Amajor stumbling block in the study of ecosystems is their bewildering complexity.

(ONeill and Waide 1981)

The perspective offered by ONeill and Waide (1981) in the above quote illustrates one of the
impacts of anthropogenic stressors on ecosystem processes. This perspective may also partially
explain the relative infrequency with which ecosystem processes are measured in biological assess-
ments. Because ecosystemsurprises (sensu Paine et al. 1998) may result fromfocusing on isolated
components, one potential solution to this bewildering complexity is to develop a comprehensive
understanding of emergent ecosystem properties (ONeill and Waide 1981). For example, we can
on abundance or biomass estimates of these functional groups. As described in the previous chapters,
the idea that behavior of a complex system often cannot be understood solely by analysis of its com-
ponents is a major thesis of hierarchy theory. The order that emerges from complex systems and
the constraints placed on the range of potential interactions in these systems are fundamental differ-
ences between randomly assembled populations and a stable ecosystem. The functional redundancy
of ecosystems that results from species replacement is a good example of our inability to predict
ecosystem responses based on understanding of components.
In one of the earlier theoretical treatments of ecosystemecotoxicology, ONeill and Waide (1981)
biological properties.
3. Treat the ecosystem as a biogeochemical system that focuses on movement of energy and
materials.
ators of contaminant effects than the amount of materials or energy stored in ecosystem
pools.
Thus, before we can understand how ecosystems respond to contaminants and other anthropo-
genic perturbations, it is necessary to develop an appreciation for the complex ecosystem processes
that are most likely to be affected by physical and chemical stressors. In the previous chapter, we
of how ecosystems respond to stressors will focus on these processes. Although general ecology
ecosystem separately, it is important to realize that these processes are intimately related. Patterns of
primary and secondary production in ecosystems are often limited by the amount of available nutri-
ents. Biogeochemical processes, the size of nutrient pools, and the rate of materials cycling in an
ecosystem can, in turn, be regulated by primary productivity. Finally, while our focus in this section
links between species richness, diversity, and ecosystem processes require that we also consider
structural features.
In addition to viewing ecosystems within a hierarchical context, contemporary ecologists routinely
characterize ecosystems based on bioenergetic and biogeochemical processes. Captured solar radi-
ation stored in chemical bonds by autotrophic organisms is made available to heterotrophs. As
described in the previous chapter, the perception that ecosystems are energy-transforming systems
emerged relatively early in the history of ecology. Eltons (1927) depiction of a tundra food web and
his recognition that a large number of herbivores are necessary to support a smaller number of pred-
izing ecosystem energetics. Although the use of calories or other units of energy as the currency to
integrate Eltons trophic levels did not occur for several decades, these early investigations helped
to formalize contemporary perspectives of ecosystem dynamics. Eltons (1927) food web became
Lindemans (1942) food cycle that was eventually formalized as a universal energy model by Odum
(1968) that also included a material-cycling component.
Flux of energy through an ecosystem is determined by the rate at which plants assimilate energy
of these conversions. Because contaminants and other stressors can affect any of these processes,
synthesis in plants is the conversion of light energy and raw materials (carbon dioxide and water) to
carbohydrates and oxygen:
6CO
2
6H
2
O Light energy C
6
H
12
O
2
6O
2
(30.1)
Although this stoichiometrically balanced chemical reaction to describe photosynthesis is correct,
it is not especially satisfying from an ecological perspective and should be expanded to include both
2002):
Inorganic carbon Nutrients Light energy Biomass Heat (30.2)
The energy necessary for the conversion of CO
2
to a reduced state in carbohydrates is provided by
and the difference between GPP and these metabolic costs is called net primary production (NPP).
accounting for respiration:
NPP GPP Respiration (30.3)
Methods for measuring NPP in terrestrial and aquatic ecosystems are diverse, but typically focus on
assessing changes in biomass, CO
2
, or O
2
. The most direct method for estimating NPP in terrestrial
ecosystems is the harvest method, which generally involves measuring the increase in plant standing
crop or biomass (B) over a growing season.
B B
2
B
1
(30.4)
where B
2
is the biomass at time 2 and B
1
is the biomass at time 1.
Note that primary production of an ecosystem is a functional measure of the instantaneous rate
of biomass generation, generally expressed as dry weight of plant material (or carbon) per unit area
per unit time (g/m
2
/year). In contrast, biomass is a structural measure of the amount of plant material
present at one particular point in time. Amore energetically appropriate measure may be obtained by
converting dry weight of plant material to calories. Estimates of both NPP and biomass have been
used as endpoints in assessing stressor impacts on ecosystem energetics.
Other approaches for estimating primary production involve measuring gas exchange
(e.g., uptake of CO
2
or release of O
2
and the use of radioactive carbon isotopes,
14
C. Although
harvest methods provide the most direct measure of NPP in terrestrial ecosystems and have been
employed to estimate production of larger marine plants (macrophytes, kelp), they are less com-
mon in aquatic ecosystems because of small size and rapid turnover of primary producers. Three
approaches have been employed in aquatic ecosystems to estimate primary productivity: light and
dark bottles oxygen techniques, radioisotopes such as
14
C, and in situ diel approaches. The traditional
approach for aquatic systems uses light and dark bottles containing water with ambient phytoplank-
ton populations. This approach compares changes in dissolved oxygen concentration ( [O
2
]) in
bottles held in the light with changes in the dark. Because [O
2
] in the light is a result of both GPP
and respiration whereas change in the dark bottle is a result of respiration only, GPPcan be estimated
by the difference between these measures:
GPP O
2
light O
2
dark (30.5)
Asimilar approach has been used to estimate metabolism in stream ecosystems in which cobble
substrate collected from the streambed is placed in light and dark chambers. This approach provides
an estimate of whole community metabolism because the cobble substrate typically includes both
autotrophic and heterotrophic organisms (e.g., algae, bacteria, fungi, and invertebrates).
The carbon-14 (
14
C) technique provides a considerably more sensitive estimate of primary
productivity, which may be necessary in oligotrophic systems where GPP is very low. The
14
C tech-
nique is also preferred by some ecologists because it allows researchers to explicitly follow carbon
phytoplankton are incubated with a tracer amount of
14
Clabeled dissolved inorganic carbon. The
accumulation of carbon in organic matter relative to the dissolved inorganic fraction provides a
measure of primary production.
Note that the light and dark bottle technique and the
14
C incubation technique may be comprom-
ised by container artifacts. Isolation of primary producers from natural systems by placement in
bottles may result in depleted nutrient concentrations, decreased turbulence and mixing, and growth
of organisms on the sides of the container. In situ approaches that measure diel changes in O
2
or CO
2
eliminate these bottle effects and provide ecosystem-level estimates of GPPand respiration. Changes
in CO
2
or O
2
during daylight are a result of GPP, whole ecosystemrespiration, and exchange with the
atmosphere. Thus, whole ecosystem GPP and respiration can be estimated by measuring changes in
O
2
or CO
2
during the daylight and at night, and correcting for atmospheric exchange. Avariation of
this approach is used in streams, where whole ecosystemmetabolismis determined by comparing O
2
or CO
2
concentrations at upstreamand downstreamlocations and measuring the travel time between
stations.
Numerous abiotic factors limit primary productivity in both terrestrial and aquatic ecosystems;
however, light, temperature, nutrients, and moisture (in terrestrial habitats) are generally considered
1%), whereas higher values are observed in forests (23.5%) (Cooper
pigments and inhibit photosynthesis. Similarly, the rate of photosynthesis generally increases with
temperature, up to some optimal value, and then declines. Because respiration also increases with
temperature, optimal temperatures for NPP and photosynthesis will likely differ, complicating our
ability to predict the precise relationship between temperature and NPP. Finally, the amount of
available nutrients, especially nitrogen (N) and phosphorus (P), limit primary production in many
ecosystems. Primary production of aquatic ecosystems is particularly sensitive to nutrient limitation,
increase algal productivity (Ryther and Dunstan 1971). Although most of the research on nutrient
limitation has focused on N and P, some ecosystems may be limited by other materials. Studies
conducted using water collected from the Sargasso Sea, a highly oligotrophic ecosystem, showed
that enrichment with N and P had relatively little effect on phytoplankton (Menzel and Ryther 1961).
Primary productivity in much of the open ocean is limited by iron, which has stimulated interest
in the use of iron to fertilize the oceans as a measure to increase sequestration of anthropogenic
CO
2
. Not surprisingly, many of the most comprehensive studies demonstrating effects of nutrients
on productivity have been conducted in lakes where the association between primary productivity
and abiotic factors has been documented experimentally (Schindler 1974).
Because light is rapidly attenuated in aquatic ecosystems, the amount of light available to primary
producers decreases as a function of depth according to the following equation:
dI dz kI (30.6)
where I amount of solar radiation, z depth, and k
amount of light at 10 m depth in open seawater is about 50% lower than at the surface. Because of
greater amounts of light absorbing materials, values of k in lakes and other productive ecosystems
are considerably greater. In deep rivers, lakes, and marine ecosystems, the reduction in light limits
as the area near the surface where photosynthesis is greater than respiration.
In addition to the direct effects of these limiting factors, combined and interactive effects of light
levels, nutrients, and other abiotic factors can affect primary production in aquatic ecosystems. In
a large-scale comparison across several ecoregions in North America, Bott et al. (1985) concluded
that the combined effects of photosynthetically active radiation (PAR), chlorophyll a, and water tem-
perature accounted for 70% of the variation in community metabolism among streams. Similarly,
Fleituch (1999) reported that benthic community metabolism along a river continuum was primar-
and conductivity. It is well established that enrichment of aquatic ecosystems caused by excessive
nutrients often stimulates primary production and causes excessive plant growth, including blooms
of potentially toxic blue-green algae. Because these dense populations of algae limit light penetra-
tion, dramatic shifts in the structure and function of major primary producers may occur. Attached
replaced by phytoplankton communities that are capable of remaining near the surface.
Because of its association with global climate change, ecologists have recently given special
2
and other abiotic factors on primary productivity. Researchers
hypothesize that if elevated CO
2
increases primary productivity, some of the excess anthropogenic
carbon released fromburning fossil fuels and land use changes may be sequestered into plant biomass.
This response remains uncertain because of the potential for other factors (e.g., nutrients, light,
temperature) to limit primary production in terrestrial and aquatic ecosystems. Melillo et al. (1993)
useda terrestrial ecosystemmodel (TEM) to predict the effects of climate change and elevatedCO
2
on
NPP. Spatially referenced information on climate, soils, vegetation, water availability, and elevation
were usedtopredict current NPPvalues for a wide varietyof ecosystems. Model predictions of current
responses of NPP to a doubling of CO
2
and associated changes in temperature, precipitation, and
cloud cover as predicted by general circulation models (Figure 30.1). Overall global NPP increased
by approximately 23%, but there was considerable variation among ecosystems. This geographic
underlying mechanisms. For example, moist temperate ecosystems responded primarily to elevated
temperature and increased nitrogen cycling whereas dry temperate ecosystems responded primarily
to elevated levels of CO
2
.
Productivity is not evenly distributed among regions of the world, and comparisons of NPP and
biomass estimated by Whittaker and Likens (1973) for some of the worlds major biomes reveal
Ecosystem type
0
1
2
3
4
5
6
7
Current NPP
Predicted NPP
N
P
P

(
1
0
g
C
/
y
)
Results froma TEMused to predict the effects of a 2 increase in CO
2
and associated changes
in temperature, precipitation, and cloud cover on NPP of different terrestrial ecosystems. (Data from Table 2 in
Melillo et al. (1993).)
Terrestrial ecosystems
Tropical forest 1800 42
Temperate forest 1250 32
Boreal forest 800 20
Temperate grassland 500 1.5
Alpine and tundra 140 0.6
Desert scrub 70 0.7
Aquatic ecosystems
Algal beds and reefs 2000 2
Estuaries 1800 1
Lakes and streams 500 0.02
Continental shelf 360 0.01
Open ocean 125 0.003
Source. Data from Whittaker and Likens (1973).
several interesting patterns (Table 30.1). Although sunlight is necessary for primary production, it
is evident from Table 30.1 that other factors contribute to global patterns. If adequate moisture or
nutrients are not available, as in arid ecosystems or the open ocean, NPP will be low regardless of
the levels of sunlight. In forest ecosystems, a general decrease in NPP is seen as we move to colder
moisture results in very high productivity for tropical forests. Despite the generally low productivity
of open ocean ecosystems, estuaries, algal beds, and coral reefs are among the most productive
aquatic habitats.
major primary producers. Biomass in terrestrial habitats is generally much greater than in aquatic
ecosystems, and this large terrestrial biomass represents an important pool of global carbon. The
lower biomass in aquatic environments results from the relatively small body size of dominant
primary producers (e.g., phytoplankton), which has important implications for trophic dynamics.
Because small primary producers in aquatic ecosystems are capable of very rapid turnover, they can
support a relatively large biomass of consumers compared to terrestrial ecosystems. The ratio of
productivity to biomass (P:B) also varies greatly among different biomes and ecosystems. As shown
the large amount of nonphotosynthetic biomass in these ecosystems (e.g., bark, trunk, and branches).
In contrast, P:Bratios in aquatic ecosystems, especially those dominated by phytoplankton, are much
higher because of their small size and rapid turnover rates. The production values of lentic and marine
point in time. These differences in growth forms and turnover rates between terrestrial and aquatic
ecosystems may also have important consequences for responses to anthropogenic disturbances.
Because of their rapid growth rates, we expect that primary producers in aquatic ecosystems would
respond more rapidly to contaminants than in terrestrial ecosystems.
predators that obtain their energy from plant or animal biomass. Consumers such as bacteria and
Consumption (C) Total amount of energy consumed
Egestion (E) Total amount of energy lost to egestion
Assimilation (CE) Total amount of energy available for production and
respiration
Production (CA) Total amount of energy available for growth and
reproduction
C 100%) Portion of consumed food that is assimilated
A 100%) Portion of assimilated food that is converted to new biomass
C 100%) Portion of consumed food that is converted to new biomass
n
A
n 1
trophic levels n, a consumer, and n 1, the resource
fungi, organisms that obtain energy from decomposing plant and animal material, should also be
included in measures of secondary production. Secondary productivity is similar to primary pro-
ductivity in that we must distinguish between the portion of energy for growth and reproduction
(and thus available to higher trophic levels) and the portion associated with maintenance costs of the
consumer (Table 30.2). As noted above, the amount of energy available to consumers is ultimately
herbivores or predators is actually assimilated. Because food quality for predators is generally greater
tion, respiration, and maintenance), is generally greater in predators. In addition to the recalcitrant
materials in plant tissue, herbivores must also contend with a diverse assortment of defensive chem-
icals produced by plants, which also limits consumption. Interestingly, coevolutionary responses
in herbivores, which coincidentally provide protection against some xenobiotics. In contrast to ter-
feeding on unprotected phytoplankton or algae.
Only a small fraction of the assimilated energy in consumers is available for growth and reproduction;
the remaining is necessary for maintenance and respiration. Because metabolic costs are generally
of assimilated food available for new biomass, is generally lower in warm-blooded organisms
amount of consumed food available for biomass, is a function of both assimilation and production
there is considerable variation among ecosystems (Pauly and Christensen 1995). The important
an ecosystem (Table 30.3). Ricklefs (1990) estimated the average length of food chains based on
Open ocean 500 0.1 25 7.1
Coastal marine 8000 10.0 20 5.1
Temperate grassland 2000 1.0 10 4.3
Tropical forest 8000 10.0 5 3.2
Source. Data from Ricklefs (1990).
the following equations:
E n NPP Eff
n 1
(30.7)
n 1
log E n log NPP
log Eff
(30.8)
where n number of trophic levels, E n energy available to a predator at a trophic level n,
and Eff
overall NPP.
Estimates of secondary production for some species can be derived from measures of feeding rates,
conditions (West 1968). However, determining secondary production in natural populations is more
challenging and generally requires estimates of consumption, growth, and reproduction. Sophist-
icated bioenergetics models have been developed for some aquatic species such as large-mouth
bass (Kitchell 1983). These individual-based models generally use laboratory-derived estimates of
consumption, respiration and elimination, and then solve for growth.
Consumption Respiration Wastes Growth (30.9)
Several practical issues complicate our ability to estimate whole ecosystem production using these
individual-based models. While estimates of secondary production for individual species, especially
those for which we have a thorough understanding of natural history (Jordan et al. 1971, Kilgore and
Armitage 1978), have been developed, integrating this information to derive secondary production
estimates for whole ecosystems or even major components of ecosystems is challenging. Wiens
(1973) estimated secondary production of grassland bird communities, and Chew and Chew (1970)
examined energy relationships of dominant mammals in a desert shrub community. Perhaps the best
examples have been developed in aquatic ecosystems where researchers have derived community-
level estimates of secondary production for major taxonomic or functional feeding groups (Benke
and Wallace 1980, 1997, Carlisle 2000, Fisher and Gray 1983). Secondary production (P) in benthic
macroinvertebrates is obtained from estimates of biomass and growth rate using the following
simple relationship:
P B
i
g
i
(30.10)
where B
i
and g
i
are biomass and growth rates of the ith species.
While the methodology for estimating secondary production in aquatic ecosystems is well estab-
lished, these are labor-intensive efforts. Estimating biomass for macroinvertebrates is relatively
straightforward; however, the intensive sampling frequency necessary to determine growth rates
of macroinvertebrates often limits application of this technique. Because of these methodological
ecotoxicological literature. An example of one such exception, Carlisle (2000) constructed food webs
based on quantitative analyses of macroinvertebrate secondary production for six different streams
along a gradient of heavy metal pollution.
trophic groups are also impediments to studies of secondary production. Early attempts to quantify
the relationship between NPP and secondary production should be evaluated cautiously because of
the failure to appreciate the dominant role of decomposers and microbial production. The opinion
of ONeill et al. (1986) that the trophic level concept is most useful as a heuristic device and tends
to obscure, rather than illuminate, organizational principles of ecosystems is likely shared by many
ecosystemecologists. The use of stable isotopes, described in Section 34.4.4, is one potential solution
to this problem; however, relatively few studies have employed this technique in ecosystem-level
studies of secondary production.
Numerous studies have reported a direct quantitative relationship between primary productivity and
secondary productivity or biomass of consumers (Coe et al. 1976, Cyr and Pace 1993, McNaughton
et al. 1989). Amajor emphasis of the International Biological Program described in Chapter 29 was
to understand the biological basis of productivity and to quantify relationships between primary and
secondary production. Much of this research focused on understanding the underlying mechanisms
and consequences of interactions between plants and consumers. Some of the strongest evidence
to support the relationship between NPP and secondary production has been obtained from exper-
imental introductions of nutrients to whole ecosystems. The predictable increases in both primary
when investigating ecosystem energetics. Intuitively, we would expect that herbivore biomass or
production would increase with NPP; however, the nature of this relationship will likely vary among
ecosystems and herbivore types. For example, because grassland herbivores consume a larger por-
tion of NPP than forest herbivores (Whittaker 1975), the relationship between NPP and herbivore
abundance in forest ecosystems is relatively weak (Figure 30.2). Concentrations of structural com-
pounds, such as lignins and other recalcitrant materials that limit herbivory in terrestrial ecosystems,
are generally lower in aquatic primary producers. Consequently, grazers in many aquatic ecosystems
consume a large fraction of available biomass (30%40%) and the relationship between primary and
secondary production in these systems is generally much stronger. Because a greater fraction of NPP
is removed in aquatic ecosystems, we also predict that predators would play a more important role
are supported by studies showing the relative importance of top-down effects in aquatic ecosys-
tems compared to terrestrial ecosystems (Strong 1992). Predator control over lower trophic levels,
termed trophic cascades, has been frequently observed in aquatic ecosystems but only rarely in ter-
restrial ecosystems. Similarly, bottom-up control of herbivores and other consumers by nutrients and
primary producers is quite common in many lentic ecosystems and is the mechanism responsible
Primary productivity
S
e
c
o
n
d
a
r
y


p
r
o
d
u
c
t
i
v
i
t
y
Forest ecosystems
Grassland ecosystems
Aquatic ecosystems
Hypothetical relationship between net primary productivity (NPP) and secondary productivity
in aquatic and terrestrial ecosystems. Stronger relationships are expected in aquatic ecosystems because grazers
consume a larger portion of plant biomass compared to terrestrial ecosystems.
for cultural eutrophication. Understanding the relationship between NPP and secondary production
in ecosystems is important for predicting potential contaminant effects. It is possible that some of
the variation in this relationship may account for differences in contaminant transfer rates among
ecosystems. These issues will be explored in Chapter 34.
The movement of materials and energy in ecosystems has been investigated using a variety of
descriptive, theoretical, and empirical approaches. Attempts to develop comprehensive explanatory
models that connect physical, chemical, and biological processes have been especially successful in
aquatic ecosystems. Vannotes classic paper The river continuum concept (Vannote et al. 1980)
recognized that patterns and processes in streams change predictably from headwaters to the mouth.
In addition to linking geomorphologic characteristics of a watershed to biological processes, this
paper elucidated mechanisms responsible for the downstream transport, utilization, and storage of
energy and materials. The major tenets of the river continuum concept (RCC) can be summarized by
consideringlongitudinal changes inthe sources of energy andmaterials fromupstreamto downstream
(Figure 30.3). The relative importance of allochthonous and autochthonous sources of energy shift
from upstream to downstream, resulting in changes in the ratio of NPP to respiration and structural
alterations in the composition of stream communities. Shaded headwater streams are generally
heterotrophic (P/R<1) because the dense riparian canopy in these systems limits primary productivity
opens, shading and the relative input of organic materials from riparian areas is reduced, and the
stream becomes autotrophic (P/R>1). Finally, large rivers may return to heterotrophic conditions
(P/R<1) because of increased depth and greater light attenuation.
Longitudinal changes in the abundance and composition of macroinvertebrate functional feeding
ance of allochthonous and autochthonous inputs. The abundance of organisms that utilize coarse
particulate organic material (CPOM) (e.g., leaf litter) is greatest in headwater streams and decreases
downstream. Grazers, organisms that consume attached algae and periphyton, are more important
Tests of the predictions of the RCCin different geographic regions have provided good support for
the major tenets in NorthAmerica (Bott et al. 1985, Minshall et al. 1983) and Europe (Fleituch 1999).
Minshall et al. (1983) measured benthic organic matter, community metabolism, decomposition,
F
P
O
M
CPOM
Shredders
Collectors
Predators
Grazers
Shredders
Collectors
Predators
Grazers
Collectors
Predators
1
2
3
4
5
6
7
8
9
S
t
r
e
a
m

o
r
d
e
r
P > R
P < R
P < R
Major predictions of the river continuumconcept showing changes in ecosystemenergetics and
in Vannote et al. (1980).)
and functional feeding group composition along longitudinal gradients in streams from four distinct
geographic areas in North America. Although regional and local variation was observed, changes in
structure and function from headwaters to downstream sites were consistent with predictions of the
ecosystems that clearly illustrated the important connections between upstream and downstream
size within the same drainage had previously been treated as completely different systems. By
visualizing streams as a continuum of processes along a longitudinal gradient, ecologists recognized
that ecosystem function occurring in small headwater streams or large rivers could be described
using similar models (Figure 30.4). Despite differences in community composition and the relative
importance of allochthonous and autochthonous inputs, similar processes operate in both headwater
and mid-order streams. The RCC provided a conceptual framework for testing hypotheses about
visualized these systems.
through biotic and abiotic compartments of ecosystems. Figure 30.5 is a simple model illustrating the
of materials through these compartments. Nutrients and other materials are assimilated from soil
or water by autotrophic organisms (plants and autotrophic bacteria), passed on to consumers, and
released back to abiotic compartments. The amount and availability of nutrients are among the most
important factors that limit primary productivity. In addition to limiting growth rates of primary pro-
CPOM
Shredders
FPOM
Collectors
Primary
production
Grazers
Headwater stream
Grazers
CPOM
Collectors
Primary
production
Grazers
Mid
CPOM
Shredders FPOM
Mid-order stream
Energy pathways and the importance of allochthonous and autochthonous materials in head-
water and mid-order streams. Despite considerable variation in the sources of energy and dominant functional
Minshall et al. (1983).)
Animals
Primary producers Bacteria
Detritus
Atmosphere
Soil
Water
Sediments
Photosynthesis
and assimilation
Organic compounds
(peat, coal, oil)
norganic compounds
(e.g., limestone)
Weathering
Erosion
Sedimentary
rock formation
Respiration
and excretion
Erosion, burning
fossil fuels
Simple model showing the connection between biotic and abiotic compartments and the dom-
from Figure 12.3 in Ricklefs (1990).)
and anthropogenic disturbances (DeAngelis et al. 1989). More importantly, an understanding of
nutrient and material cycles is essential for predicting ecosystem consequences of increased anthro-
pogenic inputs of certain materials, especially carbon, nitrogen, sulfur, and phosphorus. Predicting
the consequences of altered biogeochemical cycles also requires that we consider the ecosystem as
a unit instead of focusing only on component parts (ONeill and Waide 1981). Because the behavior
and pathway of many toxic chemicals follow those of natural elements in ecosystems, considera-
tion of biogeochemical processes is essential for understanding fate and transport of contaminants.
Finally, we will see that many toxic chemicals have direct effects on ecosystems because they alter
biogeochemical processes.
The primary difference between movement of energy and nutrients through an ecosystem is that
nutrients are retained and constantly cycled between biotic and abiotic components by a variety of
meteorological, geological, and biological processes. In other words, the earth is considered an open
nutrients such as C, N, and P are assimilated, transformed, and released back to the ecosystem,
often in a very different form, where they can be used again. Meteorological processes include
precipitation, snowmelt, and atmospheric deposition. The major geological process is weathering
of materials from soils and underlying geological formations. Biological processes are analogous
chains. If input of materials exceeds output, these unused materials may also accumulate in nutrient
is relatively constant, we can calculate the length of time an average molecule resides in this pool.
measure of the accessibility of materials to organisms. For example, the atmosphere is a relatively
active pool for oxygen, with a residence time of about 20,000 years. In contrast, the atmosphere is
accessibility of atmospheric N to organisms.
At a local level, if we assume that nutrient concentrations within a compartment are at equilibrium
(e.g., uptake is approximately equal to export), then measurement of uptake or loss provides an
estimate of turnover time within a compartment (Figure 30.6). Turnover time of organic materials and
nutrients varies among ecosystems and is closely related toclimate and temperature. Table 30.4shows
the accumulation and mean turnover times for organic material and nitrogen in several different forest
types located in different climatic zones. Nitrogen and organic matter accumulations are greatest in
The low rates of decomposition observed in cold boreal forests result in a larger fraction of organic
material in soils relative to trees. Turnover time, the average amount of time that a molecule remains
in soil before it is assimilated by plants, increases in colder climates and is longer in coniferous
forests because foliage is not replaced each year.
Nutrient Pool or
Standing Stock
Biomass
Nutrient Pool or
Standing Stock
Biomass
Production
or input
Production
or input
Yield
or output
Yield
or output
Fast turnover
Slow turnover
Nutrient pool or
standing stock
biomass
Nutrient pool or
standing stock
biomass
in Krebs (1994).)
Boreal coniferous 226,000 353 3,250 230.0
Boreal deciduous 491,000 26 3,780 27.0
Temperate coniferous 618,000 17 7,300 17.9
Temperate deciduous 389,000 4 5,619 5.5
Source. Data from Cole and Rapp (1981).
Atmospheric CO
2
(640)
Dissolved total CO
2
(30,000)
Algae
(5)
Bacteria
(1,500)
Respiration
(50)
Animals
Bacteria,
fungi
Plants
(450)
Dead organic
material (700)
Limestone, dolomite (18,000,000) Coal, oil, and natural gas (25,000,000)
Assimilation
(50)
(35)
Exchange
(84)
Assimilation
(35)
Respiration
(35)
(25)
Animals
Sedimentation (<1)
Oceans
Land
The carbon cycle. Numbers in parentheses indicate the amount of C in each compartment and
Despite fundamental differences between the movement of energy and materials, nutrients and
other elements in biogeochemical cycles are closely associated with primary production and often
ecosystem metabolism and secondary production (Figure 30.7). Energy stored in carbohydrates by
primary producers is ultimately released when these high energy compounds are oxidized to CO
2
by
consumers and decomposers. The movement of carbon among compartments integrates biological
processes such as assimilation and respiration with physical processes such as atmospheric-oceanic
2
are present
in the oceans, the largest pools of carbon are deposited in sediments (limestone, dolomite) and stored
as fossil fuels. Carbon dioxide occurs in a relatively lowconcentration in the atmosphere. Autotrophic
organisms (primarily plants) assimilate CO
2
and incorporate it into organic matter by photosynthesis,
and a portion of the CO
2
is returned to the atmosphere by respiration.
These physical, chemical, and biological processes are closely integrated in aquatic ecosystems
through the carbonatebicarbonate system. The CO
2
dissolved in water forms carbonic acid, which
readily disassociates to bicarbonate and carbonate ions in the following reactions:
CO
2
H
2
O H
2
CO
3
(30.11)
H
2
CO
3
H HCO
3
(30.12)
HCO
3
H CO
3
(30.13)
Because these reactions are dependent on pH, the amount of calcium (which equilibrates with the
bicarbonate and carbonate ions), and metabolism, they illustrate the close connection between biotic
removal of CO
2
by photosynthesis or addition of CO
2
by respiration will drive these reactions to the
and CO
3
in aquatic
alterations in the carbonatebicarbonate system have important toxicological implications.
Phosphorus is a major limiting nutrient in aquatic ecosystems and primarily responsible for eutroph-
ication of many lakes and streams. The P cycle is relatively simple because the atmosphere plays a
relatively small role. Consequently, transport of P in ecosystems is primarily sedimentary and at a
local scale. The major source of P to ecosystems is from underlying rocks (Figure 30.8), and loss
from soils is usually balanced by releases of inorganic P from weathering. Plants assimilate phos-
phorus as phosphate (PO
3
4
), and availability and rate of uptake are dependent on pH. Herbivores
Plant, animal
Tissue (organic N)
Atmospheric N
Ammonia,
nitrates
Ammonia,
Ammonium
Lightning
D
e
n
i
t
r
i
f
i
c
a
t
i
o
n
N
i
t
r
i
f
i
c
a
t
i
o
n
N

f
i
x
a
t
i
o
n
Nitrogen cycle
Plant, animal
tissue (organic N)
Ammonia,
nitrates
Ammonia,
ammonium
D
e
n
i
t
r
i
f
i
c
a
t
i
o
n
N
i
t
r
i
f
i
c
a
t
i
o
n
Herbivores
Plants
Soils
Phosphorus cycle
Rocks
PO
4
norganic
P
E
x
c
r
e
t
i
o
n
,

d
e
c
o
m
p
o
s
i
t
i
o
n
Weathering
Herbivores
Plants
Soils Rocks
The phosphorus and nitrogen cycles showing the major biotic and abiotic pathways.
obtain all of their required P from consumption of plants, and P is returned to soil by excretion
32
P in an aquatic
microcosm from primary producers (phytoplankton, periphyton) to zooplankton. This study is an
through an ecosystem. Similar to the situation for many contaminants, this study also demonstrated
that the ultimate fate of
32
P was sediments, which was shown to be a major reservoir of P in aquatic
ecosystems.
The N cycle is considerably more complex because it involves a major atmospheric compon-
ent and because N can exist in numerous oxidation states. Five basic processes drive the N cycle:
vast majority of N occurs in the atmosphere as molecular N
2
, a form that is unavailable to plants.
Rhizobium) and cyanobacteria (blue-green algae) in aquatic envir-
onments convert atmospheric N
2 3
2
and then to nitrates (NO
3
) by several different groups of bacteria,
including Nitrosomonas and Nitrobacter. Plants assimilate N primarily as nitrate, and N is released
of denitrifying bacteria (Pseudomonas) reduces nitrates and releases inorganic N back to the atmo-
ication, toxic algal blooms (Burkholder and Glasgow 1997), and the formation of anoxic conditions
in the Gulf of Mexico (Rabalais et al. 1998).
Compared with other biogeochemical cycles such as C, N, and P, anthropogenic alteration of
the global sulfur (S) cycle by combustion of fossil fuels has been extreme. Approximately 60% of
be described in Section 35.2. The S cycle is also relatively complex because S can exist in several
oxidation states, and conversion among these different forms is dependent on different types of
bacteria. Sulfur in a sedimentary phase such as organic matter and rocks can be released by natural
processes such as weathering and erosion. The gaseous form of S (H
2
S) is released from volcanoes
and decomposition of organic material. Sulfur released to the environment either by natural or
anthropogenic processes is oxidized to sulfate (SO
2
4
) and deposited. Sulfur dioxide released from
the combustion of fossil fuels is oxidized and converted to sulfuric acid (H
2
SO
4
).
Unlike the situation observed in mature, undisturbed forests where most nutrients are generally
retained, a fraction of nutrients and other materials are transported downstream in lotic ecosystems
either in dissolved or particulate forms. Instead of cycling as observed in terrestrial systems, the
movement of nutrients in lotic ecosystems is generally represented as a downstream spiral (Elwood
a molecule travels as it completes a cycle between organic and inorganic phases. The length of
including the rate of microbial mineralization, stream temperature, stream velocity, the shape of the
stream channel, and the number of snags and other woody debris that reduce downstream transport.
uptake by organisms, is an important characteristic of ecosystem function. Uptake length essentially
S
w
F
w
wU (30.14)
where S
w
uptake length, F
w
U uptake rate of nutrients
fromwater, and w average stream width. Uptake length generally increases with streamdischarge
and decreases with temperature, the amount of riparian vegetation, and biomass of detritus and algae.
In relatively small streams, attached algae, fungi, bacteria, and periphyton are responsible for
most of the uptake of nutrients, which generally follows MichaelisMenten kinetics. Some of these
materials may be released back to the water column, but a fraction enters benthic food chains through
grazing organisms. Nutrients and materials continue to spiral downstream in larger rivers, but as
stream size increases nutrient dynamics in these systems more closely resemble patterns observed
in lentic ecosystems. The same processes that determine retention and transport of nutrients have
well established that periphyton and attached algae are important sinks for contaminants in lotic
ecosystems, and highly persistent chemicals may spiral downstream as they move between biotic
and abiotic compartments.
The relative importance of allochthonous inputs to stream energy budgets has been well established.
However, the contribution of these terrestrial inputs to nutrient dynamics has received considerably
less attention. Whole ecosystem nutrient budgets have been calculated for a few relatively small
watersheds. In one of the most comprehensive studies, Triska et al. (1984) measured inputs from
stream in the H.J. Andrews Experimental Forest, Oregon, USA. Most of the total annual nitrogen
input (15.25 g/m
2 2
, with biological inputs contributing an
additional 4.19 g/m
2
.Direct and indirect biologically derived inputs from litterfall,throughfall,
tothe stream.Totalinputofnitrogenwas34%greaterthanoutput, indicatingthatthe streamwas
notoperatingata steadystate. The difference betweeninputandoutputwasprimarilya resultof storage ofnitrogenasparticulate organic matter.
Constructing nutrientbudgets for whole ecosystems requires thatwe identify and measure the
processes thatcontrolinputs and outputs. Researchers atHubbard Brook ExperimentalForest,
a deciduous forestlocated in the White Mountains of New Hampshire, USA, have developed mass
budgets for a variety of nutrients (Likens etal. 1970). Because the Hubbard Brook watershed is
by measuring stream discharge, precipitation, and concentrations of materials in precipitation and
stream water. Because terrestrial losses of nutrients were eventually released to streams, measures
mass budgets for the entire watershed (Table 30.5). With the exceptions of NH
4
and NO
3
, output
compared to the pools of materials. This result demonstrates that, in stable watersheds such as Hub-
bard Brook, the majority of nutrients are usually retained and recycled. The export of sulfur, derived
this result will be discussed in Section 35.2.
Experimental injection of nutrients and tracers is the most direct method to examine retention and
transport of nutrients in streams. The general approach is to add a small amount of a radioactive
Input 2.7 16.3 38.3 1.1 2.6 0.7 1.5
Output 0.4 8.7 48.6 1.7 11.8 2.9 6.9
Net change 2.3 7.6 10.3 0.6 9.2 2.2 5.4
Source. Data from Likens et al. (1970).
or stable isotope at one point, and measure concentrations at several points downstream. Triska
et al. (1989) reported that 29% of the N injected into a third-order forested stream in California was
retained, while the remaining portion was transported downstream. Despite uptake by autotrophs,
which was greatest during the day, nitrate concentrations increased downstream, indicating that the
stream reach was a source of dissolved N to benthic communities. Decreased respiration and tissue
C:N ratios downstream of the injection point indicated a biological response to N enrichment.
Although the preferred method to measure nutrient dynamics and uptake length in streams is to
use tracers (e.g.,
32
P,
15
N) that maintain ambient concentrations, because of expense and logistical
issues, short-term additions that increase ambient nutrient concentrations are becoming increasingly
common (Davis and Minshall 1999, Hall et al. 2002). Although this approach may be useful for com-
paring different streams, Mulholland et al. (2002) reported that uptake length was overestimated by
short-termnutrient addition experiments compared to tracer additions. Because the degree of overes-
timation was related to the level of nutrient addition, these authors concluded that nutrient additions
should be as low as possible, while maintaining the ability to accurately measure concentrations at
several locations downstream.
Our attempts to place spatial boundaries on ecosystems are compromised by our recognition that
materials and energy readily move between ecosystems. Aquatic ecologists have long recognized the
connect processes occurring in upland terrestrial systems to lakes and oceans (Fisher et al. 1998).
the quality and quantity of transported materials through input, storage, and instream biogeochem-
ical processes. The boundaries of stream ecosystems were previously considered to extend only
a short distance into the riparian zone. However, stream ecologists now recognize the important
linkages between streams and upland ecosystems (Fausch et al. 2002). Rather than visualizing
stream ecosystems as longitudinal corridors, recent studies emphasize vertical and lateral connec-
tions between the stream and surrounding landscape. Fisher et al. (1998) extended the nutrient
spiraling concept to include processes that occur outside of the stream. They suggested that the
nested, concentric arrangement of subsystems within stream ecosystems (e.g., surface water, hypo-
rheic zone, and riparian zone) is analogous to a telescope, where the length of the telescoping
required to transform an amount of material in transport. Thus, because processing length increases
with disturbance, we would expect that impacted systems would have lower rates of materials
cycling.
Salmon carcasses
Primary
production
Secondary
production
Terrestrial
scavengers
Anadromous
salmon
Riparian
vegetation
Juvenile salmon
growth and survival
Although we have traditionallyconsidered the transport of nutrients and other materials in streams
as a one-way process, in some instances nutrients exported downstream may be returned to head-
organic matter and N when the salmon return to their natal streams to spawn (Figure 30.9). Because
many of these streams are naturally oligotrophic and have a heavy canopy that limits primary pro-
ductivity, these nutrient subsidies can be very important to ecosystem productivity. Studies using
stable isotopic tracers of
15
Nand
15
to streams and adjacent riparian habitats. Because salmon are enriched with heavier isotopes of N
and C, comparisons of primary producers, consumers, riparian vegetation, and wildlife in streams
with and without spawning salmon have revealed the importance of these subsidies. In addition to
stimulation of primary producers and bottom-up effects on higher trophic levels (Bilby et al. 1996,
diversity of riparian vegetation, and provide up to 25% of the N to riparian plants and 3090% of
the N to the diet of terrestrial scavengers (Naiman et al. 2002). Increased primary and secondary
productivity associated with salmon carcasses translated to greater growth rates and survival of
juveniles inhabiting these streams, providing a positive feedback for returning salmon (Bilby et al.
1996). Similarly, declines in the abundance and biomass of spawning salmon that return to their
natal streams have important consequences for the function of both aquatic and adjacent terrestrial
ecosystems. In addition to transporting marine-derived nutrients to headwater streams, anadromous
Although there is little information on the effects of these marine-derived pollutants on headwater
(Ewald et al. 1998).
The vast majority of studies investigating processes that control rates of material cycling have been
Comparisons of nutrient cycles among different ecosystems (e.g., tropical vs. temperate forests; arid
vs. humid grasslands; headwater streams vs. large rivers), at different altitudes, and among different
geomorphological units improve our understanding of these regulating factors and provide insights
into underlying mechanisms. Comparative studies across ecosystems may also provide the best
Carpenter 2000). Cross-ecosystem comparisons are essentially an optimization problem, where the
suitability of controls decreases but the ability to make broad generalizations increases among highly
dissimilar ecosystems (Fisher and Grimm 1991).
Anthropogenic activities have resulted in increased N loading to aquatic and terrestrial ecosys-
tems primarily from agricultural activities and fossil fuel combustion. Because some of the excess
N deposited in headwaters is transported downstream, an understanding of factors that control N
uptake and export is essential for predicting ecosystem effects at the watershed level. In a compre-
hensive analysis of nutrient dynamics and metabolism in streams, researchers have used nutrient
tracer experiments to measure ammoniumuptake and retention in 11 streams ranging fromthe North
Slope of Alaska to Puerto Rico (Mulholland et al. 2001, 2002, Peterson et al. 2001, Webster et al.
2003). The Lotic Intersite Nitrogen Experiment (LINX) compared streammetabolismand Ndynam-
ics in tropical, arid, temperate, and tundra streams. The goal of this large-scale comparative study
was to relate inter-biome variability in stream metabolism and nutrient uptake to physical, chem-
ical, and biological characteristics. Stream metabolism (i.e., autotrophic primary production, and
autotrophic and heterotrophic respiration) was measured using the upstreamdownstream diurnal
dissolved oxygen technique. To measure ammonium uptake,
15
NH
4
was injected in each stream and
samples were collected at downstream sites to determine uptake length. Stream metabolism (GPP)
was closely related to PAR (400700 nm) and P concentration (Mulholland et al. 2001). Comparison
of results across different biomes indicated that ammonium uptake length varied by approximately
two orders of magnitude (141350 m) and increased with stream discharge (Webster et al. 2003). In
shallow headwater streams with a higher surface-to-volume ratio, most of the uptake and removal
processes occurred through assimilation by benthic autotrophic and heterotrophic organisms and by
sorption to sediments (Peterson et al. 2001). Because a large fraction of Ninputs to headwater streams
was retained, especially during periods of high productivity, these systems regulated downstream
transport to lakes, rivers, and estuaries and thereby may reduce eutrophication.
Comparative studies of lakes and streams provide an opportunity to assess factors that regulate move-
ment of materials in two different types of ecosystems that vary widely in their major hydrologic
dom ina te d b y ve r t ic a l e xc ha nge s be tw e e n e pilim ne tic , me ta lim ne tic , a nd h ypolim ne tic z one s. T he r e la ti ve im por ta nc e of a l loc hthono us a nd a ut oc hthonous input is a ls o q uite di f f e r e nt in la ke s a nd s tr e a ms. B e c a us e of the s e dif f e r e n c e s, a pp r oa c he s use d by e c olog is t s to quant if y nutr i e n t c yc ling in on primary production, whereasstreamecologistsare more concerned withuptake length. Inlakes,
uptakeofnutrientsoccursprimarilybyphytoplanktonandbacteria,whereasperiphytonandattached algaeplay more important rolesin loticecosystems. Grazing by zooplankton and subsequent return of nutrientstothe watercolumnviaexcretion and mortalitycanoccurrapidlyinlakes.The rate at whichnutrients are depositedto and releasedfrom sediments isdependent on lake size,the volume
Because of differences in the movement of materials in lakes and streams, and the different meth-
odological approaches, a common currency is necessary to compare factors that regulate nutrient
cycling in lentic and lotic ecosystems. In both lakes and streams, cycling of nutrients is a function
of uptake and export of dissolved and particulate materials (Figure 30.10). Essington and Carpenter
Dissolved nutrient pool
Particulate nutrient pool
Uptake Release
mport
Export
Particulate
Dissolved
Particulate
Dissolved
Conceptual model showing the import and export of dissolved and particulate nutrients in
that a nutrient molecule is used before it is exported from the system, in lakes and streams. The
recycling ratio is a dimensionless number calculated as the ratio of uptake rate (U, mass time
1
)
to export rate (E, mass time
1
in particulate form (primarily to sediments) will be greater in lakes, whereas export rates of dis-
solved materials will be greater in streams. Nutrient cycling in streams is controlled primarily by
the association of nutrients with particles and downstream transport. In lakes, nutrient cycling is
controlled by physical processes that reduce the rate of sedimentation and biological processes that
increase remineralization. Essington and Carpenters (2000) model also predicts that the effects of
consumers (grazers in streams; zooplankton in lakes) will be fundamentally different in lakes and
streams. Although the model was developed to compare nutrient cycling in lakes and streams, it can
be used to quantify the movement of contaminants through these systems. We would expect that
movement of contaminants in streams would be dominated by downstream export in the dissolved
phase, whereas movement in lakes would be dominated by sedimentation.
Relatively few studies have compared ecosystem processes across terrestrial and aquatic ecosys-
tems. In a comprehensive analysis of 800 aquatic and terrestrial systems, Cebrian and Largitue
(2004) examined factors that controlled herbivory and decomposition rates. Although NPP varied
greatly within aquatic and terrestrial ecosystems, there was surprisingly little variation between these
ecosystem types when analyzed across all studies. Nutritional quality of primary producers was an
important predictor of the proportion of NPP consumed by herbivores in both aquatic and terrestrial
ecosystems, indicating the extent of top-down regulation of producer biomass. In contrast, while
the total consumption by herbivores (g C/m
2
/year) was correlated with NPP, it was unrelated to
nutritional quality.
Energy has been considered the universal currency for studying ecosystem processes for several
decades. Although the transfer of energy through aquatic and terrestrial foodwebs can reveal much
about how ecosystems operate, our understanding of this process remains somewhat incomplete.
Ecological stoichiometry improves on this single currency approach by using ratios of certain ele-
ments (C, N, and P) to characterize how composition of organisms and their prey affect nutrient
chemical substances in ecological interactions and processes (Sterner and Elser 2002). By compar-
ing elemental ratios in abiotic compartments, primary producers, and consumers, limiting elements
obtained (Anderson and Boersma 2004). Ecological stoichiometry has been used to examine link-
ages between N and C cycles in terrestrial and aquatic ecosystems. The approach is based on the
concept that relative amounts of elements in different compartments can regulate nutrient cycling
aquatic and terrestrial ecosystems. Simple trophochemical diagrams (Sterner and Elser 2002) may
be used to depict relative amounts of elements in different trophic levels and compare spatial (among
ecosystem types) and temporal stoichiometric patterns (Figure 30.11).
The study of ecological stoichiometry is critically important to our understanding of anthropo-
genic alterations in global nutrient cycles. Anthropogenic increases in CO
2
ition. Analyses of global increases in these materials indicate that N and P are disproportionately
enriched relative to C, thus potentially disrupting natural stiochiometric ratios of these elements
describing CO
2
effects on primary productivity cannot be considered in isolation but should also
account for anthropogenic increases in other nutrients (Sterner and Elser 2002). Similarly, elevated
Concentration of P
C
o
n
c
e
n
t
r
a
t
i
o
n

o
f

N
Algae
Bacteria
Fish
Z
Concentration of C
Simple trophochemical diagramused in ecological stoichiometry to depict relative amounts of
elements in different trophic levels and to compare spatial (among ecosystemtypes) and temporal stoichiometric
Natural 61 0.13 0.003
Anthropogenic 8 0.14 0.012
Percent increase from 13 108 400
anthropogenic sources
Source. FromFalkowski, P., et al., Science, 290, 291296, 2000.
CO
2
will likely produce leaf litter with greater C:N ratios, resulting in lower rates of mineralization
and decomposition.
At a more local level, enrichment of certain elements resulting from anthropogenic releases may
alter processes occurringin ecosystems. On the basis of an analysis of major ecosystemcompartments
rates in streams located in several biomes. The C:N ratios increased from consumers to primary
eutrophication of downstreamsystems. Although undisturbed headwater streams are highly retentive
of N, anthropogenic deposition of N to these watersheds will decrease C:N ratios and increase
N export downstream (Dodds et al. 2004).
Decomposition is the process by which dead organic matter (detritus) is broken down to its com-
ponent parts. Decomposition is a critical characteristic of ecosystem function and has been shown
to vary among ecosystems and with levels of disturbance. Decomposition rates are determined
by a complex interplay of physical, chemical, and biological processes. Factors such as chemical
composition and nutritional quality of detritus, physicochemical characteristics of the habitat, and
be consumed by detritivores or further processed by bacteria and fungi, with CO
2
and inorganic
materials (e.g., NH
4
) as the end products. Organic material may be fragmented into smaller sizes
by physical processes and invertebrates, thereby increasing surface area of particles. The net res-
ult is the conversion of particulate organic materials to dissolved constituents and the transfer of
carbon to decomposers and detritivores. In terrestrial ecosystems, the most important macroinver-
tebrate decomposers include nematodes, insects, isopods, crustaceans, and oligochaetes. In aquatic
ecosystems, CPOM is processed by shredders (primarily aquatic insects) and converted to FPOM.
Secondary production of some aquatic ecosystems, particularly canopied headwater streams, may be
heavily subsidized by inputs of allochthonous materials. In contrast to autochthonous energy derived
from macrophytes, periphyton, phytoplankton, and other primary producers, allochthonous energy
sources consist of organic materials and detritus derived from outside sources. Detritus-based food
webs are common in many marine and lotic ecosystems. The relative contribution of allochthonous
and autochthonous materials to secondary production is dependent on the dimensions and shape of
the stream or lake and will be described in Section 31.2.3.
of the nutritional quality provided to detritivores is derived from bacteria and fungi that rapidly
colonize leaf surfaces during decomposition (Suberkropp and Klug 1976). High nutritional quality
of detritus is expectedtoincrease growth rates andmetabolic activity of detritivores and decomposers,
thereby increasing the rate of decomposition. In general, nutritional quality of detritus, expressed
as concentrations of N and P, is greater in aquatic ecosystems than in terrestrial systems. The rate
of detrital production is also highly correlated with NPP in both aquatic and terrestrial ecosystems
and nutrients also play a role in determining decomposition rates and the size of detritus pools.
Because the fraction of NPP consumed by herbivores is greater in aquatic than in terrestrial
ecosystems (Cyr and Pace 1993), the amount of carbon channeled to detritus is generally greater
in most ecosystems, with greater than 50% of NPP going to decomposers (Cebrian and Lartigue
2004). The fate of this material and the relative importance of invertebrate detritivores and microbes
have received considerable attention in the ecological literature. Studies conducted in aquatic and
decay. Various approaches have been employed to assess the relative importance of physical pro-
cesses, macroinvertebrates, and microbial processes in litter decomposition. Computer simulations
were used to calculate an organic matter budget for a second-order stream at Coweeta Hydrolo-
gic Laboratory, North Carolina (Webster 1983). Results showed that macroinvertebrate shredders
were responsible for approximately 27% of the downstream transport of particulate organic mater-
ial. Elimination of macroinvertebrate shredders using the pesticide methoxychlor (discussed in
of detritus (Wallace et al. 1982).
comes from experimental studies that increased nutrient concentrations or eliminated terrestrial
inputs. Elwoodet al. (1981) showedthat nutrient enrichment of a small heterotrophic streamincreased
decomposition rates, resulting in greater abundance of consumers and higher trophic levels. Wallace
et al. (1997) used an overhead canopy to eliminate leaf litter inputs to a small headwater stream for
abundance and biomass of most major taxa and declines in overall secondary production. Predator
production also declined in streams where leaf litter was eliminated, demonstrating strong bottom-up
effects and important linkages between detritus and higher trophic levels.
food web diagrams, measuring the outcome of decomposition is relatively straightforward. Changes
in mass (dry weight) or nutrient content of organic material over time is generally used as an indic-
ator of decomposition rates. Long term assessments of organic matter dynamics can be obtained in
terrestrial habitats by measuring changes in total soil organic carbon pools using combustion tech-
niques. However, these methods do not provide an estimate of carbon availability, which is often
independent of the amount of carbon stored in pools (Robertson and Paul 2000). Fractionation of
organic carbon into humic acids, fulvic acids, and other constituents may provide some insight into
the availability of carbon to microbial decomposers (McKnight 2001).
To measure decomposition rates, organic material such as leaf litter is placed in mesh bags in
mass loss and nutrient changes. Leaf or litter materials are usually collected from representative
and relevant species in the ecosystem. The decomposition rate constant, k, is estimated from the
following equation:
X
t
X
0
e
kt
(30.15)
where X
t
is the mass remaining at time t and X
0
rate constant (k) is the slope of the least squares regression of proportion mass remaining versus time.
and different ecosystems (Figure 30.12). Mesh size of litter bags is often manipulated to exclude or
include certainmacroinvertebrate groups andtodiscern the relative importance of macroinvertebrates
and microbes in decomposition. Mass loss may be related to the abundance or biomass of detritivores
that colonized the bags during the exposure period. Because small mesh of litter bags may impede
cautiously. Leaf litter decomposition studies in terrestrial ecosystems are generally conducted over
Time (days)
M
a
s
s

r
e
m
a
i
n
i
n
g

(
%
)
Arctic tundra
k = 0.005
Semiarid grassland
k = 0.25
Tropical forest
k = 1.40
Hypothetical relationship between percent mass remaining and time in three ecosystems with
varying decomposition rates. k the decomposition rate constant.
longer time periods (13 years) than in aquatic ecosystems (36 months). Because contaminants will
affect both microbial processes and abundance of macroinvertebrate decomposers, values of k are
expected to decrease in stressed ecosystems.
Flow of energy, cycling of nutrients, and decomposition of litter are fundamental processes in all
ecosystems. A basic understanding of these processes and how they vary spatially and temporally
is necessary to predict how these systems may respond to anthropogenic stressors (Howarth, 1991).
Because the components of ecosystems (e.g., populations and communities) may respond quite dif-
ferently to anthropogenic disturbances, an ecosystem perspective is necessary to limit the likelihood
biogeochemical cycles in ecosystems are intimately coupled with the movement of contaminants,
basic models developed by ecosystem ecologists may also help explain fate of xenobiotic chemicals.
Each of the processes we have discussed in this chapter is likely to be affected by physical or chemical
stressors to some degree. The relative sensitivity and variability of these responses compared with
more traditional population- and community-level endpoints will likely vary among stressors and
among ecosystemtypes. In the following two chapters we will consider descriptive and experimental
approaches that have examined effects of stressors on these ecosystem processes.
The perspective that ecosystems are highly complex may explain the relative infrequency
with which ecosystem processes are measured in biological assessments.
Before we can understand how ecosystems respond to contaminants and other anthropo-
genic perturbations, it is necessary to develop an appreciation for the ecosystemprocesses
that are most likely to be affected.
In addition to viewing ecosystems within a hierarchical context, ecologists routinely
characterize ecosystems based on bioenergetic and biogeochemical processes.
Flux of energy through an ecosystem is determined by the rate at which plants assimilate
energy by photosynthesis, the transfer of this energy to herbivores and other consumers,
The energy necessary for the conversion of CO
2
to a reduced state in carbohydrates is
as GPP.
reproduction after accounting for respiration.
through an ecosystem is an important indicator in ecosystem-level assessments.
In addition to the direct effects of limiting factors on NPP, combined and interactive
effects of light, nutrients, and other abiotic factors can affect primary production in aquatic
ecosystems.
2
on primary
productivity because of its association with global climate change.
and predators that obtain their energy from plant or animal biomass.
levels in an ecosystem.
While the methodology for estimating secondary production in aquatic ecosystems is
well established, most techniques are highly labor-intensive and consequently secondary
production.
Intuitively, we would expect that herbivore biomass or production would increase with
NPP; however, the nature of this relationship varies among ecosystems and herbivore
types.
Because a greater fraction of NPP is removed in aquatic ecosystems, it is likely that
ecosystems.
Variation in the relationship between NPP and secondary production may account for
differences in contaminant transfer rates among ecosystems.
The amount and availability of nutrients are among the most important factors that limit
primary productivity and decomposition rates.
Because the behavior of many toxic chemicals is similar to that of natural elements in
ecosystems, consideration of biogeochemical processes is essential for understanding
fate and transport of contaminants.
The earth is considered an open system with respect to energy but a closed system with
respect to materials such as C, N, and P.
Despite fundamental differences between the movement of energy and materials, nutri-
ents and other elements in biogeochemical cycles are closely associated with primary
Unlike the situation observed in mature, undisturbed forests where most nutrients are
downstream in lotic ecosystems, a process known as nutrient spiraling.
Experimental injection of nutrients and tracers is the most direct method to examine
retention and spiraling of nutrients in streams.
Rather than visualizing lotic ecosystems as longitudinal conduits for transporting nutri-
ents and other chemicals downstream, ecologists now recognize the important linkages
between streams and upland ecosystems.
Although we have traditionally considered the transport of nutrients and other materials
in streams as a one-way process, in some instances materials exported downstream may
be returned to headwaters.
Comparative studies across ecosystems such as the LINX project may provide the best
logical interactions and processes (Sterner and Elser 2002), uses ratios of certain elements
(C, N, and P) to characterize how composition of organisms and their prey affect nutrient
broken down to its component parts, is a fundamental process of ecosystems that varies
Because the fraction of NPP consumed by herbivores is greater in aquatic than terrestrial
ecosystems, the amount of carbon channeled to detritus is generally greater in terrestrial
systems.
Changes in mass (dry weight) or nutrient content of organic material over time is generally
used as an indicator of decomposition rates.
Anderson, T.R. and Boersma, M.R.D., Stoichiometry: Linking elements to biochemicals, Ecology, 85,
11931202, 2004.
Appalachian stream, Ecology, 61, 108118, 1980.
web analysis, Ecology, 78, 11321145, 1997.
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