1. Ecosystems and Restoration Ecology
Chapter 55
Ecosystems and Restoration Ecology

2. Overview: Cool Ecosystem
An ecosystem consists of all the organisms living in a community, as well as the abiotic factors with which they interact
An example is the unusual community of organisms, including chemoautotrophic bacteria, living below a glacier in Antarctica
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3. Figure 55.1
Figure 55.1 Why is this Antarctic ice blood red? 3

4. Ecosystems range from a microcosm, such as an aquarium, to a large area, such as a lake or forest
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5. Figure 55.2
Figure 55.2 A desert spring ecosystem. 5

6. Energy flows through ecosystems, whereas matter cycles within them
Regardless of an ecosystem’s size, its dynamics involve two main processes: energy flow and chemical cycling
Energy flows through ecosystems, whereas matter cycles within them
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7. Concept 55.1: Physical laws govern energy flow and chemical cycling in ecosystems
Ecologists study the transformations of energy and matter within ecosystems
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8. Conservation of Energy
Laws of physics and chemistry apply to ecosystems, particularly energy flow
The first law of thermodynamics states that energy cannot be created or destroyed, only transformed
Energy enters an ecosystem as solar radiation, is conserved, and is lost from organisms as heat
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9. The second law of thermodynamics states that every exchange of energy increases the entropy of the universe
In an ecosystem, energy conversions are not completely efficient, and some energy is always lost as heat
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10. Conservation of Mass
The law of conservation of mass states that matter cannot be created or destroyed
Chemical elements are continually recycled within ecosystems
In a forest ecosystem, most nutrients enter as dust or solutes in rain and are carried away in water
Ecosystems are open systems, absorbing energy and mass and releasing heat and waste products
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11. Energy, Mass, and Trophic Levels
Autotrophs build molecules themselves using photosynthesis or chemosynthesis as an energy source
Heterotrophs depend on the biosynthetic output of other organisms
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12. Energy and nutrients pass from primary producers (autotrophs) to primary consumers (herbivores) to secondary consumers (carnivores) to tertiary consumers (carnivores that feed on other carnivores)
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13. Prokaryotes and fungi are important detritivores
Detritivores, or decomposers, are consumers that derive their energy from detritus, nonliving organic matter
Prokaryotes and fungi are important detritivores
Decomposition connects all trophic levels
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14. Figure 55.3
Figure 55.3 Fungi decomposing a dead tree. 14

15. Microorganisms and other detritivores Secondary and tertiary consumers
Figure 55.4
Sun
Key
Chemical cycling
Energy flow
Heat
Primary producers
Primary consumers
Detritus
Figure 55.4 An overview of energy and nutrient dynamics in an ecosystem.
Microorganisms and other detritivores
Secondary and tertiary consumers 15

16. Concept 55.2: Energy and other limiting factors control primary production in ecosystems
In most ecosystems, primary production is the amount of light energy converted to chemical energy by autotrophs during a given time period
In a few ecosystems, chemoautotrophs are the primary producers
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17. Ecosystem Energy Budgets
The extent of photosynthetic production sets the spending limit for an ecosystem’s energy budget
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18. The Global Energy Budget
The amount of solar radiation reaching Earth’s surface limits the photosynthetic output of ecosystems
Only a small fraction of solar energy actually strikes photosynthetic organisms, and even less is of a usable wavelength
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19. Gross and Net Production
Total primary production is known as the ecosystem’s gross primary production (GPP)
GPP is measured as the conversion of chemical energy from photosynthesis per unit time
Net primary production (NPP) is GPP minus energy used by primary producers for respiration
NPP is expressed as
Energy per unit area per unit time (J/m2yr), or
Biomass added per unit area per unit time (g/m2yr)
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20. NPP is the amount of new biomass added in a given time period
Only NPP is available to consumers
Standing crop is the total biomass of photosynthetic autotrophs at a given time
Ecosystems vary greatly in NPP and contribution to the total NPP on Earth
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21. TECHNIQUE 80 60 40 20 Snow Clouds Vegetation Percent reflectance Soil
Figure 55.5
TECHNIQUE 80 60 40 20
Snow
Clouds
Vegetation
Percent reflectance
Soil
Figure 55.5 Research Method: Determining Primary Production with Satellites
Liquid water
400
600
800
1,000
1,200
Visible
Near-infrared
Wavelength (nm) 21

22. Tropical rain forests, estuaries, and coral reefs are among the most productive ecosystems per unit area
Marine ecosystems are relatively unproductive per unit area but contribute much to global net primary production because of their volume
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23. Net primary production (kg carbon/m2yr)
Figure 55.6
Net primary production (kg carbon/m2yr) 3 2 1
Figure 55.6 Global net primary production. 23

24. Net ecosystem production (NEP) is a measure of the total biomass accumulation during a given period
NEP is gross primary production minus the total respiration of all organisms (producers and consumers) in an ecosystem
NEP is estimated by comparing the net flux of CO2 and O2 in an ecosystem, two molecules connected by photosynthesis
The release of O2 by a system is an indication that it is also storing CO2
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25. Float surfaces for 6–12 hours to transmit data to satellite.
Figure 55.7
Float surfaces for 6–12 hours to transmit data to satellite.
Total cycle time: 10 days
Float descends to 1,000 m and “parks.”
O2 concentration is recorded as float ascends.
Figure 55.7 Impact: Ocean Production Revealed
Drift time: 9 days 25

26. Primary Production in Aquatic Ecosystems
In marine and freshwater ecosystems, both light and nutrients control primary production
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27. Light Limitation
Depth of light penetration affects primary production in the photic zone of an ocean or lake
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28. Nutrient Limitation
More than light, nutrients limit primary production in geographic regions of the ocean and in lakes
A limiting nutrient is the element that must be added for production to increase in an area
Nitrogen and phosphorous are the nutrients that most often limit marine production
Nutrient enrichment experiments confirmed that nitrogen was limiting phytoplankton growth off the shore of Long Island, New York
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29. Phytoplankton density (millions of cells per mL)
Figure 55.8
RESULTS 30 24 18 12 6
Ammonium enriched
Phosphate enriched
Unenriched control
Phytoplankton density (millions of cells per mL)
Figure 55.8 Inquiry: Which nutrient limits phytoplankton production along the coast of Long Island? A B C D E F G
Collection site 29

30. Experiments in the Sargasso Sea in the subtropical Atlantic Ocean showed that iron limited primary production
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31. Table 55.1
Table 55.1 Nutrient Enrichment Experiment for Sargasso Sea Samples 31

32. Upwelling of nutrient-rich waters in parts of the oceans contributes to regions of high primary production
The addition of large amounts of nutrients to lakes has a wide range of ecological impacts
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33. This has led to the use of phosphate-free detergents
In some areas, sewage runoff has caused eutrophication of lakes, which can lead to loss of most fish species
In lakes, phosphorus limits cyanobacterial growth more often than nitrogen
This has led to the use of phosphate-free detergents
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34. Video: Cyanobacteria (Oscillatoria)
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35. Primary Production in Terrestrial Ecosystems
In terrestrial ecosystems, temperature and moisture affect primary production on a large scale
Primary production increases with moisture
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36. Net annual primary production (above ground, dry g/m2 yr)
Figure 55.9
1,400
1,200
1,000
800
600
400
200
Net annual primary production (above ground, dry g/m2 yr)
Figure 55.9 A global relationship between net primary production and mean annual precipitation for terrestrial ecosystems. 20 40 60 80
100
120
140
160
180
200
Mean annual precipitation (cm) 36

37. It is affected by precipitation, temperature, and solar energy
Actual evapotranspiration is the water transpired by plants and evaporated from a landscape
It is affected by precipitation, temperature, and solar energy
It is related to net primary production
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38. Nutrient Limitations and Adaptations That Reduce Them
On a more local scale, a soil nutrient is often the limiting factor in primary production
In terrestrial ecosystems, nitrogen is the most common limiting nutrient
Phosphorus can also be a limiting nutrient, especially in older soils
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39. Various adaptations help plants access limiting nutrients from soil
Some plants form mutualisms with nitrogen-fixing bacteria
Many plants form mutualisms with mycorrhizal fungi; these fungi supply plants with phosphorus and other limiting elements
Roots have root hairs that increase surface area
Many plants release enzymes that increase the availability of limiting nutrients
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40. Concept 55.3: Energy transfer between trophic levels is typically only 10% efficient
Secondary production of an ecosystem is the amount of chemical energy in food converted to new biomass during a given period of time
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41. Production Efficiency
When a caterpillar feeds on a leaf, only about one-sixth of the leaf’s energy is used for secondary production
An organism’s production efficiency is the fraction of energy stored in food that is not used for respiration
Production
efficiency 
Net secondary production  100%
Assimilation of primary production
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42. Plant material eaten by caterpillar
Figure 55.10
Plant material eaten by caterpillar
200 J
67 J
Figure Energy partitioning within a link of the food chain.
Cellular respiration
100 J
Feces
33 J
Not assimilated
Growth (new biomass; secondary production)
Assimilated 42

43. Figure 55.10a
Figure Energy partitioning within a link of the food chain. 43

44. Fishes have production efficiencies of around 10%
Birds and mammals have efficiencies in the range of 13% because of the high cost of endothermy
Fishes have production efficiencies of around 10%
Insects and microorganisms have efficiencies of 40% or more
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45. Trophic Efficiency and Ecological Pyramids
Trophic efficiency is the percentage of production transferred from one trophic level to the next
It is usually about 10%, with a range of 5% to 20%
Trophic efficiency is multiplied over the length of a food chain
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46. Approximately 0.1% of chemical energy fixed by photosynthesis reaches a tertiary consumer
A pyramid of net production represents the loss of energy with each transfer in a food chain
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47. Tertiary consumers 10 J Secondary consumers 100 J Primary consumers
Figure 55.11
Tertiary consumers
10 J
Secondary consumers
100 J
Primary consumers
1,000 J
Primary producers
Figure An idealized pyramid of net production.
10,000 J
1,000,000 J of sunlight 47

48. In a biomass pyramid, each tier represents the dry mass of all organisms in one trophic level
Most biomass pyramids show a sharp decrease at successively higher trophic levels
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49. Dry mass (g/m2) Dry mass (g/m2)
Figure 55.12
Trophic level
Dry mass (g/m2)
Tertiary consumers
Secondary consumers
Primary consumers
Primary producers
1.5 11 37
809
(a) Most ecosystems (data from a Florida bog)
Figure Pyramids of biomass (standing crop).
Trophic level
Dry mass (g/m2)
Primary consumers (zooplankton)
Primary producers (phytoplankton) 21 4
(b) Some aquatic ecosystems (data from the English Channel) 49

50. Turnover time is the ratio of the standing crop biomass to production
Certain aquatic ecosystems have inverted biomass pyramids: producers (phytoplankton) are consumed so quickly that they are outweighed by primary consumers
Turnover time is the ratio of the standing crop biomass to production
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51. Dynamics of energy flow in ecosystems have important implications for the human population
Eating meat is a relatively inefficient way of tapping photosynthetic production
Worldwide agriculture could feed many more people if humans ate only plant material
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52. Concept 55.4: Biological and geochemical processes cycle nutrients and water in ecosystems
Life depends on recycling chemical elements
Nutrient cycles in ecosystems involve biotic and abiotic components and are often called biogeochemical cycles
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53. Biogeochemical Cycles
Gaseous carbon, oxygen, sulfur, and nitrogen occur in the atmosphere and cycle globally
Less mobile elements include phosphorus, potassium, and calcium
These elements cycle locally in terrestrial systems but more broadly when dissolved in aquatic systems
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54. All elements cycle between organic and inorganic reservoirs
A model of nutrient cycling includes main reservoirs of elements and processes that transfer elements between reservoirs
All elements cycle between organic and inorganic reservoirs
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55. Burning of fossil fuels Formation of sedimentary rock
Figure 55.13
Reservoir A
Organic materials available as nutrients
Reservoir B Organic materials unavailable as nutrients
Fossilization
Living organisms, detritus
Peat
Coal
Oil
Respiration, decomposition, excretion
Burning of fossil fuels
Assimilation, photosynthesis
Reservoir D Inorganic materials unavailable as nutrients
Reservoir C Inorganic materials available as nutrients
Figure A general model of nutrient cycling.
Weathering, erosion
Atmosphere
Minerals in rocks
Water
Formation of sedimentary rock
Soil 55

56. In studying cycling of water, carbon, nitrogen, and phosphorus, ecologists focus on four factors
Each chemical’s biological importance
Forms in which each chemical is available or used by organisms
Major reservoirs for each chemical
Key processes driving movement of each chemical through its cycle
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57. The Water Cycle Water is essential to all organisms
Liquid water is the primary physical phase in which water is used
The oceans contain 97% of the biosphere’s water; 2% is in glaciers and polar ice caps, and 1% is in lakes, rivers, and groundwater
Water moves by the processes of evaporation, transpiration, condensation, precipitation, and movement through surface and groundwater
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58. Movement over land by wind
Figure 55.14a
Movement over land by wind
Evaporation from ocean
Precipitation over land
Precipitation over ocean
Evapotranspira- tion from land
Figure Exploring: Water and Nutrient Cycling
Percolation through soil
Runoff and groundwater 58

59. The Carbon Cycle
Carbon-based organic molecules are essential to all organisms
Photosynthetic organisms convert CO2 to organic molecules that are used by heterotrophs
Carbon reservoirs include fossil fuels, soils and sediments, solutes in oceans, plant and animal biomass, the atmosphere, and sedimentary rocks
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60. CO2 is taken up and released through photosynthesis and respiration; additionally, volcanoes and the burning of fossil fuels contribute CO2 to the atmosphere
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61. Burning of fossil fuels and wood
Figure 55.14b
CO2 in atmosphere
Photosynthesis
Photo- synthesis
Cellular respiration
Burning of fossil fuels and wood
Phyto- plankton
Consumers
Figure Exploring: Water and Nutrient Cycling
Consumers
Decomposition 61

62. The Nitrogen Cycle
Nitrogen is a component of amino acids, proteins, and nucleic acids
The main reservoir of nitrogen is the atmosphere (N2), though this nitrogen must be converted to NH4+ or NO3– for uptake by plants, via nitrogen fixation by bacteria
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63. Denitrification converts NO3– back to N2
Organic nitrogen is decomposed to NH4+ by ammonification, and NH4+ is decomposed to NO3– by nitrification
Denitrification converts NO3– back to N2
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64. Decomposition and sedimentation
Figure 55.14c
N2 in atmosphere
Reactive N gases
Industrial fixation
Denitrification
N fertilizers
Fixation
Runoff
Dissolved organic N
NO3–
Terrestrial cycling N2
NH4+
NO3–
Aquatic cycling
Denitri- fication
Figure Exploring: Water and Nutrient Cycling
Decomposition and sedimentation
Assimilation
Decom- position
NO3–
Fixation in root nodules
Uptake of amino acids
Ammonification
Nitrification
NH3
NH4+
NO2– 64

65. Decomposition and sedimentation
Figure 55.14ca
N2 in atmosphere
Reactive N gases
Industrial fixation
Denitrification
N fertilizers
Fixation
Runoff
Dissolved organic N
NO3–
NH4+
NO3–
Terrestrial cycling
Figure Exploring: Water and Nutrient Cycling
Aquatic cycling
Decomposition and sedimentation 65

66. Fixation in root nodules
Figure 55.14cb
Terrestrial cycling N2
Denitri- fication
Assimilation
Decom- position
NO3–
Uptake of amino acids
Fixation in root nodules
Figure Exploring: Water and Nutrient Cycling
Ammonification
Nitrification
NH3
NH4+
NO2– 66

67. The Phosphorus Cycle
Phosphorus is a major constituent of nucleic acids, phospholipids, and ATP
Phosphate (PO43–) is the most important inorganic form of phosphorus
The largest reservoirs are sedimentary rocks of marine origin, the oceans, and organisms
Phosphate binds with soil particles, and movement is often localized
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68. Wind-blown dust Geologic uplift Weathering of rocks Runoff Consumption
Figure 55.14d
Wind-blown dust
Geologic uplift
Weathering of rocks
Runoff
Consumption
Decomposition
Plant uptake of PO43–
Plankton
Dissolved PO43–
Uptake
Leaching
Figure Exploring: Water and Nutrient Cycling
Sedimentation
Decomposition 68

69. Decomposition and Nutrient Cycling Rates
Decomposers (detritivores) play a key role in the general pattern of chemical cycling
Rates at which nutrients cycle in different ecosystems vary greatly, mostly as a result of differing rates of decomposition
The rate of decomposition is controlled by temperature, moisture, and nutrient availability
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70. Mean annual temperature (C)
Figure 55.15
EXPERIMENT
Ecosystem type
Arctic
Subarctic
Boreal
Temperate
Grassland
Mountain A G M
B,C D P T
H,I S
E,F U N L O J K Q R
RESULTS 80 70 60 50 40 30 20 10
Figure Inquiry: How does temperature affect litter decomposition in an ecosystem? R U
Percent of mass lost K O Q T J P D S N F C I M L A H B E G
–15
–10 –5 5 10 15
Mean annual temperature (C) 70

71. EXPERIMENT Ecosystem type Arctic Subarctic Boreal Temperate Grassland
Figure 55.15a
EXPERIMENT
Ecosystem type
Arctic
Subarctic
Boreal
Temperate
Grassland
Mountain A G M D
B,C P T
H,I
Figure Inquiry: How does temperature affect litter decomposition in an ecosystem?
E,F S O U N L J K Q R 71

72. Mean annual temperature (C)
Figure 55.15b
RESULTS 80 70 60 50 40 30 20 10 U R O Q K T
Percent of mass lost J P S D N F C I M L H
Figure Inquiry: How does temperature affect litter decomposition in an ecosystem? A B E G
–15
–10 –5 5 10 15
Mean annual temperature (C) 72

73. Decomposition is slow in anaerobic muds
Rapid decomposition results in relatively low levels of nutrients in the soil
For example, in a tropical rain forest, material decomposes rapidly, and most nutrients are tied up in trees other living organisms
Cold and wet ecosystems store large amounts of undecomposed organic matter as decomposition rates are low
Decomposition is slow in anaerobic muds
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74. Case Study: Nutrient Cycling in the Hubbard Brook Experimental Forest
The Hubbard Brook Experimental Forest has been used to study nutrient cycling in a forest ecosystem since 1963
The research team constructed a dam on the site to monitor loss of water and minerals
They found that 60% of the precipitation exits through streams and 40% is lost by evapotranspiration
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75. Nitrate concentration in runoff (mg/L) Completion of tree cutting
Figure 55.16
(a) Concrete dam and weir
(b) Clear-cut watershed 80 60 40 20
Deforested
Figure Nutrient cycling in the Hubbard Brook Experimental Forest: an example of long-term ecological research.
Nitrate concentration in runoff (mg/L)
Completion of tree cutting 4 3 2 1
Control
1965
1966
1967
1968
(c) Nitrate in runoff from watersheds 75

76. (a) Concrete dam and weir
Figure 55.16a
Figure Nutrient cycling in the Hubbard Brook Experimental Forest: an example of long-term ecological research.
(a) Concrete dam and weir 76

77. In one experiment, the trees in one valley were cut down, and the valley was sprayed with herbicides
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78. (b) Clear-cut watershed
Figure 55.16b
Figure Nutrient cycling in the Hubbard Brook Experimental Forest: an example of long-term ecological research.
(b) Clear-cut watershed 78

79. These results showed how human activity can affect ecosystems
Net losses of water were 3040% greater in the deforested site than the undisturbed (control) site
Nutrient loss was also much greater in the deforested site compared with the undisturbed site
For example, nitrate levels increased 60 times in the outflow of the deforested site
These results showed how human activity can affect ecosystems
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80. Nitrate concentration in runoff (mg/L) Completion of tree cutting
Figure 55.16c 80 60 40 20
Deforested
Nitrate concentration in runoff (mg/L)
Completion of tree cutting 4 3 2 1
Control
Figure Nutrient cycling in the Hubbard Brook Experimental Forest: an example of long-term ecological research.
1965
1966
1967
1968
(c) Nitrate in runoff from watersheds 80

81. Concept 55.5: Restoration ecologists help return degraded ecosystems to a more natural state
Given enough time, biological communities can recover from many types of disturbances
Restoration ecology seeks to initiate or speed up the recovery of degraded ecosystems
Two key strategies are bioremediation and augmentation of ecosystem processes
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82. (a) In 1991, before restoration
Figure 55.17
Figure A gravel and clay mine site in New Jersey before and after restoration.
(a) In 1991, before restoration
(b) In 2000, near the completion of restoration 82

83. (a) In 1991, before restoration
Figure 55.17a
Figure A gravel and clay mine site in New Jersey before and after restoration.
(a) In 1991, before restoration 83

84. (b) In 2000, near the completion of restoration
Figure 55.17b
Figure A gravel and clay mine site in New Jersey before and after restoration.
(b) In 2000, near the completion of restoration 84

85. Bioremediation
Bioremediation is the use of organisms to detoxify ecosystems
The organisms most often used are prokaryotes, fungi, or plants
These organisms can take up, and sometimes metabolize, toxic molecules
For example, the bacterium Shewanella oneidensis can metabolize uranium and other elements to insoluble forms that are less likely to leach into streams and groundwater
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86. Concentration of soluble uranium (M)
Figure 55.18 6 5 4 3 2 1
Concentration of soluble uranium (M)
Figure Bioremediation of groundwater contaminated with uranium at Oak Ridge National Laboratory, Tennessee. 50
100
150
200
250
300
350
400
Days after adding ethanol 86

87. Figure 55.18a
Figure Bioremediation of groundwater contaminated with uranium at Oak Ridge National Laboratory, Tennessee. 87

88. Biological Augmentation
Biological augmentation uses organisms to add essential materials to a degraded ecosystem
For example, nitrogen-fixing plants can increase the available nitrogen in soil
For example, adding mycorrhizal fungi can help plants to access nutrients from soil
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89. Restoration Projects Worldwide
The newness and complexity of restoration ecology require that ecologists consider alternative solutions and adjust approaches based on experience
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90. Figure 55.19a
Equator
Figure Exploring: Restoration Ecology Worldwide 90

91. Kissimmee River, Florida
Figure 55.19b
Figure Exploring: Restoration Ecology Worldwide
Kissimmee River, Florida 91

92. Truckee River, Nevada Figure 55.19c
Figure Exploring: Restoration Ecology Worldwide
Truckee River, Nevada 92

93. Tropical dry forest, Costa Rica
Figure 55.19d
Figure Exploring: Restoration Ecology Worldwide
Tropical dry forest, Costa Rica 93

94. Rhine River, Europe Figure 55.19e
Figure Exploring: Restoration Ecology Worldwide
Rhine River, Europe 94

95. Succulent Karoo, South Africa
Figure 55.19f
Figure Exploring: Restoration Ecology Worldwide
Succulent Karoo, South Africa 95

96. Figure 55.19g
Figure Exploring: Restoration Ecology Worldwide
Coastal Japan 96

97. Maungatautari, New Zealand
Figure 55.19h
Figure Exploring: Restoration Ecology Worldwide
Maungatautari, New Zealand 97

98. Microorganisms and other detritivores Secondary and tertiary consumers
Figure 55.UN01
Sun
Key
Chemical cycling
Energy flow
Heat
Primary producers
Primary consumers
Detritus
Figure 55.UN01 Summary figure, Concept 55.1
Microorganisms and other detritivores
Secondary and tertiary consumers 98

99. Formation of sedimentary rock
Figure 55.UN02
Reservoir A
Organic materials available as nutrients:
Living organisms, detritus
Reservoir B
Organic materials unavailable as nutrients:
Peat, coal, oil
Fossilization
Respiration, decomposition, excretion
Burning of fossil fuels
Assimilation, photosynthesis
Figure 55.UN02 Summary figure, Concept 55.4
Reservoir D
Inorganic materials unavailable as nutrients:
Minerals in rocks
Reservoir C
Inorganic materials available as nutrients:
Atmosphere, water, soil
Weathering, erosion
Formation of sedimentary rock 99

100. Figure 55.UN03
Figure 55.UN03 Appendix A: answer to Concept Check 55.4, question 1
100

101. Figure 55.UN04
Figure 55.UN04 Appendix A: Answer to Test Your Understanding, question 9
101