+

) BIOL 207 students:

 One more Make up session: April 3rd
(Thursday)
 Exam: April 10th (Thursday)
PDHP 212 students:
 Make up session: April 4th (afternoon)
 Exam: April 11th (Friday)
+

Robert E. Ricklefs

The Economy of Nature, Fifth Edition Chapter 6: Energy in the

Ecosystem

2
+ 3

Background: Organizing Concepts

 In 1920s, English ecologist Charles Elton and others promoted a

revolutionary concept:
 organisms living in the same place not only have similar tolerances of
physical factors, but
 feeding relationships link these organisms into a single functional entity

 This system of feeding relationships is called a food web.

+ 4

The Ecosystem Concept

 The English ecologist A.G. Tansley took Elton’s ideas one step
further:
 in 1935 Tansley coined the term ecosystem, the fundamental unit of
ecological organization
 the ecosystem concept: “the biological and physical parts of nature
together, unified by the dependence of animals and plants on their
physical surroundings and by their contributions to maintaining the
conditions and composition of the physical world.”
-R.E. Ricklefs
+ 5

Alfred J. Lotka, the Thermodynamic

Concept, and Lindeman’s concept
 Alfred J. Lotka introduced the concept of the ecosystem
as an energy-transforming machine:
 described by a set of equations representing exchanges of matter
and energy among components, and
 obeying thermodynamic principles that govern all energy
transformations
 In 1942, Raymond Lindeman brought Lotka’s ideas of the ecosystem
as an energy-transforming machine to the attention of ecologists. He
incorporated:
 Lotka’s thermodynamic concepts
 Elton’s concept of the food web as expression of the ecosystem’s structure
 Tansley’s concept of the ecosystem as the fundamental unit in ecology
+ 6

Lindeman’s Foundations of Ecosystem

Ecology
 The ecosystem is the fundamental unit of ecology.
 Within the ecosystem, energy passes through many steps or links
in a food chain.
 Each link in the food chain is a trophic level (or feeding level).
 Inefficiencies
in energy transformation lead to a pyramid of
energy in the ecosystem.
+
An ecological pyramid of energy
+ 8

Odum’s Energy Flux Model

 Eugene P. Odum popularized ecology to a generation of ecologists.
 Odum further developed the emerging framework of ecosystem
ecology:
 he recognized the utility of energy and masses of elements as common
“currencies” in comparative analysis of ecosystem structure and function

 Odum extended his models to incorporate nutrient cycling.

 Fluxes of energy and materials are closely linked in ecosystem function.
However, they are fundamentally different:
 energy enters ecosystems as light and is degraded into heat
 nutrients cycle indefinitely, converted from inorganic to organic forms and
back again
 Studies of nutrient cycling provide an index to fluxes of energy.
+ 9

Primary Production: reminder

 Primary production is the process whereby plants,algae, and some bacteria (primary
producers) capture the energy of light and transform it into the energy of chemical bonds in
carbohydrate:
 its rate is referred to as primary productivity
 6CO2 + 6H2O  C6H12O6 + 6O2
 for each g of C assimilated, 39 kJ energy stored

 The rate of primary production determines the rate of energy supply to the rest of the
ecosystem:
 gross primary production = total energy assimilated by primary producers
 net primary production = energy accumulated (in stored form) by primary
producers
 gross - net = respiration, the energy consumed by producers for
maintenance and biosynthesis
+ 10

Measurement of Primary Production 1

 How much energy has been accumulated by net production?

 harvest techniques determine dry mass accumulated (net
production)
 gas exchange techniques determine net uptake of CO2 in light
(net production), production of CO2 in dark (respiration) and
gross production as their sum
 Radioactive carbon (14C) may also determine net uptake of
carbon by plants
+ 11

Measurement of Primary Production 2

 Aquatic systems pose special problems:

 harvest approach is not practical for small organisms, such as
phytoplankton
 carbon is too abundant for practical measurement of small
changes

 Alternatives for aquatic systems:

 light and dark bottles may be used to determine changes in
O2
 C approach may also be used in unproductive waters
14
+
Measurements of Carbon Dioxide flux in dark and light can
provide an estimate of GPP
+ 13

Effects of Light and Temperature 1

 Plants are not usually light-limited in full sun.

 Shading (by other leaves or plants) may reduce photosynthetic rate below its
maximum.

 Overall, photosynthetic efficiency of the ecosystem is typically 1-2%:

 remaining energy is either reflected or absorbed and dissipated
 Leaves reflect 25 to 75%
 Molecules other than photosynthetic pigments absorb remainder – converted to heat and
radiated, or conducted across leaf surface, or transpired
 Photosynthetic efficiency
 Percentage of the energy in sunlight that is converted to net primary production during
the growing season
+ 14

Effects of Light and Temperature 2

 Optimum temperature for photosynthesis varies with

system:
 about 16oC for many temperate species
 as high as 38oC for some tropical species

 Rate of photosynthesis increases with temperature, up

to a point:
 rate of respiration also increases with temperature
 net assimilation may thus decrease at high temperatures
+ 15

Water limits primary production

(reminder)
 Photosynthesis in terrestrial systems is water-limited:
 under water stress, stomates close and gas exchange ceases, stopping
photosynthesis

 Transpiration or water-use efficiency:

 typically 2g production per kg of water transpired (4g for drought-tolerant
crops)
 ecosystem-level efficiency may be an order of magnitude poorer (0.2 g/kg)
 Most precipitation is not taken up by plants
+ 16

Nutrients stimulate primary production

– terrestrial and aquatic.
 Terrestrial production may be nutrient -limited:
 fertilizers stimulate crop production
 N is the most common limiting element

 Aquatic systems are often strongly nutrient -limited:

 especially true of open ocean
 inadvertent addition of nutrients may stimulate unwanted
production
+ 17

Primary production varies among

ecosystems.
 Primary
production is maximum under favorable
combinations of:
 intense sunlight
 warm temperatures
 abundant rainfall
 ample nutrients

 Onland, production is highest in humid tropics, lowest in

tundra and desert.
+ NPP among ecosystems
+ 19

Only 5% to 20% of energy passes between

trophic levels.
 Energy reaching each trophic level depends on:
 net primary production (base of food chain)
 efficiencies of transfers between trophic levels
 Plant
use between 15% and 70% of light energy
assimilated for maintenance – thus that portion is
unavailable to consumers
 Herbivoresand carnivores expend more energy on
maintenance than do plants: production of each
trophic level is only 5% to 20% that of the level
below it.
+ 20

Ecological Efficiency
 Ecological efficiency
(food chain efficiency)  el
is the percentage of
energy transferred from
one trophic level to the
next:
 range of 5% to 20% is
typical, as we’ve seen
 to understand this more
fully, we must study the
use of energy within a
trophic level

Undigested plant fibers in elephant dung

+ 21

Intratrophic Energy Transfers

 Intratrophic transfers involve several components:

 ingestion (energy content of food ingested)
 egestion (energy content of indigestible materials
regurgitated or defecated) (the elephant dung)
 assimilation (energy content of food digested and absorbed)
 excretion (energy content of organic wastes)
 respiration (energy consumed for maintenance)
 production (residual energy content for growth and
reproduction)
+ 22

Fundamental Energy Relationships

 Components of an animal’s energy budget are related

by:
 ingested energy - egested energy = assimilated energy
 assimilated energy - respiration - excretion = production
+ 23

Assimilation Efficiency

 Assimilation efficiency = assimilation/ingestion

 primarily a function of food quality:
 seeds: 80%
 young vegetation: 60-70%
 plant foods of grazers, browsers: 30-40%
 decaying wood: 15%
 animal foods: 60-90%
+ 24

Net Production Efficiency

 Net production efficiency = production/assimilation

 depends largely on metabolic activity:
 birds: <1%
 small mammals: <6%
 sedentary, cold-blooded animals: as much as 75%

 Gross production efficiency = assimilation efficiency

x net production efficiency = production/ingestion,
ranges from below 1% (birds and mammals) to >30%
(aquatic animals).
+ Active, warm-blooded animals – low net production
efficiencies; hummingbird: <1%
+ 26

Production Efficiency in Plants

 The concept of production efficiency is somewhat

different for plants because plants do not digest
and assimilate food:
 net production efficiency = net production/gross
production; varies between 30% and 85%
 rapidly growing plants in temperate zone have net
production efficiencies of 75-85%; their counterparts in
the tropics are 40-60% efficient
+ 27

Detritus Food Chains

 Ecosystems support two parallel food chains:
 herbivore-based (relatively large animals feed on
leaves, fruits, seeds)
 detritus-based (microorganisms and small animals
consume dead remains of plants and indigestible
excreta of herbivores)
 herbivores consume:
 1.5-2.5% of net primary production in temperate forests
 12% in old-field habitats
 60-99% in plankton communities
+ 28

Exploitation Efficiency

 When production and consumption are not balanced,

energy may accumulate in the ecosystem (as organic
sediments).
 Exploitation efficiency = ingestion by one trophic
level/production of the trophic level below it.
 Tothe extent that exploitation efficiency is <100%,
ecological efficiency = exploitation efficiency x gross
production efficiency.
+ 29

Energy moves through ecosystems at

different rates.

 Otherindices address how rapidly energy cycles through an

ecosystem:
 residence time measures the average time a packet of energy resides in
storage:
 residence time (yr) = energy stored in biomass/net productivity

 biomass accumulation ratio is a similar index based on biomass rather than

energy:
 biomass accumulation ratio (yr) = biomass/rate of biomass production
 + 30

Biomass Accumulation Ratios

 Biomass accumulation ratios
become larger as amount of stored
energy increases:
 humid tropical forests have net
production of 1.8 kg/m2/yr and
biomass of 43 kg/m2, yielding
biomass accumulation ratio of
23yr
 ratios for forested terrestrial
communities are typically >20 yr
 ratios for planktonic aquatic
ecosystems are <20 days
+ 31

Residence Time for Litter

 Decomposition of litter is dependent on conditions of

temperature and moisture.
 Index is residence time = mass of litter accumulation/rate
of litter fall:
 3 months in humid tropics
 1-2 yr in dry and montane tropics
 4-16 yr in southeastern US
 >100 yr in boreal ecosystems
+ 32

Ecosystem Energetics

 Comparative studies of ecosystem energetics now exist for

various systems.
 Many systems are supported mainly by autochthonous
materials (produced within system).
 Some ecosystems are subsidized by input of allochthonous
materials (produced outside system).
 autochthonous production dominates in large rivers, lakes, marine ecosystems
 allochthonous production dominates in small streams, springs, and caves
(100%)
 + 33

Cedar Bog Lake

 Lindeman’s study of a small lake
in Minnesota uncovered
surprisingly low exploitation
efficiencies:
 herbivores: 20%
 carnivores: 33%
 residual production of plants and
herbivores accumulates as
bottom sediment
+ 34

Some General Rules

 Assimilation efficiency increases at higher trophic

levels.
 Netand gross production efficiencies decrease at
higher trophic levels.
 Ecological efficiency averages about 10%.
 About1% of net production of plants ends up as
production on the third trophic level: the pyramid of
energy narrows quickly.
 Toincrease human food supplies means eating lower
on food chain!
+ 35

Summary 1

 Ecosystems are giant energy-transforming machines;

energy enters ecosystems through production by
plants and is transferred to animals and heterotrophic
microorganisms through food chain and detrital
pathways.
 Ourunderstanding of ecosystem function was greatly
enhanced by the pioneering work of Elton, Tansley,
Lotka, Lindeman, and Odum.
+ 36

Summary 2

 Primary production is the foundation of energy flux

through ecosystems. Primary production varies
widely among systems because of differences in
light, temperature, precipitation, and availability of
nutrients.
 Secondary production results from food chain
processing of energy by animals. We monitor energy
fluxes by considering various efficiencies and the
residence times of energy and materials.
+
37

As usual

Do the quiz for Chapter 6. Due by next class period.