ECOLOGY

Primary Production and Energy Flow

How do I become
more productive?
 Ecosystem Ecology
The term ecosystem first proposed by Arthur
Tansley (1935):
Though the organisms may claim our primary interest,we cannot
separate them from their special environment, with which they form
one system. It is the [eco]systems so formed which, from the point
of view of the ecologist, are the basic units of nature on the face of
the earth.
Ecosystem Ecology: The study of energy, water
and nutrient flows (flux) in ecosystems:
Biotic and abiotic processes

Fundamental areas of interest include primary

production, energy flow and nutrient cycling

 Definitions

Population: a collection of individuals of the

same species
Community: an assemblage of populations
in an area or habitat
Ecosystem: a biological community plus all
of the abiotic factors influencing that
community
 Definitions
Production: creation of new organic matter

Primary Production (PP): fixation of energy by autotrophs

PP measured in various ways:
2 1
usu. rate of CO2 uptake (g/m yr )

Or -- biomass or O2 produced

Gross Primary Production (GPP): total amount of energy

fixed (or CO2 taken up) by all autotrophs in an ecosystem.

Net Primary Production (NPP): amount of energy left over

after autotrophs have met their own energetic needs
(respiration and, ie, the amount of energy available to
consumers).
NPP = GPP RPP
 Trophic Dynamics
Lindeman (1942): The Trophic Dynamic Aspect of
Ecology
Trophic dynamics: transfer of energy from one

part of an ecosystem to another

First suggested grouping organisms within an

ecosystem into trophic levels.

The number of trophic levels in an ecosystem is

limited by energy losses with each transfer or
conversion of energy between trophic levels
 Trophic Dynamics
Each T.L. feeds on T.L. immediately below.
As energy is transferred from one T.L to another,
energy is degraded/lost due to:
Limited assimilation

Consumer respiration

Heat production

Energy quantity decreases with each

successive trophic level

A Simplified Food Web

Secondary Consumers
(Predators)

Primary Consumers
(Herbivores)

Primary Producers
(Plants)
A Simplified Food Web

Secondary Consumers
(Predators) PAR

Primary Consumers
(Herbivores)

Primary Producers
(Plants)
A Simplified Food Web

Secondary Consumers
(Predators) PAR

Primary Consumers
(Herbivores)

reflected

Primary Producers
absorbed
(Plants) by Chl a
 A Simplified Food Web

Secondary Consumers
(Predators) PAR

Primary Consumers
(Herbivores)

reflected

photorespiration Primary Producers

absorbed
heat production (Plants) by Chl a
 A Simplified Food Web

Secondary Consumers
(Predators) PAR

Primary Consumers
(Herbivores)

reflected

photorespiration Primary Producers

absorbed
heat production (Plants) by Chl a
 A Simplified Food Web

Secondary Consumers
(Predators) PAR

respiration Primary Consumers

heat production
(Herbivores)

limited assimilation
reflected

photorespiration Primary Producers

absorbed
heat production (Plants) by Chl a
 A Simplified Food Web

Secondary Consumers
(Predators) PAR

respiration Primary Consumers

heat production
(Herbivores)

limited assimilation
reflected

photorespiration Primary Producers

absorbed
heat production (Plants) by Chl a
Hence the Trophic Pyramid

Biomass and productivity

generally decrease with
trophic level
Energy Flow In A Temperate Deciduous Forest
Gosz et al. (1978): study of solar energy flux at
Hubbard Brook experimental forest:
Of the total energy input via solar radiation:

15% reflected

41% converted to heat.

42% absorbed during evapotranspiration.

2.2% fixed by plants as GPP

1.2% used in plant respiration.

1% left for NPP

Energy Flow In A Temperate Deciduous Forest
Gosz et al. 1978:
< 1% of total energy input converted to NPP

Of the NPP available to consumers (herbivores),

96% lost as consumer respiration

Insufficient energy left to support a viable

population at a 3rd trophic level.

 Factors Controlling Terrestrial PP
PP variable but most strongly correlated with temperature
and moisture.
Highest PP under warm, moist conditions

An indicator of PP is actual evapotranspiration (AET):

Total amount of water that evaporates and transpires off a
landscape during the course of a year (mm H2O/yr)
affected by temp and precipitation

ecosystems with high AET tend to be warm, receive

large amounts of precipitation
ecosystems with low AET tend to be cold, receive little
precipitation or both
eg hot deserts and cold tundra have low AET
 Among different types of
ecosystems, AET is correlated
with NPP

Rosenzweig (1968) estimated

influence of moisture and
temperature on rates of PP by
plotting the relationship
between annual NPP and AET.
 Factors Controlling Terrestrial PP
Within similar ecosystems, temps tend to be similar, so
moisture (precipitation) tends to be the controlling factor

eg Sala et al. (1988) study of

9,498 sites in the grasslands
of central North America:
 Factors Controlling Terrestrial PP
Patterns of soil fertility also explain significant variation in
NPP within terrestrial ecosystems (assuming equal temp.
and moisture)
Liebig (1840): concept of nutrient limitation

Nutrient availability
controls patterns of PP
in agricultural
ecosystems...

NB: Liebigs law somewhat simplistic:

2 or more factors may be simultaneously
limiting in many systems Liebigs Law of the Minimum
 Factors Controlling Terrestrial PP
...and in natural ecosystems:
Shaver and Chapin (1986):
Added commercial fertilizer (N+P+K) to several tundra systems in

Alaska
NPP 23 30% higher on fertilized plots.
 Factors Controlling Terrestrial PP

Effects of nutrient additions depend on prior nutrient availability

PP responds to additions of limiting nutrients

eg Bowman et al. (1993):

Experimental fertilization of wet and dry alpine meadows,

Niwot Ridge, CO.

Wet meadows had higher initial [N], [P]

4 treatments:

N; P; N+P; control
 Factors Controlling Terrestrial PP
Bowman et al. (1993):
Results more dramatic in dry

meadow (lower initial [N], [P])

N, N+P both produced

signif. biomass *
In contrast, weaker response in

wet meadow
only N+P had an effect
*

sugg. N-limitation in dry meadow; *

co-limitation in wet meadow

Light might also limit NPP in wet

meadow; biomass might
produce enough shading to
inhibit growth response to nutrient
additions. * = statistically significant response
 Factors Controlling Aquatic PP
Phytoplankton are the dominant primary producers
in aquatic ecosystems

Aquatic NPP generally limited by nutrient

availability
temperatures generally less variable in the
ocean than on land

Several studies have found proportional

relationship between [P] and phytoplankton
biomass, chlorophyll a and NPP in lakes.
Observational
studies in Japan,
North America:

eg:
Hogetsu and Ichimura 1954
Ichimura 1956
Sakamoto 1966
Dillon and Rigler 1974
Smith 1979
 Lake Fertilization Experiments
eg. studies at Experimental Lakes Area, Ontario (Mills and Schindler
1987, Findlay and Kasian 1987):
Lake 226 divided by vinyl curtain
Each basin = 8 ha, 500,000 m
3

One side fertilized with P:

 Marine NPP is also controlled by nutrient availability

Highest rates of NPP in areas with greatest nutrient availability:

continental margins (runoff from land, bioturbation of bottom sediments)

areas of upwelling (deep nutrient-laden waters rise to euphotic zone)

Open ocean tends to be nutrient poor (relies on vertical mixing for nutrients)
 Unlike lakes, marine NPP appears
to be limited primarily by [N]:

Granli et al. (1990) nutrient enrichment

experiments, Baltic Sea:
Fertilized flasks containing indigenous

phytoplankton spp.
N, P, control treatments

N treatments led to increased

chlorophyll concentrations.
 Factors Controlling NPP
Temperature and precipitation (terrestrial)
and nutrient availability (aquatic) explain
most of the variation seen in NPP, but not
all:
Residual Variation: proportion of variation
not explained by the independent
variable.
Dillon and Rigler (1974) suggested
environmental factors besides nutrient
availability significantly influence
phytoplankton biomass.
 Summary

Terrestrial Primary Production is generally

limited by temperature and moisture.
Aquatic Primary Production is generally
limited by nutrient availability.
usually P in freshwater ecosystems

N in marine ecosystems

Energy losses limit the number of trophic

levels found in ecosystems.