BIOLS340

CH18: Primary & secondary production

Done by: Yousif Adel
-A world of green: the workings of the biosphere depend on the
production of energy-rich biomass by photosynthetic primary producers
such as green plants

Introduction
-The interactions between organisms and their environments are fueled
by complex fluxes and transformations of energy
-Carbon fixation: forest plants use photosynthetically active radiation
(PAR) to convert CO2 into sugars and other forms of biomass
-Some chemical energy used for: Metabolism, Growth
-Some chemical energy is consumed by: Herbivores, Detritivores
-Some chemical energy (biomass) ends up as soil organic matter

-In most ecosystems, sunlight provides the ultimate source of energy to

power all biological activity, such as the frog and the growth of the plant
on which it sits

-Ecosystem: a biological community plus all of the abiotic factors

influencing that community. (Ecosystem ecologists study the flow of
energy, water and nutrients in ecosystems)
-Primary production: the production of new organic matter or biomass,
by autotrophs in an ecosystem during some period of time

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  Gross primary production: the total amount of biomass
produced by all the autotrophs in the ecosystem
 Net primary production: the amount of biomass left over
after autotrophs have met their own energetic needs
-Secondary production: the production of biomass by heterotrophic
consumer organisms feeding on plants, animals, microbes, fungi or
detritus during some period of time
-Ecologists have measured primary production mainly as the rate of
carbon uptake by primary producers or by the amount of biomass or
oxygen produced
-Trophic Level: a position in a food web and is determined by the
number of transfers of energy from primary producers to that level

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 18.1: Terrestrial primary production patterns
 Which limits terrestrial primary production?
1.Temperature
2.Moisture
3.Nutrients
-The variables most highly correlated with variation in terrestrial
primary production are temperature and moisture. Highest rates of
terrestrial primary production generally occur under warm, moist
conditions

Actual evapotranspiration & terrestrial primary production

-Rosenzweig (1968) estimated influence of moisture and temperature on
rates of primary production by plotting relationship between annual net
primary production and annual actual evapotranspiration (AET)
-AET: annual amount of water that evaporates and transpires off a
landscape during a period of one year
 Cold dry ecosystems tend to have low AET
 Sala (1988): what controls variation in primary production within
similar ecosystem (grassland)?
 Sala found that east-west variation in primary production among
grassland ecosystems correlated significantly with the amount of
rainfall
-Generally, there is a positive relationship between net primary
production and AET
-The researchers found strong correlation between AET or precipitation

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and rates of terrestrial primary production

Soil fertility & terrestrial primary production

 How we can explain the significant variation in terrestrial primary
production?
 By differences in soil fertility
-Justus Liebig (1840): pointed out that nutrient supplies often limit plant
growth
-Liebig’s Law of the Minimum: nutrient limitation to plant growth
could be traced to a single limiting nutrient
-Several factors, including a number of nutrients, simultaneously affect
levels of terrestrial primary production
-Shaver and Chapin (1986) fertilization experiments by addition of
commercial fertilizers containing nitrogen, phosphorus and potassium to
arctic tundra in Alaska → increased net primary production by 23-300%

-Bowman et al. (1993) added nutrients to tundra in Colorado. Nutrients

were added in the following combination:
 Control (no nutrients added)
 Nitrogen added
 Phosphorus added
 Nitrogen and phosphorus added
 Results showed variable increase in net primary production

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 18.2: Aquatic primary production patterns
-Studies on lake ecosystems in Japan (Hogetsu and Ichimura, 1954;
Ichimura, 1956; Sakamoto, 1966) and in Northern Hemisphere (Dillon
and Rigler,1974) described a positive relationship between phosphorous
and phytoplankton biomass
 What is the relationship between phytoplankton biomass and the
rate of primary production?
 Studies on lakes by Val Smith (1979) showed a strong positive
correlation between chlorophyll concentrations and photosynthetic
rates
-Whole Lake Experiments on Primary Production:
-Experimental lake in Canada was divided in halves and each half was
fertilized separately from 1973-1980
-Results show that nutrient availability controls rates of primary
production in freshwater ecosystems

Marine primary production global patterns

-Oceanographers have observed that the highest rates of primary
production by marine phytoplankton are concentrated in areas with
higher levels of nutrient availability
-Highest rates of primary production:
 Margins of continents: runoff, sediment disturbance
 Upwelling areas: deep water currents

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-Low rates of primary production:
 Open oceans: vertical mixing (blocked in tropics by
thermocline = temperature changes rapidly with depth)

-Granéli (1990) lab culture experiments using single species showed that
addition of nitrate increased chlorophyll a concentrations at all sites
-Repeating the above experiments using indigenous species in another
site gave similar results
 N appears to be limiting nutrient

-Residual variation: that proportion of variation not explained by the

independent variable (nutrient concentration)
-Dillon and Rigler (1972) suggested that environmental factors besides
nutrient availability significantly influence phytoplankton biomass
-One of those factors is the intensity of predation on the zooplankton
that feed on phytoplankton

18.3: Primary producer diversity

-Primary producer diversity contributes to higher primary production
-Tilman et al. (2001) examined plant functional groups, plants with
similar physiological and anatomical characteristics
 Manipulated number of species and functional groups that
were present in 168 plots

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  Higher species richness resulted in higher primary production.
Plots with 16 species had 3x the production of single species
plots
 The functional groups with N-fixing legumes and C4 grasses
had higher productivity

18.4: Consumer influences

-Consumers can influence rates of primary production in aquatic and
terrestrial ecosystems through trophic cascades
-Bottom-up controls: when ecologists refer to the influence of physical
and chemical factors, such as temperature and nutrients on ecosystems
-Top-down controls: the influences of consumers on ecosystems

Lake primary production

-Carpenter et al. (1985) proposed piscivores (eats primarily fish) and
planktivorous fish can cause significant deviations in primary
productivity
-Carpenter and Kitchell (1993) proposed the influence of
consumers on lake primary productivity propagate through food
webs
 Trophic Cascade Hypothesis - effects of predators on prey
can alter more than one trophic level

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The trophic cascade
hypothesis, a result of
“cascading” indirect
interactions

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Predicted effects of piscivores on planktivore, herbivore, &
phytoplankton biomass and production

-Carpenter and Kitchell (1993)

 Reduction in planktivorous fish populations led to reduced
rates of primary production
 In absence of planktivorous small, predaceous invertebrates
became more numerous
 In presence of abundant, large herbivorous zooplankton,
phytoplankton biomass and rate of primary production
declined

18.5: Secondary production

-Lindeman (1942) concluded the ecosystem concept is fundamental to
the study of energy transfer within an ecosystem (trophic dynamics)
-Suggested grouping organisms within an ecosystem into trophic
levels
-Each feeds on level immediately below
 As energy is transferred from one trophic level to
another, energy is degraded

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  Ecological Efficiency is the percentage of energy
transferred from one trophic level to the one
above it. Varies from about 5-20%

Trophic dynamic view of ecosystems

-As energy is transferred from one trophic level to another, energy is
degraded:
 Limited assimilation
 Consumer respiration
 Heat production
-Energy quality decreases with each successive trophic level
- Pyramid shaped energy distribution

Linking primary & secondary production

-Predictions of the relationship between primary and secondary
production depend on which predator-prey model is used

The prey-dependent model The ratio-dependent model

Predicts that increased primary
production will lead to an increase
in secondary consumer (predators)
abundance but no change in
primary consumers (herbivores)
Predicts that increased primary
production will lead to an increase
in both primary and secondary
consumers

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Contrasting predictions by prey-dependent and ratio-dependent models
of functional response. While prey-dependent models predict varying
responses to enrichment by different trophic levels, ratio-dependent
models predict proportional increases in abundance at all trophic levels
regardless of the number of trophic levels

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