13.3 Energy Flow in Ecosystems E X P E C TAT I O N S Explain how autotrophs sustain ecosystems by supporting higher trophic levels. Describe the concept of primary productivity and explain why it varies among ecosystems. Use the energy pyramid to explain production, distribution, and use of food resources. Using the ecological hierarchy of living things, evaluate how a change in one population can affect the entire hierarchy. Relate the pyramid of primary productivity to the pyramids of biomass and numbers. All organisms require energy for growth, body maintenance (such as repairing damage to body parts), and reproduction, and many species require energy for locomotion. Energy to support these activities is released from large, energy-rich organic molecules during the process of cellular respiration (or, in a few species, fermentation) and stored in the form of ATP. Most primary producers use energy from the Sun to drive the production of energy-rich molecules in their own bodies. Consumers subsequently obtain these molecules by eating either the bodies of primary producers or the bodies of other consumers. Therefore, the amount of energy available to an ecosystem is determined by the amount of energy captured by the autotrophs. Every day the Sun bombards Earth with about 1022 joules of solar radiation. The majority of this radiation is absorbed, scattered, or reflected by the atmosphere. Much of the radiation that does reach the biosphere hits bare ground or water, with only a small fraction landing on photosynthetic organisms. Of this, only a portion is of a wavelength suitable for photosynthesis. The result is that only one to two percent of the total radiation emitted by the Sun is converted into chemical energy. The amount varies somewhat from place to place, depending on the type of organisms found in each region, the intensity of the light (recall Figure 13.9 on page 434), and many other factors (see Figure 13.17). Even though the fraction of solar radiation actually captured by primary producers is very small, these organisms produce between 150 and 200 billion tonnes of organic material each year. This amount supports the majority of life on Earth. 444 MHR • Unit 5 Population Dynamics 100% of solar radiation reaching Earth reflected back by clouds, dust, and Earth’s surface 34% 23% drives the water cycle 42% absorbed by the atmosphere and the surface of Earth 1% drives air and water currents 1 to 2% captured by photosynthesis Figure 13.17 Very little of the total amount of radiation leaving the Sun is converted to chemical energy during photosynthesis. Primary Productivity of rainfall the system receives (since water is a raw material of photosynthesis). In addition, photosynthetic organisms need certain nutrients, such as nitrogen, potassium, and phosphorus, to grow. Even though many of these nutrients are not directly involved in photosynthesis, they contribute to limiting the rate of primary productivity by affecting plant growth. Primary productivity is the amount of light energy that autotrophs in an ecosystem convert to chemical energy (and store in organic compounds) during a specific period of time. It is commonly measured in terms of energy per area, per year (J/m2/a). It can also be expressed as biomass (mass) of vegetation added to an ecosystem per area, per year (g/m2/a). It is important to remember that primary productivity is the rate at which organisms produce new biomass, which is not the same as the total mass of all photosynthetic autotrophs present in an area at one time. For example, a forest has a very large biomass — the mass of its vegetation is greater than that of a grassland of equal size. But primary productivity of the grassland may actually be higher because animals are constantly eating the plants and new ones are being produced. Thus, new mass is being accumulated in the grassland at a higher rate than in the forest. The amount of primary productivity (as shown in Figure 13.18) can vary significantly, both among ecosystems and within an ecosystem over time. The rate of productivity depends on many factors, including the number of autotrophs present in the ecosystem, the amount of light and heat present (the process slows during winter), and the amount Not all solar energy captured by primary producers is passed on to higher trophic levels. A substantial portion of the energy captured by producers is used in their own cellular respiration reactions. Another large portion is simply never eaten by consumers — which is why many ecosystems look green. Only some of the total biomass eaten by consumers is converted into the body tissues of the organism that ate it. Figure 13.19 on the following page shows what happens to the energy a herbivore (caterpillar) obtains from the plant material it eats. Approximately half the plant tissue is indigestible, and the energy from this portion is expelled with the caterpillar’s feces. Although some of this energy will be consumed by decomposers and will continue to be part of the ecosystem, it will not 3 4 5 6 A Percentage (%) of Earth’s surface area Figure 13.18 Why do the oceans contribute such a high proportion of Earth’s total primary productivity when their average productivity is low compared with that of algal beds and reefs? Net primary productivity is the total amount of 0 B Average net primary productivity (g/m2 /a) 2500 2 2000 1 1500 0 1000 65.0 5.2 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 0.3 0.1 0.1 500 open ocean continental shelf extreme desert, rock, sand, or ice desert and semi-desert 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 land and stream estuary algal beds and reefs upwelling zones Energy Transfer at Higher Trophic Levels 0 5 10 15 20 25 C Percentage (%) of Earth’s net primary productivity solar energy transformed to chemical energy by autotrophs (called gross primary productivity), minus the amount used by the autotrophs during cellular respiration. Chapter 13 Ecological Principles • MHR 445 be available to the secondary consumers that eat caterpillars. Approximately one third of the energy the caterpillar obtains is used in its own cellular respiration (providing energy for locomotion, maintaining body temperature, and other body processes), and therefore lost to the ecosystem. In fact, only about one sixth of the energy is incorporated into new caterpillar tissue — tissue that can be eaten by secondary consumers. This energy loss (and its related unusable heat) occurs between all trophic levels in a food web (see Figure 13.20). Although the efficiency with which energy is transferred from one level to the next varies among different types of organisms, it usually ranges between 5 and 20 percent. In other words, roughly 80 to 95 percent of the potential energy available at one trophic level is not transferred to the next one. This pattern of energy loss is often illustrated as a pyramid of productivity (see Figure 13.21 on page 448). 100 J feces 200 J plant material eaten by caterpillar 33 J cellular respiration 67 J growth Figure 13.19 Why is so much energy lost as waste from a caterpillar? The next MiniLab will help you understand pyramids of productivity by examining three typical food chains. heat (energy) loss to environment decomposers 100% energy to environment loss = 23.8% loss = 76.2% 5.5% loss = 23.5% loss = 71% carnivores 11.4% loss = 21.4% respiratory heat loss organic wastes/dead tissue top carnivores loss = 67.2% herbivores loss = 20.4% loss = 63.4% 16.2% producers photosynthesis Figure 13.20 Why is the respiratory heat loss of consumers higher than that of producers? Note that it is highest for top-level carnivores. 446 MHR • Unit 5 Population Dynamics 1–2% solar energy Earth Sun Since progressively less energy is transferred from lower to higher levels in a food web, less biomass can be produced at the higher trophic levels. This concept can be represented in a biomass pyramid, in which each tier represents the biomass of that trophic level (see Figure 13.22A on the following page). Typically, the shape of a biomass pyramid is similar to that of a pyramid of productivity. However, in some aquatic ecosystems, a relatively low biomass of primary producers (called phytoplankton) supports a higher biomass of primary consumers (zooplankton), as shown in Figure 13.22B. This BIO FACT Meat is more easily digested than most plant materials, so carnivores are slightly more efficient at converting food into biomass. However, carnivores typically use up more of this biomass during their own cellular respiration because their energy needs are higher than the energy needs of herbivores. Carnivores tend to move around more to find food, and many of them are endothermic. MINI LAB Where Do You Fit in the Food Chain? 2. How might diet influence the number of humans Earth can ultimately support? How might your knowledge of pyramids of productivity influence your decision about the type of foods you include in your diet? Recall that only a very small fraction of the energy released by the Sun is assimilated into plant material (see Figure 13.17 on page 444). For ease of calculation, assume that the amount of energy captured by plants and contained in their tissues is two percent of the total energy available from sunlight. Additionally (although it is a simplification), assume that 10 percent of the energy at one trophic level is transferred to the next level. Study the diagram and determine the percentage of the Sun’s energy available to humans (as shown at the top of each of the three food chains). 3. A square metre of land planted with rice produces about 5200 kJ of energy per year. A chicken farm produces about 800 kJ/m2 of potential food energy per year. Assume that a human must consume 2400 kJ per day to survive. Although it is an oversimplification to imply that a person could survive by eating only one type of food, calculate the total area of land needed to support the student population of your school for one year on a diet of: Analyze 1. About 80 percent of the world’s population eat mostly grain-based foods. Why do you think this is the case? (a) rice (b) chicken 4. Research the differences between the food used to feed chickens and other poultry in small family-run farms and the type of feed used in large, commercial agribusiness operations. Which do you think is more environmentally friendly? 2% of original energy Eighty percent of humans have a diet mostly of grain… Sun grain humans 2% …but many Canadians, like Americans and Europeans, also consume much meat and fish. 0.2% Sun grain beef 2% 0.02% 0.2% Sun phytoplankton humans zooplankton copepods 0.002% herring 0.0002% tuna humans Three typical food chains for humans with different diets. Chapter 13 Ecological Principles • MHR 447 Tertiary consumers 10 J Secondary consumers 100 J Primary consumers 1000 J Primary producers 10 000 J 1 000 000 J of sunlight occurs because the phytoplankton are eaten so quickly that there is no time for a large population to develop. The population of zooplankton can only exist because the phytoplankton have an extremely high reproductive rate — new organisms appear as fast as others are eaten. This means that the productivity of phytoplankton is very high, and the pyramid of productivity for this ecosystem is therefore wide at the top and narrow at the bottom. In every ecosystem, the biomass of carnivores at the highest trophic level is very limited. Only a tiny fraction of the chemical energy captured by photosynthesis flows all the way through a food web to a tertiary or higher-level consumer. Thus, most food webs are limited to five or fewer trophic levels — there is just not enough energy left to support more levels. Trophic level Dry weight (g/m2 ) 1.5 11 37 809 Tertiary consumers Secondary consumers Primary consumers Primary producers A Florida bog 21 4 Primary consumers (zooplankton) Primary producers (phytoplankton) B English Channel Figure 13.22 The biomass pyramid in (A) is based on data collected from a Florida bog. In the English Channel ecosystem, the pyramid of biomass is inverted (B). 448 MHR • Unit 5 Population Dynamics Figure 13.21 This figure is drawn to show a 10 percent efficiency of energy transfer from one trophic level to the next. Although the rate of efficiency varies (from 5 to 20 percent), 10 percent is a commonly used average figure. This gives rise to what is sometimes called “the rule of 10” when describing the shape of this pyramid. Animals that make up the highest trophic level in an ecosystem tend to be large, predatory species such as lions, whales, hawks, and eagles. Since biomass is limited at the top of the pyramid, there can only be a few of these large animals in any ecosystem at one time. In fact, when you compare the number of individual organisms at each trophic level, you will find that the same pyramidal shape appears. The pyramid of numbers in Figure 13.23A shows the effect of the decreasing energy supply on the number of individuals at each level. Although the supply of solar energy is almost limitless, almost all of the energy is eventually lost from ecosystems as a result of inefficient transfers between trophic levels. Not all pyramids of numbers have this shape. In a forest, for example, a few individual primary producers (trees) have enough biomass to support a large population of herbivores. As is true for the pyramid of biomass in some aquatic ecosystems, this could result in a pyramid of a different shape (see Figure 13.23B). Completing the Thinking Lab on the following page will demonstrate that it is not only energy that can be passed through a food web. Certain toxic compounds can cause serious damage to species in an ecosystem when those compounds are passed from one trophic level to another. In the next section, you will learn that nutrients are essential for proper growth and the repair of body tissue, and that they also cycle in ecosystems. Grassland top level consumer tertiary consumer secondary consumer primary consumer producer A Deciduous forest tertiary consumer secondary consumer primary consumer producer B BIO FACT The pyramid of productivity shows that primary consumers harvest more of the energy trapped during the process of photosynthesis (by eating photosynthetic organisms directly) than secondary consumers (who eat the organisms who ate the photosynthetic organisms). For humans, it is far more efficient in terms of obtaining energy to eat grain directly rather than to eat grain-fed beef. The pyramid of numbers shows that the biosphere could successfully feed far more humans if they were herbivores. Figure 13.23 (A) Many ecosystems have a trophic structure that produces a pyramid of numbers with a broad base. (B) In some ecosystems, however, there can be fewer producers than primary consumers. THINKING LAB Biological Magnification Background Certain toxic compounds are not easily broken down by decomposers, so they remain in the water or soil for long periods of time. This increases the probability that these compounds will be ingested by small organisms — which are then eaten by increasingly larger organisms and passed through food webs. Since each animal on a trophic level tends to eat many organisms from the level below, the amount of toxins taken in becomes magnified with each step up the food chain. These compounds, which accumulate in the fatty tissues of animals, have been observed to have diverse (and often harmful) effects on these organisms. These effects include damage to the nervous and reproductive systems as well as the production of genetic mutations leading to various forms of cancer. One of the first cases in which the phenomenon of biological magnification was observed involved DDT. DDT is a powerful insecticide that was used widely during World War II to control mosquitoes and other insects that transmitted diseases (such as malaria) to humans. Signs of damage caused by this chemical started to turn up about 10 years after it was first used. Mounting evidence of its deleterious effects caused the Canadian government to restrict the use of DDT after 1969. Study the the table on the right and the diagram on the following page, and answer the following questions. You Try It 1. What is the relationship between an organism’s trophic level and the concentration of DDT in its body? 2. How might an animal that lives a long distance from an area sprayed with DDT accumulate the chemical in its body? 3. Describe the general patterns you find in the data in the table. Speculate on the possible reasons for the differences in concentrations of DDT measured among species and localities and over time. 4. Research the specific types of prey consumed by one of the bird species listed in the table. Draw a biomass pyramid involving this species, incorporating trophic levels. Include the correct quantity of DDT (in ppb) at each level of your chart. DDT concentration in eggs (ppb) Species Year Bay of Fundy Atlantic Ocean Leach’s storm-petrel (Oceanodroma leucorhoa) (feeds on small organisms near the water surface) 1968 no data 1460 1972 6810 2480 1976 1750 750 1980 1130 460 1984 1050 400 1968 no data 890 1972 2570 760 1976 1270 590 1980 1030 550 1984 740 300 1972 6510 2850 1976 1490 2180 1980 1910 1340 1984 1070 1880 Atlantic puffin (Fratercula arctica) (feeds on small fish) double-crested cormorant (Phalacrocorax auritus) (feeds on larger fish) DDT concentration in the eggs of three species of sea birds breeding along Canada’s east coast Chapter 13 Ecological Principles • MHR 449 sea birds 2 800 ppb fish 43 ppb dolphins 5 200 ppb squid 22 ppb plankton 1.7 ppb seawater 0.0001 ppb Figure 13.24 An example of data collected on DDT in a food web of organisms in the north Pacific Ocean. Concentration of DDT is measured in parts per billion (ppb). The arrows indicate the flow of energy from one trophic level to another. SECTION 450 REVIEW 1. K/U Explain why the primary productivity of a grassland usually exceeds the primary productivity of a forest ecosystem covering an area of the same size. 2. Identify the factors that might cause annual fluctuations in the primary productivity of a specific grassland ecosystem. 3. On average, less than 20 percent of the food energy consumed by a grasshopper is converted into new grasshopper tissue. What happens to the rest of the food energy that was contained in the tissues of the producer species consumed by the grasshopper? 4. Explain why many models showing the relationships among trophic levels in an ecosystem are shown in the shape of pyramids. Describe the difference between the information depicted in a biomass pyramid, a pyramid of productivity, and a pyramid of numbers. 5. K/U Explain why the primary productivity levels of equatorial ecosystems generally exceed those of ecosystems situated farther north. 6. K/U Explain why the energy transfer from herbivores to carnivores is more efficient than the energy transfer from producers to herbivores within the same food chain. 7. K/U Suppose that the biomass for primary producers is less than the biomass for primary consumers. Can this ecosystem survive? Explain your answer. 8. Describe the factors that determine the shape of a pyramid of numbers diagram for a specific type of ecosystem. Explain why a grassland pyramid would have a wider base than a forest pyramid. 9. MC We have seen that there are fewer carnivores than herbivores in ecosystems because of the inefficiency of energy transfer between trophic levels, and that the world could support more people if we ate only plant material. Some people feel this means that humans should switch to a vegetarian diet; others disagree. There are, in fact, a variety of issues to consider in addition to the relatively simple one of energy transfer. Take a stand. Prepare your arguments carefully and be prepared to debate the issue in class. You might want to prepare a pamphlet that could be used to educate the public (or the rest of the class) about your point of view. K/U K/U K/U MHR • Unit 5 Population Dynamics K/U
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