Chapter 54

Ecosystems

PowerPoint Lectures for Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero
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• Overview: Ecosystems, Energy, and Matter • An ecosystem consists of all the organisms living in a community
– As well as all the abiotic factors with which they interact

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• Ecosystems can range from a microcosm, such as an aquarium
– To a large area such as a lake or forest

Figure 54.1
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• Regardless of an ecosystem’s size
– Its dynamics involve two main processes: energy flow and chemical cycling

• Energy flows through ecosystems
– While matter cycles within them

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• Concept 54.1: Ecosystem ecology emphasizes energy flow and chemical cycling • Ecosystem ecologists view ecosystems
– As transformers of energy and processors of matter

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Ecosystems and Physical Laws • The laws of physics and chemistry apply to ecosystems
– Particularly in regard to the flow of energy

• Energy is conserved
– But degraded to heat during ecosystem processes

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Trophic Relationships • Energy and nutrients pass from primary producers (autotrophs)
– To primary consumers (herbivores) and then to secondary consumers (carnivores)

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• Energy flows through an ecosystem
– Entering as light and exiting as heat
Tertiary consumers Microorganisms and other detritivores

Secondary consumers

Detritus

Primary consumers

Primary producers Heat Sun

Key Chemical cycling Energy flow

Figure 54.2
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• Nutrients cycle within an ecosystem

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Decomposition • Decomposition
– Connects all trophic levels

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• Detritivores, mainly bacteria and fungi, recycle essential chemical elements
– By decomposing organic material and returning elements to inorganic reservoirs

Figure 54.3
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• Concept 54.2: Physical and chemical factors limit primary production in ecosystems • Primary production in an ecosystem
– Is the amount of light energy converted to chemical energy by autotrophs during a given time period

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Ecosystem Energy Budgets • The extent of photosynthetic production
– Sets the spending limit for the energy budget of the entire ecosystem

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The Global Energy Budget • The amount of solar radiation reaching the surface of the Earth
– Limits the photosynthetic output of ecosystems

• Only a small fraction of solar energy
– Actually strikes photosynthetic organisms

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Gross and Net Primary Production • Total primary production in an ecosystem
– Is known as that ecosystem’s gross primary production (GPP)

• Not all of this production
– Is stored as organic material in the growing plants

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• Net primary production (NPP)
– Is equal to GPP minus the energy used by the primary producers for respiration

• Only NPP
– Is available to consumers

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• Different ecosystems vary considerably in their net primary production
– And in their contribution to the total NPP on Earth
Open ocean Continental shelf Estuary 65.0 Algal beds and reefs Upwelling zones Extreme desert, rock, sand, ice Desert and semidesert scrub Tropical rain forest Savanna Cultivated land Boreal forest (taiga) Temperate grassland Woodland and shrubland Tundra Tropical seasonal forest Temperate deciduous forest Temperate evergreen forest Swamp and marsh Lake and stream 0 Key Marine Terrestrial Freshwater (on continents) 5.2 0.3 0.1 0.1 4.7 3.5 3.3 2.9 2.7 2.4 1.8 1.7 1.6 1.5 1.3 1.0 0.4 0.4 10 20 30 40 50 60 0 250 500 1,000 1,500 2,000 2,500 (b) Average net primary production (g/m2/yr) 0 140 1,600 1,200 1,300 2,000 125 360 24.4 5.6 1,500 2,500 500 3.0 90 2,200 900 600 800 600 700 0.6 7.1 4.9 3.8 2.3 0.3 5 10 15 20 25 7.9 9.1 9.6 5.4 3.5 1.2 0.9 0.1 0.04 0.9 22

(a) Percentage of Earth’s surface area

(c) Percentage of Earth’s net primary production

Figure 54.4a–c

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• Overall, terrestrial ecosystems
– Contribute about two-thirds of global NPP and marine ecosystems about one-third
North Pole 60°N 30°N Equator 30°S 60°S South Pole 180° Figure 54.5
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120°W

60°W

60°E

120°E

180°

Primary Production in Marine and Freshwater Ecosystems • In marine and freshwater ecosystems
– Both light and nutrients are important in controlling primary production

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Light Limitation • The depth of light penetration
– Affects primary production throughout the photic zone of an ocean or lake

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Nutrient Limitation • More than light, nutrients limit primary production
– Both in different geographic regions of the ocean and in lakes

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• A limiting nutrient is the element that must be added
– In order for production to increase in a particular area

• Nitrogen and phosphorous
– Are typically the nutrients that most often limit marine production

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• Nutrient enrichment experiments
– Confirmed that nitrogen was limiting phytoplankton growth in an area of the ocean
Pollution from duck farms concentrated near Moriches Bay adds both nitrogen and phosphorus to the coastal water off Long Island. Researchers cultured the phytoplankton Nannochloris atomus with water collected from several bays.
EXPERIMENT

30
I ng Lo nd sla

21 15 19 Shinnecock Bay

Coast of Long Island, New York. The numbers on the map indicate the data collection stations.

B outh at S Gre

4

5

ay

11

Moriches Bay Atlantic Ocean

2

Figure 54.6
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RESULTS Phytoplankton abundance parallels the abundance of phosphorus in the water (a). Nitrogen, however, is immediately taken up by algae, and no free nitrogen is measured in the coastal waters. The addition of ammonium (NH4+) caused heavy phytoplankton growth in bay water, but the addition of phosphate (PO43+) did not induce algal growth (b). 8 7 6 5 4 3 2 1 0 Inorganic phosphorus (µg atoms/L) Phytoplankton (millions of cells/mL) Phytoplankton Inorganic phosphorus 8 7 6 5 4 3 2 1 0 30 Phytoplankton (millions of cells per mL) 24 18 12 6 0 Shinnecock Bay Ammonium enriched Phosphate enriched Unenriched control

2

11 30 15 19 21 Station number Great Moriches South Bay Bay

4

5

Starting 2 algal density

4

5 11 30 Station number

15

19

21

(a) Phytoplankton biomass and phosphorus concentration

(b) Phytoplankton response to nutrient enrichment

CONCLUSION Since adding phosphorus, which was already in rich supply, had no effect on Nannochloris growth, whereas adding nitrogen increased algal density dramatically, researchers concluded that nitrogen was the nutrient limiting phytoplankton growth in this ecosystem.

Figure 54.6
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• Experiments in another ocean region
– Showed that iron limited primary production

Table 54.1
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• The addition of large amounts of nutrients to lakes
– Has a wide range of ecological impacts

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• In some areas, sewage runoff
– Has caused eutrophication of lakes, which can lead to the eventual loss of most fish species from the lakes

Figure 54.7
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Primary Production in Terrestrial and Wetland Ecosystems • In terrestrial and wetland ecosystems climatic factors
– Such as temperature and moisture, affect primary production on a large geographic scale

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• The contrast between wet and dry climates
– Can be represented by a measure called actual evapotranspiration

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• Actual evapotranspiration
– Is the amount of water annually transpired by plants and evaporated from a landscape – Is related to net primary production
3,000 Net primary production (g/m2/yr) Tropical forest

2,000

Temperate forest 1,000 Desert shrubland 0 0 Arctic tundra 500 1,000 1,500 Mountain coniferous forest Temperate grassland

Figure 54.8

Actual evapotranspiration (mm H2O/yr)

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• On a more local scale
– A soil nutrient is often the limiting factor in primary production
EXPERIMENT Over the summer of 1980, researchers added phosphorus to some experimental plots in the salt marsh, nitrogen to other plots, and both phosphorus and nitrogen to others. Some plots were left unfertilized as controls. Adding nitrogen (N) boosts net primary RESULTS production. Live, above-ground biomass (g dry wt/m2) 300 250 200 150 100 50 0 July N only N+P

Control P only

June

August 1980

Experimental plots receiving just phosphorus (P) do not outproduce the unfertilized control plots. CONCLUSION These nutrient enrichment experiments confirmed that nitrogen was the nutrient limiting plant growth in this salt marsh.

Figure 54.9

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• Concept 54.3: Energy transfer between trophic levels is usually less than 20% efficient • The secondary production of an ecosystem
– Is the amount of chemical energy in consumers’ food that is converted to their own new biomass during a given period of time

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Production Efficiency • When a caterpillar feeds on a plant leaf
– Only about one-sixth of the energy in the leaf is used for secondary production

Plant material eaten by caterpillar

200 J

67 J Feces 100 J 33 J Growth (new biomass)

Cellular respiration

Figure 54.10

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• The production efficiency of an organism
– Is the fraction of energy stored in food that is not used for respiration

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Trophic Efficiency and Ecological Pyramids • Trophic efficiency
– Is the percentage of production transferred from one trophic level to the next – Usually ranges from 5% to 20%

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Pyramids of Production
• This loss of energy with each transfer in a food chain
– Can be represented by a pyramid of net production
Tertiary consumers

10 J

Secondary consumers

100 J

Primary consumers

1,000 J

Primary producers

10,000 J

Figure 54.11
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1,000,000 J of sunlight

Pyramids of Biomass • One important ecological consequence of low trophic efficiencies
– Can be represented in a biomass pyramid

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• Most biomass pyramids
– Show a sharp decrease at successively higher trophic levels
Trophic level Tertiary consumers Secondary consumers Primary consumers Primary producers (a) Most biomass pyramids show a sharp decrease in biomass at successively higher trophic levels, as illustrated by data from a bog at Silver Springs, Florida. Dry weight (g/m2) 1.5 11 37 809

Figure 54.12a
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• Certain aquatic ecosystems
– Have inverted biomass pyramids
Trophic level Dry weight (g/m2) 21 4

Primary consumers (zooplankton) Primary producers (phytoplankton)

(b) In some aquatic ecosystems, such as the English Channel, a small standing crop of primary producers (phytoplankton) supports a larger standing crop of primary consumers (zooplankton).

Figire 54.12b

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Pyramids of Numbers • A pyramid of numbers
– Represents the number of individual organisms in each trophic level

Trophic level

Number of individual organisms 3 354,904 708,624 5,842,424

Tertiary consumers Secondary consumers Primary consumers Primary producers

Figure 54.13
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• The dynamics of energy flow through ecosystems
– Have important implications for the human population

• Eating meat
– Is a relatively inefficient way of tapping photosynthetic production

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• Worldwide agriculture could successfully feed many more people
– If humans all fed more efficiently, eating only plant material
Trophic level Secondary consumers

Primary consumers

Primary producers Figure 54.14
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The Green World Hypothesis • According to the green world hypothesis
– Terrestrial herbivores consume relatively little plant biomass because they are held in check by a variety of factors

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• Most terrestrial ecosystems
– Have large standing crops despite the large numbers of herbivores

Figure 54.15
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• The green world hypothesis proposes several factors that keep herbivores in check
– Plants have defenses against herbivores – Nutrients, not energy supply, usually limit herbivores – Abiotic factors limit herbivores – Intraspecific competition can limit herbivore numbers – Interspecific interactions check herbivore densities
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• Concept 54.4: Biological and geochemical processes move nutrients between organic and inorganic parts of the ecosystem • Life on Earth
– Depends on the recycling of essential chemical elements

• Nutrient circuits that cycle matter through an ecosystem
– Involve both biotic and abiotic components and are often called biogeochemical cycles
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A General Model of Chemical Cycling • Gaseous forms of carbon, oxygen, sulfur, and nitrogen
– Occur in the atmosphere and cycle globally

• Less mobile elements, including phosphorous, potassium, and calcium
– Cycle on a more local level

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• A general model of nutrient cycling
– Includes the main reservoirs of elements and the processes that transfer elements between reservoirs
Reservoir a Organic materials available as nutrients Living organisms, detritus Reservoir b Organic materials unavailable as nutrients Coal, oil, peat Fossilization

Assimilation, photosynthesis

Respiration, decomposition, excretion

Burning of fossil fuels Reservoir d Inorganic materials unavailable as nutrients Minerals in rocks

Reservoir c Inorganic materials available as nutrients

Weathering, erosion Formation of sedimentary rock

Figure 54.16

Atmosphere, soil, water

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• All elements
– Cycle between organic and inorganic reservoirs

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Biogeochemical Cycles • The water cycle and the carbon cycle
THE WATER CYCLE THE CARBON CYCLE

Transport over land Solar energy Net movement of water vapor by wind Precipitation over ocean Evaporation from ocean Evapotranspiration from land Precipitation over land Burning of fossil fuels and wood

CO2 in atmosphere Photosynthesis Cellular respiration

Higher-level Primary consumers consumers Detritus

Percolation through soil Runoff and groundwater

Carbon compounds in water

Decomposition

Figure 54.17
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• Water moves in a global cycle
– Driven by solar energy

• The carbon cycle
– Reflects the reciprocal processes of photosynthesis and cellular respiration

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• The nitrogen cycle and the phosphorous cycle
THE NITROGEN CYCLE THE PHOSPHORUS CYCLE

N2 in atmosphere Rain

Geologic uplift Assimilation NO3− Decomposers Nitrification NH4+ Nitrifying bacteria NO2 − Denitrifying bacteria

Weathering of rocks Runoff

Plants

Consumption Sedimentation Soil Leaching Plant uptake of PO43−

Nitrogen-fixing bacteria in root nodules of legumes

Ammonification NH3 Nitrogen-fixing soil bacteria

Nitrifying bacteria

Decomposition

Figure 54.17
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• Most of the nitrogen cycling in natural ecosystems
– Involves local cycles between organisms and soil or water

• The phosphorus cycle
– Is relatively localized

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Decomposition and Nutrient Cycling Rates • Decomposers (detritivores) play a key role
– In the general pattern of chemical cycling
Consumers

Producers Decomposers Nutrients available to producers

Abiotic reservoir

Figure 54.18
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Geologic processes

• The rates at which nutrients cycle in different ecosystems
– Are extremely variable, mostly as a result of differences in rates of decomposition

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Vegetation and Nutrient Cycling: The Hubbard Brook Experimental Forest • Nutrient cycling
– Is strongly regulated by vegetation

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• Long-term ecological research projects
– Monitor ecosystem dynamics over relatively long periods of time

• The Hubbard Brook Experimental Forest
– Has been used to study nutrient cycling in a forest ecosystem since 1963

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• The research team constructed a dam on the site
– To monitor water and mineral loss

Figure 54.19a

(a) Concrete dams and weirs built across streams at the bottom of watersheds enabled researchers to monitor the outflow of water and nutrients from the ecosystem.

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• In one experiment, the trees in one valley were cut down
– And the valley was sprayed with herbicides

Figure 54.19b

(b) One watershed was clear cut to study the effects of the loss of vegetation on drainage and nutrient cycling.

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• Net losses of water and minerals were studied
– And found to be greater than in an undisturbed area

• These results showed how human activity
– Can affect ecosystems
Nitrate concentration in runoff (mg/L) 80.0 60.0 40.0 20.0 4.0 3.0 2.0 1.0 0 Completion of tree cutting Deforested

Control

1965

1966

1967

1968

Figure 54.19c

(c) The concentration of nitrate in runoff from the deforested watershed was 60 times greater than in a control (unlogged) watershed.

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• Concept 54.5: The human population is disrupting chemical cycles throughout the biosphere • As the human population has grown in size
– Our activities have disrupted the trophic structure, energy flow, and chemical cycling of ecosystems in most parts of the world

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Nutrient Enrichment • In addition to transporting nutrients from one location to another
– Humans have added entirely new materials, some of them toxins, to ecosystems

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Agriculture and Nitrogen Cycling
• Agriculture constantly removes nutrients from ecosystems
– That would ordinarily be cycled back into the soil

Figure 54.20
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• Nitrogen is the main nutrient lost through agriculture
– Thus, agriculture has a great impact on the nitrogen cycle

• Industrially produced fertilizer is typically used to replace lost nitrogen
– But the effects on an ecosystem can be harmful

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Contamination of Aquatic Ecosystems • The critical load for a nutrient
– Is the amount of that nutrient that can be absorbed by plants in an ecosystem without damaging it

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• When excess nutrients are added to an ecosystem, the critical load is exceeded
– And the remaining nutrients can contaminate groundwater and freshwater and marine ecosystems

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• Sewage runoff contaminates freshwater ecosystems
– Causing cultural eutrophication, excessive algal growth, which can cause significant harm to these ecosystems

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Acid Precipitation • Combustion of fossil fuels
– Is the main cause of acid precipitation

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• North American and European ecosystems downwind from industrial regions
– Have been damaged by rain and snow containing nitric and sulfuric acid

4.6

4.3

4.6 4.3 4.6 4.3 4.6 Europe 4.1

Figure 54.21

North America

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• By the year 2000
– The entire contiguous United States was affected by acid precipitation

Figure 54.22
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Field pH ≥5.3 5.2–5.3 5.1–5.2 5.0–5.1 4.9–5.0 4.8–4.9 4.7–4.8 4.6–4.7 4.5–4.6 4.4–4.5 4.3–4.4 <4.3

• Environmental regulations and new industrial technologies
– Have allowed many developed countries to reduce sulfur dioxide emissions in the past 30 years

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Toxins in the Environment • Humans release an immense variety of toxic chemicals
– Including thousands of synthetics previously unknown to nature

• One of the reasons such toxins are so harmful
– Is that they become more concentrated in successive trophic levels of a food web

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• In biological magnification
– Toxins concentrate at higher trophic levels because at these levels biomass tends to be lower
Herring gull eggs 124 ppm Concentration of PCBs

Lake trout 4.83 ppm

Smelt 1.04 ppm

Figure 54.23

Zooplankton 0.123 ppm

Phytoplankton 0.025 ppm

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• In some cases, harmful substances
– Persist for long periods of time in an ecosystem and continue to cause harm

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Atmospheric Carbon Dioxide • One pressing problem caused by human activities
– Is the rising level of atmospheric carbon dioxide

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Rising Atmospheric CO2
• Due to the increased burning of fossil fuels and other human activities
– The concentration of atmospheric CO2 has been steadily increasing
390 380 370 360 350 0.30 340 330 320 310 300 CO2 0.15 0 1.05 0.90 0.75 Temperature 0.60 0.45 Temperature variation (°C)

CO2 concentration (ppm)

−0.15 − 0.30 − 0.45 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 Year

Figure 54.24

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How Elevated CO2 Affects Forest Ecology: The FACTS-I Experiment
• The FACTS-I experiment is testing how elevated CO2
– Influences tree growth, carbon concentration in soils, and other factors over a ten-year period

Figure 54.25
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The Greenhouse Effect and Global Warming • The greenhouse effect is caused by atmospheric CO2
– But is necessary to keep the surface of the Earth at a habitable temperature

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• Increased levels of atmospheric CO2 are magnifying the greenhouse effect
– Which could cause global warming and significant climatic change

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Depletion of Atmospheric Ozone • Life on Earth is protected from the damaging effects of UV radiation
– By a protective layer or ozone molecules present in the atmosphere

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• Satellite studies of the atmosphere
– Suggest that the ozone layer has been gradually thinning since 1975
350 Ozone layer thickness (Dobson units) 300

250

200

150

100

50

0 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005

Figure 54.26

Year (Average for the month of October)

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• The destruction of atmospheric ozone
– Probably results from chlorine-releasing pollutants produced by human activity
1 Chlorine from CFCs interacts with ozone (O3), forming chlorine monoxide (ClO) and oxygen (O2). Chlorine atoms O2 Chlorine O3

ClO O2 3 Sunlight causes Cl2O2 to break down into O2 and free chlorine atoms. The chlorine atoms can begin the cycle again.

ClO Cl2O2 2 Two ClO molecules react, forming chlorine peroxide (Cl2O2).

Figure 54.27

Sunlight

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• Scientists first described an “ozone hole”
– Over Antarctica in 1985; it has increased in size as ozone depletion has increased

(a) October 1979 Figure 54.28a, b
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(b) October 2000