408 PART SIX • ECOSYSTEM ECOLOGY

ECOLOG ICAL IS S U ES Human Appropriation of Net Primary Productivity

A lthough we represent only one of more than 1.5 million

known species inhabiting our planet, humans use a vastly
disproportionate share of Earth’s resources. This pattern of
Nations on products consumed in 1995 for 230 countries in
seven categories (plant foods, meat, milk, eggs, wood, paper,
and fiber), a per capita value of HANPP was calculated for each
resource use is most evident in the human appropriation of net country. The per capita estimate of HANPP was then applied to
primary productivity (NPP). In a paper first published in 1986, a gridded database of the human population at a spatial reso-
ecologist Peter Vitousek and colleagues at Stanford University lution of 0.25° (latitude and longitude). This spatial scale was
used three different approaches to estimate the fraction of NPP chosen to match the spatial resolution of NPP derived from the
that humans have appropriated. A low estimate was derived by satellite-derived vegetation index (NDVI). The resulting map
calculating the amount of NPP people use directly—as food, depicts the spatial pattern of HANPP, showing where the prod-
fuel, fiber, or timber. An intermediate estimate was also calcu- ucts of terrestrial photosynthesis are consumed (Figure 1a). By
lated that includes all the productivity of lands devoted entirely combining the maps of global NPP and HANPP, the research-
to human activities (such as the NPP of croplands, as opposed to ers were able to map HANPP as a percentage of NPP, provid-
the portion of crops actually eaten). This estimate also included ing a spatially explicit balance of NPP “supply” and “demand”
the energy human activity consumes, such as in setting fires to (Figure 1b). The resulting map reveals a great deal of regional
clear land. A third approach provided a high estimate, which variation in the appropriation of NPP.
further included productive capacity lost as a result of convert- Summing for the globe, the researchers estimate annual
ing open land to cities and forests to pastures or because of de- HANPP to be 24.2 Pg organic matter (1 Pg = 1015g), or ap-
sertification or overuse (overgrazing, excessive erosion). Results proximately 20 percent of terrestrial annual NPP (with high
of the three approaches (low, intermediate, and high) yielded a and low estimates of 14–26 percent). Some regions, however,
wide range of estimates: 3 percent, 19 percent, and 40 percent, such as Western Europe and south central Asia, consume
respectively. These estimates are a remarkable level of use for more than 70 percent of their NPP. In contrast, HANPP in
a species that represents approximately 0.5 percent of the total other regions (typically in the wet tropics) is less than 15
heterotrophic biomass on Earth. percent. The lowest value (approximately 6 percent) is found in
Since the initial analyses conducted by Vitousek and col- South America.
leagues in the mid-1980s, advances in satellite technology have Both population and per capita consumption interact to de-
greatly enhanced scientists’ ability to monitor patterns of land use termine the human ecological impact at a regional scale. The
and primary productivity at a continental to global scale. Marc role of population is clear from Figure 1b despite vast differ-
Imhoff and Lahouari Bounoua, researchers at NASA’s Goddard ences in consumption among nations. For example, east and
Space Flight Center (GSFC; Greenbelt, Maryland), and col- south central Asia, with almost half of the world’s population,
leagues have recently undertaken an analysis of patterns of human appropriates 72 percent of its regional NPP despite having the
use of NPP at a global scale using satellite-derived data (see lowest per capita consumption of any region. Affluence also
http://earthobservatory.nasa.gov/Features/MeasuringVegetation). plays an important role. The average annual per capita HANPP
The researchers defined human appropriation of terrestrial for industrialized countries is almost double that of developing
NPP (HANPP) as the amount of terrestrial NPP required to de- nations, which are home to 83 percent of the global population.
rive food and fiber products consumed by humans, including If the per capita HANPP of the developing nations increased
the organic matter that is lost during harvesting and process- to match that of the industrialized countries, global HANPP
ing of whole plants into end products. Using data compiled by would increase by 75 percent to a value of 35 percent of the
the Food and Agricultural Organization (FAO) of the United current global NPP.

21.9 Consumers Vary in Efficiency Of the food ingested by a consumer (I ), a portion is assimi-
lated across the gut wall (A ), and the remainder is expelled from
of Production the body as waste products (W ). Of the energy that is assimi-
Although there is a general relationship between the availability lated, some is used in respiration (R) and the remainder goes
of primary productivity and the productivity of consumer organ- to production (P), which includes production of new tissues
isms (secondary productivity) across a variety of terrestrial and as well as reproduction. The ratio of assimilation to ingestion
aquatic ecosystems, within a given ecosystem there is consider- (A/I ) the assimilation efficiency, is a measure of the efficiency
able variation among consumer organisms in their efficiency to with which the consumer extracts energy from food. The ratio
transform energy consumed into secondary production. These of production to assimilation (P/A) the production efficiency,
differences can be illustrated using the following simple model is a measure of how efficiently the consumer incorporates
of energy flow through a consumer organism. assimilated energy into secondary production.
 CHAPTER 21 • ECOSYSTEM ENERGETICS 409

1. Use Figure 21.5 together with Figure 1 to answer 2. Since the mid-20th century, the efficiency of agricultural
the following question. Why is the HANPP production (the quantity of crops produced per hectare of
expressed as a percentage (Figure 1b) low for the land under cultivation) has increased dramatically. How
wet tropical regions of the world (regions of tropical would this increase have changed the values of HANPP
rain forest)? shown in Figures 1a and 1b?

HANPP
Units: log10 (g C/yr)
13.4
9.8

5.8
Excluded area

(a)

HANPP
Units: % of NPP
200–40,000
100–200
80–100
60–80
45–60
30–45
20–30
10–20
0.1–10
0

(b)

Figure 1 Spatial distribution of the annual NPP resources required by the human population,
as measured by (a) HANPP, and (b) HANPP as a percentage of local NPP.
(Nature Publishing Group.)

A consumer’s ability to convert the energy it ingests into little energy in respiratory heat and converting more assimilated
secondary production varies with species and the type of con- energy into production. Among the vertebrates, ectotherms
sumer. Assimilation efficiencies vary widely among ectotherms have intermediate values of production efficiency (approxi-
and endotherms. Endotherms are much more efficient than mately 10 percent). In contrast, endotherms, with their high
ectotherms. However, carnivorous animals, even ectothermic energy expenditure associated with maintaining a constant
ones, have a higher assimilation efficiency (approximately 80 body temperature, convert only 1 to 2 percent of their assimi-
percent) than herbivores (20 percent to 50 percent). Predatory lated energy into production (see Section 7.11).
spiders feeding on invertebrates have assimilation efficiencies For endotherms, body size also influences production
of more than 90 percent. efficiency. You may recall that the mass-specific metabolic rate
Production efficiency varies mainly according to (kcal/g body weight/hr) increases exponentially with decreasing
taxonomic class (Table 21.2). Invertebrates in general have body mass (Chapter 7, Section 7.10). An increase in mass-
high efficiencies (30 percent to 40 percent), losing relatively specific metabolic rate lowers production efficiency.
410 PART SIX • ECOSYSTEM ECOLOGY

Table 21.2 Production Efficiency (P/A • 100) Grazing food chain Detrital food chain
of Various Animal Groups R R

Group P/A (%)

Mice 4.10
Voles 2.63 Carnivores Carnivores

Other mammals 2.92

R R
Birds 1.26
Fish 9.74
Social insects 8.31 Decomposer
Herbivores
Orthoptera 41.67 herbivores
Hemiptera 41.90
R R
All other insects 41.23
Mollusca 21.59
Crustacea 24.96 Primary
Detritus
All other noninsect invertebrates 27.68 producers
Noninsect invertebrates
Herbivores 18.81 Figure 21.19 Two parts of any ecosystem: a grazing food
Carnivores 25.05 chain and a detrital food chain. Orange arrows linking trophic levels
represent the flow of energy associated with ingestion. The blue
Source: Data from Humphreys 1979. arrows from each trophic level represent the loss of energy through
respiration. The brown arrows represent a combination of dead organic
matter (unconsumed biomass) and waste products (feces and urine).
21.10 Ecosystems Have Two Major
Food Chains in each food chain are the source of energy for the carnivores,
Energy fixed by plants is the base that the rest of life on Earth and so on. Cattle grazing on pastureland, deer browsing in the
depends on. This energy stored by plants is passed along forest, insects feeding on leaves in the forest canopy, or zoo-
through the ecosystem in a series of steps of eating and being plankton feeding on phytoplankton in the water column all
eaten—known as a food chain (see Section 17.4). Feeding rela- represent first-level consumers of the grazing food chain. In
tionships within a food chain are defined in terms of trophic or contrast, a variety of invertebrates—such as snails, beetles, milli-
consumer levels. From a functional rather than a species view- pedes, and earthworms, as well as fungi and bacteria—represent
point, all organisms that obtain their energy in the same num- first-level consumers of the detrital food chain (see Chapter 22).
ber of steps from the autotrophs or primary producers belong Figure 21.19 combines the two food chains to produce a
to the same trophic level in the ecosystem. The first trophic generalized model of trophic structure and energy flow through
level belongs to the primary producers, the second level to the an ecosystem. The two food chains are linked. The initial source
herbivores (first-level consumers), and the higher levels to the of energy for the detrital food chain is the input of dead organic
carnivores (second-level consumers). Some consumers occupy matter and waste materials from the grazing food chain. This
a single trophic level, but many others, such as omnivores, linkage appears as a series of brown arrows from each of the
occupy more than one trophic level (see Section 7.1). trophic levels in the grazing food chain, leading to the box des-
Food chains are descriptive. They represent a more abstract ignated as detritus or dead organic matter. There is one notable
expression of the food webs presented earlier (Chapters 17 difference in the flow of energy between trophic levels in the
and 18). Major feeding groups are defined based on a common grazing and decomposer food chains. In the grazing food chain,
source of energy, such as autotrophs, herbivores, and carnivores. the flow is unidirectional, with net primary production provid-
Each feeding group is then linked to others in a manner that ing the energy source for herbivores, herbivores providing the
represents the flow of energy (see Figure 18.6 for a simple food energy for carnivores, and so on. In the decomposer food chain,
chain). Boxes represent the three feeding groups: autotrophs, the flow of energy is not unidirectional. The waste materials and
herbivores, and carnivores. The arrows linking the boxes repre- dead organic matter (organisms) in each of the consumer trophic
sent the direction of energy flow. levels are “recycled,” returning as an input to the dead organic
Within any ecosystem there are two major food chains: the matter box at the base of the detrital food chain. In addition,
grazing food chain and the detrital food chain (Figure 21.19). the distinction between the grazer and consumer food chains is
The distinction between these two food chains is the source of often blurred at the higher trophic levels (carnivores), as preda-
energy for the first-level consumers, the herbivores. In the graz- tors rarely distinguish whether prey draw their resources from
ing food chain, the source of energy is living plant biomass or primary producers or detritus. For example, the diet of an insec-
net primary production. In the detrital food chain, the source of tivorous bird might include beetle species that feed on detritus,
energy is dead organic matter or detritus. In turn, the herbivores as well as species that feed on green leaf tissues.
 CHAPTER 21 • ECOSYSTEM ENERGETICS 411

21.11 Energy Flows through We quantify this flow with the energetic efficiencies defined in
Section 21.9: the assimilation efficiency (A/I ) and the produc-
Trophic Levels Can Be Quantified tion efficiency (P/A) One additional index of energetic efficiency,
To quantify the flux of energy through the ecosystem using the however, must be introduced: consumption efficiency. The con-
conceptual model of the food chains just discussed, we need to sumption efficiency, the ratio of ingestion to production at the
return to the processes involved in secondary production dis- next lower trophic level (In/Pn - 1) defines the amount of available
cussed in Section 21.8: consumption, ingestion, assimilation, energy being consumed. Sample values of these efficiencies for
respiration, and production. We will examine a single trophic an invertebrate herbivore in the grazing food chain are provided in
compartment (Figure 21.20a). The energy available to a given Figure 21.20b. Using these efficiency values, we can track the fate
trophic level (designated as n) is the production of the next- of a given amount of energy (1000 kcal) available to herbivores in
lower level (n - 1); for example, net primary production (P1) the form of NPP through the herbivore trophic level.
is the available energy for grazing herbivores (trophic level 2). If we apply efficiency values for each trophic level in the
Following the simple model of energy flow through a consumer grazing and detrital food chains, we can calculate the flow of
organism presented in Section 21.9, some proportion of that pro- energy through the whole ecosystem. The production from
ductivity is consumed or ingested (I ); the remainder makes its each trophic level provides the input to the next-higher level,
way to the dead organic matter of the detrital food chain. Some and unconsumed production (dead individuals) and waste
portion of the energy consumed is assimilated by the organ- products from each trophic level provide input into the dead
isms (A), and the remainder is lost as waste materials (W ) to organic matter compartment. The entire flow of energy through
the detrital food chain. Of the energy assimilated, some is lost to the ecosystem is a function of the initial transformation of solar
respiration, shown as the arrow labeled R that is leaving the upper energy into NPP. All energy entering the ecosystem as NPP
left corner of the box, and the remainder goes to production (P). eventually is lost through respiration.

Pn 14

R 56

A 70

W 130

I 200

800
Not
consumed
Pn −1 1000

(a) (b)

Invertebrate herbivore
efficiencies
I/Pn–1 = 20%
A/I = 35%
Pn/A = 20%
W = Waste
R = Respiration

Figure 21.20 (a) Energy flow within a single trophic compartment. (b) An example of
quantifying energy flow for an invertebrate herbivore using estimates of efficiencies provided in
table. Values are kilocalories (kcal).

I N T E R P R E T I N G E C O L O G I C A L D ATA
Q1. Assume that the assimilation efficiency is changed from 35 percent to 25 percent; what is
the new value for respiration?
Q2. What is the trophic efficiency (TE) for the example presented in the figure above? (See
Section 21.12.)
412 PART SIX • ECOSYSTEM ECOLOGY

FI ELD S TUD IES Brian Silliman

Department of Zoology, University of Florida, Gainesville, Florida

T he salt marshes that fringe the coastline of eastern North

America are among the most productive ecosystems in the
world. For the past half century, the prevailing theory found in
on Spartina growth
(individual plants)
and productivity
the ecological literature and textbooks has been that productivity (collective growth
in these coastal ecosystems is controlled by physical conditions of plants). He used
(water depth, frequency of inundation, salinity, etc.) and nutri- cages, 1 m2 in size,
ent availability, referred to as “bottom-up” control (see Sections made from a fine-
18.4 and 21.4). But as a result of research conducted by ecologist mesh wire fencing
Brian Silliman of the University of Florida, this long-held view is material, to establish
being brought into question. The focus of Silliman’s research is different experimen-
the role of consumer organisms in the salt marshes of the south- tal treatments (see
eastern United States. What is emerging from his work is a rich, Figure 2). In some of
complex picture of salt marshes that involves the interactions of the cages, snails were
marsh plants, “fungus-farming” snails, and an array of predators, excluded; in others, snails were added to establish populations
including the blue crab. of differing densities. The results of Silliman’s experiments were
The salt-marsh tidal zones of eastern North America are dramatic. Moderate to high snail densities led to runaway graz-
dominated by salt-marsh cordgrass (Spartina alterniflora). The ing effects, ultimately transforming one of the most productive
most abundant and widespread grazer in these communities is ecosystems in the world into a barren mudflat (Figure 2). The
the marsh periwinkle (Littoraria irrorata; Figure 1). The marsh effect of snails on plant growth and productivity is not through
periwinkle is a small snail, reaching 2.5 cm in length, with pop- the direct consumption of green plant tissues, but by preparing
ulation densities upward of 500 individuals per square meter the leaf tissue for colonization by their preferred food: fungus.
within the tidal zone dominated by Spartina. As the snail crawls along the leaf surface, it scrapes the surface
While he was a graduate student working in the salt with its band of sawlike teeth called radulae, creating wounds
marshes along the eastern shore of Virginia, Silliman explored that run lengthwise on the leaf surface and kill the surrounding
the role of the herbivory on patterns of net primary productiv- tissues. While it travels, the snail also deposits feces containing
ity (NPP) and standing biomass. Before Silliman’s work, it was fungal spores and nutrients, effectively stimulating the estab-
assumed that the grazers had little influence on the growth of lishment and growth of fungus.
Spartina plants and the overall productivity of the marsh. Snails
were believed to function largely as part of the decomposer
food chain, feeding on dead and dying plant tissues. Silliman
designed an experiment to assess the influence of grazing snails

Figure 2 Effects of snail grazing on Spartina standing crop

and canopy structure after eight months. Photographs show
Figure 1 The marsh periwinkle on Spartina leaf. experimental exclosure in area of high snail density.
 CHAPTER 21 • ECOSYSTEM ENERGETICS 413

In a series of follow-up experiments with colleague Steven

Newell of the University of Georgia Marine Institute, Silliman
has demonstrated a mutualistic relationship between snails
and fungi at the expense of the Spartina plants, the resource
Predators
upon which both depend. The snails employ a low-level food
production strategy whereby they prepare a favorable envi-
ronment for fungal growth, provide substrate to promote
growth, add supplemental nutrients and propagules, and –
consume fungus. Although this type of facilitation, known as
“fungus farming,” has been reported for some beetle, ter-
mite, and ant species, the work by Silliman and his colleague
is the first reported case of this type of cultivation behavior
Snails
+
outside of insects.
Given the potential of the snails to so dramatically re-
duce plant growth and marsh primary productivity at even
–
moderate population densities, how do the salt marshes re-
main so productive? To answer this question, Silliman has
turned his attention to the structure of the marsh’s food
chain. The periwinkle has several potential predators, includ-
ing the terrapin, mud crab, and blue crab. Silliman hypoth-
esized that predation maintains snail populations below the
Spartina
densities at which they have a devastating impact on Spartina
plants. This type of control on plant productivity, in which
predators limit populations of herbivores, is called “top-
down” control.
To establish the role of predators in determining the dis-
tribution of snails within the marsh, Silliman once again used
wire mesh cages—this time not to exclude the snails, but to Figure 3 The proposed mechanism of the marsh trophic
exclude their predators. Other plots (areas of marsh) without cascade. Grazing by snails has a negative impact on Spartina
cages were monitored for comparison (control plots). The populations. Likewise, predators have a negative impact on
experiment ran for a year, after which the snail populations snail populations. The net effect is that predators have a positive
were estimated in both the exclosure and control plots. The indirect effect on Spartina populations by reducing rates of
experiment results supported the hypothesis that predators herbivory by snails.
control the distribution and abundance of snail populations
(density of adults as well as recruitment of juveniles into the
Bibliography
population). Once again, the results were dramatic. The den-
Silliman, B. R., and J. C. Zieman. 2001. Top-down control on
sity of juvenile snails was 100 times greater in the plots where
Spartina alterniflora production by periwinkle grazing in a
predators were excluded as compared to the control plots.
Virginia salt marsh. Ecology 82:2830–2845.
Predators such as the blue crab do indeed control snail popu-
Silliman, B. R., and M. D. Bertness. 2002. A trophic cascade
lations in the marsh.
regulates salt marsh primary production. Proceedings of the
Thanks to Silliman’s research, a new, more complex picture
National Academy of Sciences USA 99:10500–10505.
of the salt marshes of the eastern U.S. shorelines has emerged.
Silliman, B. R., and S. Y. Newell. 2003. Fungal farming in a
Rather than the classical view of a bottom-up control on NPP,
snail. Proceedings of the National Academy of Sciences USA
where plant growth and productivity are controlled primarily
100:15643–15648.
by the physical conditions and nutrient availability, a new top-
down model of control on plant growth and productivity has 1. How does the model of trophic cascade presented in
emerged in which consumers exert a strong control on plant Figure 3 relate to the concept of indirect mutualism
structure and productivity within the ecosystem. This phenom- developed in Chapter 18 (Section 18.3)?
enon is referred to as a trophic cascade, because the effects 2. Suppose we were to introduce a top predator to this ecosys-
of predators cascade down the food web to influence primary tem that feeds on blue crabs. How might this action change
productivity (Figure 3). the nature of the trophic cascade outlined in Figure 3?
414 PART SIX • ECOSYSTEM ECOLOGY

21.12 Consumption Efficiency 9

Aquatic algae
8
Determines the Pathway of Energy

Number of observations
7
Flow through the Ecosystem 6
Although the general model of energy flow presented in Figure 5
20.18 pertains to all ecosystems, the relative importance of the 4
two major food chains and the rate of energy flow through the 3
various trophic levels can vary widely among different types of
2
ecosystems. The consumption efficiency (In/Pn - 1) defines the
1
amount of available energy produced by any given trophic level
0
(Pn - 1) that is consumed by the next-higher level (In). Values of 0 20 40 60 80 100
(a)
consumption efficiency for the various consumer trophic levels
therefore determine the pathway of energy flow through the food 9
chain, providing a basis for comparison of energy flow through Aquatic macrophytes
8

Number of observations
different ecosystems. 7
Despite its conspicuousness, the grazing food chain is not
6
the major one in most terrestrial and many aquatic ecosystems.
5
Only in some open-water aquatic ecosystems do the grazing
herbivores play the dominant role in energy flow. Ecologists 4
Helene Cyr of the University of Toronto (Canada) and Michael 3
Pace of the University of Virginia compiled published mea- 2
surements of herbivore consumption rates (herbivore consump- 1
tion efficiency), herbivore biomass, and primary productivity 0
0 20 40 60 80 100
for a wide range of aquatic and terrestrial ecosystems (Figure (b)
21.21). Although there is considerable variation in both envi-
21
ronments, some generalizations do emerge from their analysis. Terrestrial plants
Aquatic ecosystems dominated by phytoplankton have higher 18
Number of observations

rates of herbivory (median value of 79 percent) than do those

15
in which vascular plants (submerged and emergent) dominate
(median value of 30 percent). In contrast, only 17 percent of 12
primary productivity (median value) is removed by herbivores 9
in terrestrial ecosystems. Therefore, in most terrestrial and
shallow-water ecosystems, with their high standing biomass 6
and relatively low harvest of primary production by herbivores, 3
the detrital food chain is dominant. In deep-water aquatic eco-
0
systems, with their low standing biomass, rapid turnover of or- 0 20 40 60 80 100
ganisms, and high rate of harvest, the grazing food chain may Primary production removed by herbivores (%)
(c)
be dominant.
In terrestrial ecosystems, distinct differences in con- Figure 21.21 Results from a review of studies that examined
sumption efficiency and energy flow exist between for- rates of herbivory in different ecosystems. Histograms represent
est and grassland ecosystems. Nelson Hairston of Cornell the percentage of net primary productivity consumed by herbivores
University reviewed a wide range of studies that examined in ecosystems dominated by (a) algae (phytoplankton), (b) rooted
patterns of energy flow through terrestrial ecosystems, provid- aquatic plants, and (c) terrestrial plants. Number of observations refers
ing a comparison of consumption efficiencies for herbivores to the number of experiments having a given level of consumption.
(primary producer S herbivore) and their predators (herbi- Red arrows indicate the median value. Note that herbivores
vore S carnivore) The author found an average consumption consume a significantly greater proportion of phytoplankton
efficiency of 3.7 percent for herbivores inhabiting deciduous productivity than do either aquatic or terrestrial plants. Go to
at www.ecologyplace.com to perform a chi-square test.
forest ecosystems, whereas herbivores inhabiting grassland
(Nature Publishing Group.)
ecosystems had a value of 9.3 percent (both values lower than
(Adapted from Cyr and Pace 1993.)
the average for terrestrial ecosystems reported by Cyr and
Pace). Much smaller differences were observed for the con-
sumption efficiency of predators inhabiting the two ecosystem stream and river ecosystems have extremely low NPP, and the
types. Predators inhabiting forests had a value of 89.9 percent, grazing food chain is minor (see Chapter 25). The detrital food
whereas predators inhabiting grassland ecosystems had an chain dominates and depends on inputs of dead organic matter
average value of 77 percent. from adjacent terrestrial ecosystems (see Section 20.4).
Patterns of energy flow through flowing-water ecosystems Figure 21.22 graphically represents the different patterns
(streams and rivers) differ markedly from both terrestrial and of energy transfer in the four different ecosystems just dis-
standing-water ecosystems (lakes and oceans). By comparison, cussed: forest, grassland, standing water, and running water.
 CHAPTER 21 • ECOSYSTEM ENERGETICS 415

Respiration Respiration Respiration Respiration

Grazer Decomposer Grazer Decomposer

system system system system

Net primary Dead organic Net primary Dead organic

productivity matter productivity matter

(a) Forest (b) Grassland

Respiration Respiration Respiration

Respiration

Grazer Decomposer Grazer Decomposer

system system system system

Net primary Dead organic Net primary Dead organic

productivity matter productivity matter
(c) Phytoplankton community

From terrestrial catchment

(d) Stream community

Figure 21.22 General patterns of energy flow through four ecosystems: (a) forest,
(b) terrestrial grassland, (c) ocean (phytoplankton community), and (d) stream. Relative sizes of
boxes and arrows are proportional to the relative magnitude of compartments and flow.
(Adapted from Begon et al. 1986.)

21.13 Energy Decreases in Each Dry weight

(g/m2) Tertiary consumers
Successive Trophic Level 1.5 Secondary consumers
Based on the preceding discussion and the analysis presented in 11
Primary consumers
Figure 21.20, we can conclude that the quantity of energy flow- 37
ing into a trophic level decreases with each successive trophic 809 Producers
level in the food chain. This pattern occurs because not all energy
(a) Florida bog
is used for production. An ecological rule of thumb is that only
10 percent of the energy stored as biomass in a given trophic
Dry weight
level is converted to biomass at the next-higher trophic level. If, (g/m2) Consumers (zooplankton)
for example, herbivores eat 1000 kcal of plant energy, only about 21
100 kcal is converted into herbivore tissue, 10 kcal into first-level 4 Producers (phytoplankton)
carnivore production, and 1 kcal into second-level carnivore pro-
duction. However, ecosystems are not governed by some simple (b) English Channel
principle that regulates a constant proportion of energy reaching
Figure 21.23 Biomass pyramids for the consumer food chain
successive trophic levels. of (a) a bog ecosystem in Florida and (b) the marine ecosystem of the
As we have seen thus far in our discussion, differences English Channel. The pyramid for the marine ecosystem is inverted due
in the consumption efficiency as well as the efficiency of en- to the high productivity but fast turnover of phytoplankton populations
ergy conversion (assimilation and production efficiencies) exist (short life span and high rate of consumption by zooplankton).
among different feeding groups (see Table 21.2). These differ-
ences will directly influence the rate of energy transfer from considerable variation among studies and trophic levels, but the
one trophic level to the next-higher level. A measure of effi- mean value of 10.13 percent is close to the general rule of 10
ciency used to describe the transfer of energy between trophic percent transfer between trophic levels.
levels is called the trophic efficiency. The trophic efficiency An important consequence of decreasing energy transfers
(TE) is the ratio of productivity in a given trophic level (Pn) to through the food web is a corresponding decrease in the standing
the trophic level it feeds on (Pn - 1): TE = Pn/Pn - 1. biomass of organisms within each successive trophic level. If we
Daniel Pauly and Villy Christensen of the University of sum all of the biomass or energy contained in each trophic level,
British Columbia examined the energy transfer efficiency re- we can construct pyramids for the ecosystem (Figure 21.23).
ported in 48 different studies of aquatic ecosystems. There is The pyramid of biomass indicates by weight, or other means of
416 PART SIX • ECOSYSTEM ECOLOGY

measuring living material, the total bulk of organisms or fixed en- This arrangement does not hold for all ecosystems. In such
ergy present at any one time—the standing crop. Because some ecosystems as lakes and open seas, primary production is concen-
energy or material is lost at each successive trophic level, the total trated in the phytoplankton. These microscopic organisms have a
mass supported at each level is limited by the rate at which en- short life cycle and rapid reproduction. They are heavily grazed
ergy is being stored at the next-lower level. In general, the bio- by herbivorous zooplankton that are larger and longer-lived.
mass of producers must be greater than that of the herbivores they Thus, despite the high productivity of algae, their biomass is low
support, and the biomass of herbivores must be greater than that compared to that of zooplankton herbivores (Figure 21.23b). The
of carnivores. That circumstance results in a narrowing pyramid result is an inverted pyramid, with a lower standing biomass of
for most ecosystems (Figure 21.23a). primary producers (phytoplankton) and herbivores (zooplankton).

Summary
Laws of Thermodynamics 21.1 External Inputs 21.5
Energy flow in ecosystems supports life. Energy is governed by In many aquatic ecosystems a substantial proportion of
the laws of thermodynamics. The first law states that although organic carbon is derived from dead organic matter from
energy can be transferred, it cannot be created or destroyed. adjacent terrestrial ecosystems. The relative importance of
The second law states that as energy is transferred, a portion external sources of organic carbon varies widely among dif-
ceases to be usable. As energy moves through an ecosystem, ferent aquatic ecosystems. In large rivers, lakes, and most
much of it is lost as heat of respiration. Energy is degraded marine systems, the majority of organic carbon is derived
from a more organized to a less organized state, or entropy. internally from photosynthesis by autotrophs. In contrast, in
However, a continuous flux of energy from the Sun prevents smaller streams and lakes the dominant source is often exter-
ecosystems from running down. nal sources of organic carbon.

Primary Production 21.2 Energy Allocation 21.6

The flow of energy through an ecosystem starts with the har- Energy fixed by plants is allocated to different parts of the plant
nessing of sunlight by green plants through a process referred and to reproduction. How much is allocated to each component
to as primary production. The total amount of energy fixed is a function of the plant life-form as well as the environmen-
by plants is gross primary production. The amount of energy tal conditions. The pattern of allocation will directly influence
remaining after plants have met their respiratory need is net standing biomass and productivity rate.
primary production in the form of plant biomass. The rate of
primary production is net primary productivity, which is mea- Temporal Variation 21.7
sured in units of weight per unit area per unit time. Primary production in an ecosystem varies with time. Seasonal
and yearly variations in moisture and temperature directly
Terrestrial Ecosystems 21.3 influence primary production. In ecosystems dominated by
Productivity of terrestrial ecosystems is influenced by climate, woody vegetation, net primary production declines with age.
especially temperature and precipitation. Temperature influ- As the ratio of woody biomass to foliage increases, more of
ences the photosynthetic rate and the amount of available water gross production goes into maintenance.
limits photosynthesis and the amount of leaves that can be sup-
ported. Warm, wet conditions make the tropical rain forest the Secondary Production 21.8
most productive terrestrial ecosystem. Nutrient availability also Net primary production is available to consumers directly as
directly influences rates of primary productivity. plant tissue or indirectly through animal tissue. Once consumed
and assimilated, energy is diverted to maintenance, growth, and
Aquatic Ecosystems 21.4 reproduction, and to feces, urine, and gas. Change in biomass,
Light is a primary factor limiting productivity in aquatic eco- including weight change and reproduction, is secondary pro-
systems, and the depth to which light penetrates is crucial to duction. Secondary production depends upon primary produc-
determining the zone of primary productivity. Nutrient avail- tion. Any environmental constraint on primary production will
ability is the most pervasive influence on the productivity of constrain secondary production in the ecosystem.
oceans. The most productive ecosystems are shallow coastal
waters, coral reefs, and estuaries, where nutrients are more Efficiency of Energy Use 21.9
available. Nutrient availability is also a dominant factor limit- Efficiency of production varies. Endotherms have high assimi-
ing net primary productivity in lake ecosystems. In rivers and lation efficiency but low production efficiency because they
streams, net primary productivity is low, with inputs of dead have to expend so much energy in maintenance. Ectotherms
organic matter from adjacent terrestrial ecosystems being an have low assimilation efficiency but high production efficiency;
important source of energy input. they put more energy into growth.
 CHAPTER 21 • ECOSYSTEM ENERGETICS 417

Food Chains and Energy Flow 21.10 These estimates of efficiency can be used to quantify the flow
A basic function of the ecosystem is to move energy from the of energy through the food chain.
Sun through various consumers to its final dissipation in a se-
ries of energy transfers known as the food chain. The various Consumption Efficiency 21.12
members of a food web can be grouped into categories called Consumption efficiency determines the flow of energy through
trophic or feeding levels. Autotrophs occupy the first trophic the ecosystem. The detrital food chain dominates in terrestrial
level. Herbivores that feed on autotrophs make up the next tro- ecosystems, with only a small proportion of net primary pro-
phic level. Carnivores that feed on herbivores make up the third ductivity being consumed by herbivores. In open-water ecosys-
and higher trophic levels. tems, such as lakes and oceans, a greater proportion of primary
Energy flow in ecosystems takes two routes: one through productivity is consumed by herbivores. Consumption effi-
the grazing food chain, the other through the detrital food ciency of predators is more similar among these ecosystems.
chain. The bulk of production is used by organisms that feed
on dead organic matter. The two food chains are linked by the Energy Pyramids 21.13
input of dead organic matter and wastes from the consumer The quantity of energy flowing into a trophic level decreases
food chain functioning as input into the detrital food chain. with each successive trophic level in the food chain. This pat-
tern occurs because not all energy is used for production. An
Quantifying Energy Flow 21.11 ecological rule of thumb is that only 10 percent of the energy
At each trophic level, estimates of the efficiency of energy ex- stored as biomass in a given trophic level is converted to bio-
change are defined as consumption efficiency, the proportion mass at the next-higher trophic level. A plot of the total weight
of available energy being consumed; assimilation efficiency, of individuals at each successive level produces a tapering pyr-
the portion of energy ingested that is assimilated and not lost amid. In aquatic ecosystems, however, where there is a rapid
as waste material; and production efficiency, the portion of as- turnover of small aquatic producers, the pyramid of biomass
similated energy that goes to growth rather than respiration. becomes inverted.

Study Questions
1. How do the concepts of community and ecosystem 7. How does primary productivity function as a constraint
differ? on secondary productivity in ecosystems?
2. Relate the following terms: gross primary productivity, 8. What does the top-down model of food chain structure
autotrophic respiration, and net primary productivity. imply about the role of secondary producers in control-
3. Contrast net primary productivity and standing bio- ling net primary productivity and standing biomass
mass for an ecosystem. within ecosystems?
4. How do temperature and precipitation interact to 9. How do assimilation efficiency and production ef-
influence net primary productivity in terrestrial ficiency relate to the flow of energy through a trophic
ecosystems? level?
5. How does net primary productivity vary with water 10. What is the difference in energy allocation and produc-
depth in standing-water ecosystems (lakes and tion efficiency between endotherms and ectotherms?
oceans)? What is the basis for the vertical profile of net 11. What are the two major food chains, and how are they
primary productivity in these ecosystems? related?
6. What environmental factors might influence the (light) 12. How does consumption efficiency differ between ter-
compensation depth of a lake ecosystem? restrial and marine ecosystems?

Further Readings
Aber, J. D., and J. M. Melillo. 2001. 2nd edition. Terrestrial Gates, D. M. 1985. Energy and ecology. Sunderland, MA:
ecosystems. Philadelphia: Saunders College Publishing. Sinauer Associates.
This well-written and illustrated text gives readers an excellent This text shows the function of ecological systems in terms of
introduction to ecosystem ecology. energy flow and covers a range of topics from photosynthesis to
Chapin, F. S., P. A. Matson, and H. A. Mooney. 2002. Principles ecosystem productivity.
of terrestrial ecosystem ecology. New York: Springer-Verlag. Golley, F. B. 1994. A history of the ecosystem concept in
This well-written and illustrated text is aimed at a more ecology. New Haven, CT: Yale University Press.
advanced readership but provides an excellent reference for This book presents a historical overview of the development of
exploring topics covered in Part Six in more depth. the study of ecosystems in the broader framework of ecology.