water provide resistance to changes in interactive
availability, disturbance regime, and biotic controls and maintain the characteristics of
diversity. Humans have been a natural compo- ecosystems in their current state. The acquisi-
nent of many ecosystems for thousands of years. tion of water, nutrients, and light to support
Since the Industrial Revolution, however, the growth of one plant, for example, reduces avail-
magnitude of human impact has been so great ability of these resources to other plants,
and so distinct from that of other organisms that thereby constraining community productivity
the modern effects of human activities warrant (Fig. 1.4). Similarly, animal populations cannot
particular attention. The cumulative impact of sustain exponential population growth indefi-
human activities extend well beyond an individ- nitely, because declining food supply and
ual ecosystem and affect state factors such as increasing predation reduce the rate of popu-
climate, through changes in atmospheric com- lation increase. If these negative feedbacks
position, and potential biota, through the intro- are weak or absent (a low predation rate due
duction and extinction of species. The large to predator control, for example), population
magnitude of these effects blurs the distinction cycles can amplify and lead to extinction of one
between “independent” state factors and inter- or both of the interacting species. Community
active controls at regional and global scales. dynamics, which operate within a single eco-
Human activities are causing major changes in system patch, primarily involve feedbacks
the structure and functioning of all ecosystems, among soil resources and functional types of
resulting in novel conditions that lead to new organisms. Landscape dynamics, which govern
types of ecosystems. The major human effects changes in ecosystems through cycles of dis-
are summarized in the next section. turbance and recovery, involve additional
Feedbacks analogous to those in simple phys- feedbacks with microclimate and disturbance
ical systems regulate the internal dynamics of regime (see Chapter 14).
ecosystems. A thermostat is an example of a
simple physical feedback. It causes a furnace to
switch on when a house gets cold. The house Human-Caused Changes in
then warms until the thermostat switches the Earth’s Ecosystems
furnace off. Natural ecosystems are complex
networks of interacting feedbacks (DeAngelis Human activities transform the land surface,
and Post 1991). Negative feedbacks occur when add or remove species, and alter biogeochemi-
two components of a system have opposite cal cycles. Some human activities directly affect
effects on one another. Consumption of prey by ecosystems through activities such as resource
a predator, for example, has a positive effect on harvest, land use change, and management;
the consumer but a negative effect on the prey. other effects are indirect, as a result of changes
The negative effect of predators on prey pre- in atmospheric chemistry, hydrology, and
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14 1. The Ecosystem Concept

+ half of the world’s accessible runoff (see


Nature of Chapter 15), and humans use about 8% of the
Process
feedback primary production of the oceans (Pauly and
F
Resource uptake A - Christensen 1995). Commercial fishing reduces
Predator Competition A+B - the size and abundance of target species and
Mutualism C + alters the population characteristics of species
Herbivory D - that are incidentally caught in the fishery. In the
Predation E -
mid-1990s, about 22% of marine fisheries
+ E - Population growth F +
were overexploited or already depleted, and an
additional 44% were at their limit of exploita-
Herbivore Mycorrhizal tion (Vitousek et al. 1997c). About 60% of the
fungus human population resides within 100 km of a
+ D - + C +
coast, so the coastal margins of oceans are
strongly influenced by many human activities.
Plant A Plant B Nutrient enrichment of many coastal waters, for
example, has increased algal production and
+ A - - B + created anaerobic conditions that kill fish and
Shared resources
other animals, due largely to transport of nutri-
ents derived from agricultural fertilizers and
Figure 1.4. Examples of linked positive and nega- from human and livestock sewage.
tive feedbacks in ecosystems. The effect of each Land use change, and the resulting loss of
organism (or resource) on other organisms can be habitat, is the primary driving force causing
positive (+) or negative (-). Feedbacks are positive species extinctions and loss of biological diver-
when the reciprocal effects of each organism (or
sity (Sala et al. 2000a) (see Chapter 12). The
resource) have the same sign (both positive or both
time lag between ecosystem change and species
negative). Feedbacks are negative when reciprocal
effects differ in sign. Negative feedbacks resist the loss makes it likely that species will continue to
tendencies for ecosystems to change, whereas posi- be driven to extinction even where rates of land
tive feedbacks tend to push ecosystems toward a new use change have stabilized. Transport of species
state. (Modified with permission from American Nat- around the world is homogenizing Earth’s
uralist, Vol. 148 © 1996 University of Chicago Press, biota. The frequency of biological invasions
Chapin et al. 1996.) is increasing, due to the globalization of the
economy and increased international transport
climate (Fig. 1.5) (Vitousek et al. 1997c). At of products. Nonindigenous species now
least some of these anthropogenic (i.e., human- account for 20% or more of the plant species
caused) effects influence all ecosystems on in many continental areas and 50% or more of
Earth. the plant species on many islands (Vitousek
The most direct and substantial human alter- et al. 1997c). International commerce breaks
ation of ecosystems is through the transforma- down biogeographic barriers, through both
tion of land for production of food, fiber, and purposeful trade in live organisms and inad-
other goods used by people. About 50% of vertent introductions. Purposeful introduc-
Earth’s ice-free land surface has been directly tions deliberately select species that are likely
altered by human activities (Kates et al. 1990). to grow and reproduce effectively in their
Agricultural fields and urban areas cover 10 to new environment. Many biological invasions
15%, and pastures cover 6 to 8% of the land. are irreversible because it is difficult or
Even more land is used for forestry and grazing prohibitively expensive to remove invasive
systems. All except the most extreme environ- species. Some species invasions degrade
ments of Earth experience some form of direct human health or cause large economic losses.
human impact. Others alter the structure and functioning of
Human activities have also altered fresh- ecosystems, leading to further loss of species
water and marine ecosystems. We use about diversity.
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Human-Caused Changes in Earth’s Ecosystems 15

Figure 1.5. Direct and indirect


effects of human activities on Earth’s Human population
Size Resource use
ecosystems. (Redrawn with permis-
sion from Science, Vol. 277 © 1997
American Association for the Human enterprises
Advancement of Science; Vitousek Agriculture Industry Recreation International commerce
et al. 1997c.)

Land Biotic additions


transformation and losses
Land clearing Invasion
Intensification Global Hunting
Forestry biochemistry Fishing
Grazing
Water
Carbon
Nitrogen
Other elements
Synthetic chemicals
Radionuclides

Climate change Loss of biological


diversity
Enhanced
greenhouse effect Extinction of species
Aerosols and populations
Land cover Loss of ecosystems

Human activities have influenced biogeo- less anthropogenic gases have had drastic
chemical cycles in many ways. Use of fossil fuels effects on the atmosphere and ecosystems.
and the expansion and intensification of agri- Chlorofluorocarbons (CFCs), for example,
culture have altered the cycles of carbon, nitro- were first produced in the 1950s as refrigerants,
gen, phosphorus, sulfur, and water on a global propellants, and solvents. They were heralded
scale (see Chapter 15). These changes in bio- for their nonreactivity in the lower atmosphere.
geochemical cycles not only alter the ecosys- In the upper atmosphere, however, where there
tems in which they occur but also influence is greater UV radiation, CFCs react with ozone.
unmanaged ecosystems through changes in The resulting ozone destruction, which occurs
lateral fluxes of nutrients and other materials primarily over the poles, creates a hole in the
through the atmosphere and surface waters protective blanket of ozone that shields Earth’s
(see Chapter 14). Land use changes, including surface from UV radiation. This ozone hole
deforestation and intensive use of fertilizers was initially observed near the South Pole. It
and irrigation, have increased the concentra- has expanded to lower latitudes in the South-
tions of atmospheric gases that influence ern Hemisphere and now also occurs at high
climate (see Chapter 2). Land transformations northern latitudes. As a result of the Montreal
also cause runoff and erosion of sediments and Protocol, the production of many CFCs has
nutrients that lead to substantial changes in ceased. Due to their low reactivity, however,
lakes, rivers, and coastal oceans. their concentrations in the atmosphere are only
Human activities introduce novel chemicals now beginning to decline, so their ecological
into the environment. Some apparently harm- effects will persist for decades. Persistent novel
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16 1. The Ecosystem Concept

chemicals, such as CFCs, often have long-lasting centrate cesium and strontium, as do people
ecological effects than cannot be predicted at who feed on reindeer. For this reason, the input
the time they are first produced and which of radioisotopes into the atmosphere or water
extend far beyond their region and duration of from nuclear power plants, submarines, and
use. weapons has had impacts that extend far
Other synthetic organic chemicals include beyond the regions where they were used.
DDT (an insecticide) and polychlorinated The growing scale and extent of human activ-
biphenyls (PCBs; industrial compounds), which ities suggest that all ecosystems are being influ-
were used extensively in the developed world enced, directly or indirectly, by our activities.
in the 1960s before their ecological conse- No ecosystem functions in isolation, and all are
quences were widely recognized. Many of these influenced by human activities that take place
compounds continue to be used in some devel- in adjacent communities and around the world.
oping nations. They are mobile and degrade Human activities are leading to global changes
slowly, causing them to persist and to be trans- in most major ecosystem controls: climate
ported to all ecosystems of the globe. Many (global warming), soil and water resources
of these compounds are fat soluble, so they (nitrogen deposition, erosion, diversions), dis-
accumulate in organisms and become increas- turbance regime (land use change, fire control),
ingly concentrated as they move through food and functional types of organisms (species
chains (see Chapter 11). When these com- introductions and extinctions). Many of these
pounds reach critical concentrations, they can global changes interact with each other at
cause reproductive failure. This occurs most regional and local scales. Therefore, all eco-
frequently in higher trophic levels and in systems are experiencing directional changes
animals that feed on fat-rich species. Some in ecosystem controls, creating novel condi-
processes, such as eggshell formation in birds, tions and, in many cases, positive feedbacks
are particularly sensitive to pesticide accumu- that lead to new types of ecosystems. These
lations, and population declines in predatory changes in interactive controls will inevit-
birds like the perigrine falcon have been noted ably change the properties of ecosystems and
in regions far removed from the locations of may lead to unpredictable losses of ecosys-
pesticide use. tem functions on which human communities
Atmospheric testing of atomic weapons in depend. In the following chapters we point out
the 1950s and 1960s increased the concentra- many of the ecosystem processes that have
tions of radioactive forms of many elements. been affected.
Explosions and leaks in nuclear reactors used
to generate electricity continue to be regional
or global sources of radioactivity. The explosion Summary
of a power-generating plant in 1986 at
Chernobyl in Ukraine, for example, released Ecosystem ecology addresses the interactions
substantial radioactivity that directly affected among organisms and their environment as an
human health in the region and increased the integrated system through study of the factors
atmospheric deposition of radioactive mate- that regulate the pools and fluxes of materials
rials over eastern Europe and Scandinavia. and energy through ecological systems. The
Some radioactive isotopes of atoms, such as spatial scale at which we study ecosystems is
strontium (which is chemically similar to chosen to facilitate the measurement of impor-
calcium) and cesium (which is chemically tant fluxes into, within, and out of the ecosys-
similar to potassium) are actively accumulated tem. The functioning of ecosystems depends
and retained by organisms. Lichens, for not only on their current structure and envi-
example, acquire their minerals primarily from ronment but also on past events and distur-
the atmosphere rather than from the soil and bances and the rate at which ecosystems
actively accumulate cesium and strontium. respond to past events. The study of ecosystem
Reindeer, which feed on lichens, further con- ecology is highly interdisciplinary and builds on
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Additional Reading 17

many aspects of ecology, hydrology, climatol- rainfall, soil carbon, consumption of plants
ogy, and geology and contributes to current by animals?
efforts to understand Earth as an integrated 3. What are the state factors that control the
system. Many unresolved problems in ecosys- structure and rates of processes in ecosys-
tem ecology require an integration of systems tems? What are the strengths and limitations
approaches, process understanding, and global of the state factor approach to answering
analysis. this question.
Most ecosystems ultimately acquire their 4. What is the difference between state factors
energy from the sun and their materials from and interactive controls? If you were asked
the atmosphere and rock minerals. The energy to write a management plan for a region,
and materials are transferred among compo- why would you treat a state factor and
nents within the ecosystem and are then an interactive control differently in your
released to the environment. The essential plan?
biotic components of ecosystems include 5. Using a forest or a lake as an example,
plants, which bring carbon and energy into the explain how climatic warming or the harvest
ecosystem; decomposers, which break down of trees or fish by people might change the
dead organic matter and release CO2 and nutri- major interactive controls. How might these
ents; and animals, which transfer energy and changes in controls alter the structure of or
materials within ecosystems and modulate the processes in these ecosystems?
activity of plants and decomposers. The essen- 6. Use examples to show how positive and neg-
tial abiotic components of ecosystems are the ative feedbacks might affect the responses of
atmosphere, water, and rock minerals. Ecosys- an ecosystem to climatic change.
tem processes are controlled by a set of rela-
tively independent state factors (climate, parent
material, topography, potential biota, and time) Additional Reading
and by a group of interactive controls (includ-
Chapin, F.S. III, M.S. Torn, and M. Tateno. 1996. Prin-
ing resource supply, modulators, disturbance ciples of ecosystem sustainability. American Natu-
regime, functional types of organisms, and ralist 148:1016–1037.
human activities) that are the immediate con- Golley, F.B. 1993. A History of the Ecosystem
trols over ecosystem processes. The interactive Concept in Ecology: More Than the Sum of the
controls both respond to and affect ecosystem Parts. Yale University Press, New Haven, CT.
processes. The stability and resilience of eco- Gorham, E. 1991. Biogeochemistry: Its origins and
systems depend on the strength of negative development. Biogeochemistry 13:199–239.
feedbacks that maintain the characteristics of Hagen, J.B. 1992. An Entangled Bank: The Origins
ecosystems in their current state. of Ecosystem Ecology. Rutgers University Press,
New Brunswick, NJ.
Jenny, H. 1980. The Soil Resources: Origin and
Behavior. Springer-Verlag, New York.
Review Questions Lindeman, R.L. 1942.The trophic-dynamic aspects of
ecology. Ecology 23:399–418.
1. What is an ecosystem? How does it differ Schlesinger, W.H. 1997. Biogeochemistry: An Ana-
from a community? What kinds of environ- lysis of Global Change. Academic Press, San
Diego.
mental questions can be addressed by
Sousa, W.P. 1985. The role of disturbance in natural
ecosystem ecology that are not readily communities. Annual Review of Ecology and
addressed by population or community Systematics 15:353–391.
ecology? Tansley, A.G. 1935. The use and abuse of vegetational
2. What is the difference between a pool and a concepts and terms. Ecology 16:284–307.
flux? Which of the following are pools and Vitousek, P.M. 1994. Beyond global warming:
which are fluxes: plants, plant respiration, Ecology and global change. Ecology 75:1861–1876.