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through food-web relationships (pp. 100–101). Matter is recy-  results in geographic patterns across the globe (Figure 5.8).
                        cled because when organisms die and decay, their nutrients   In terrestrial ecosystems, net primary productivity tends to
                        remain in the system. In contrast, most energy that organisms   increase with temperature and precipitation. In aquatic eco-
                        take in drives cellular respiration and is released as heat.  systems, net primary productivity tends to rise with light and
                                                                             the availability of nutrients.
                        Energy is converted to biomass

                        As autotrophs, such as green plants and phytoplankton, con-  Nutrients influence productivity
                        vert solar energy to the energy of chemical bonds in sug-
                        ars through photosynthesis (p. 50), they perform  primary   Nutrients are elements and compounds (pp. 41–44) that organ-
                          production. Specifically, the total amount of chemical energy   isms consume and require for survival. Organisms need sev-
                        produced by autotrophs is termed gross primary production.   eral dozen naturally occurring nutrients to survive. Elements
                        Autotrophs use most of this production to power their own   and compounds required in relatively large amounts (such as
                        metabolism by cellular respiration (p. 50). The energy that   nitrogen, carbon, and phosphorus) are called  macronutrients.
                        remains after respiration and that is used to generate bio-  Nutrients needed in small amounts are called micronutrients
                        mass (such as leaves, stems, and roots; p. 100) ecologists   (examples include zinc, copper, and iron).
                        call net primary production. Thus, net primary production   Nutrients  stimulate  production  by  plants,  and  lack  of
                        equals gross primary production minus the energy used in   nutrients can limit production. As mentioned earlier, the avail-
                        respiration.                                         ability of nitrogen or phosphorus frequently is a limiting fac-
                            Another way to think of net primary production is that it   tor (p. 85) for plant or algal growth. When these nutrients are
                        represents the energy or biomass available for consumption   added to a system, producers show the greatest response to
                        by heterotrophs. Some of this plant biomass is eaten by her-  whichever nutrient has been in shortest supply. Nitrogen tends
                        bivores. Plant matter not eaten by herbivores becomes fodder   to be limiting in marine systems, and phosphorus in fresh-
                        for detritivores and decomposers once the plant dies or drops   water systems, though both contribute to eutrophication in all
                        its leaves. Heterotrophs use the energy they gain from plant   waters. Thus eutrophication in the Chesapeake Bay is driven
                        biomass for their own metabolism, growth, and reproduction.   by excess nitrogen, whereas  eutrophication  in the  freshwa-
                        Some of this energy is used by heterotrophs to generate bio-  ter ponds and lakes in the bay’s watershed are spurred by
                        mass in their bodies (such as skin, muscle, or bone), which is   increases in phosphorus.
                        termed secondary production.                             Canadian ecologist David Schindler and others demon-
                            Ecosystems vary in the rate at which autotrophs convert   strated the effects of phosphorus on freshwater systems in
                        energy to biomass. The rate at which this conversion occurs   the 1970s by experimentally manipulating entire lakes. In
                        is termed productivity, and ecosystems whose plants  convert   one experiment, his team bisected a 16-ha (40-acre) lake in
                        solar energy to biomass rapidly are said to have high  net   Ontario with a plastic barrier. To one half the researchers
                          primary  productivity. Freshwater wetlands, tropical forests,   added carbon, nitrate, and phosphate; to the other they added
                        coral reefs, and algal beds tend to have the highest net primary   only carbon and nitrate. Soon after the experiment began,
                        productivities, whereas deserts, tundra, and open ocean tend   they witnessed a dramatic increase in algae in the half of
                        to have the lowest (Figure 5.7). Variation among ecosystems   the lake that received phosphate, whereas the other half (the
                        and among biomes (Chapter 4) in net primary productivity   control for the experiment, p. 29) continued to host algal





                                                                                                       Figure 5.7 Net primary
                                2500      Terrestrial ecosystems                                       productivity varies greatly   CHAPTER 5 • Envi R onm E n TA l S y STE m S   A nd E C o S y STE m E C ology
                                          Aquatic ecosystems
                             Net primary productivity  (g C/m 2 /yr)  1500                             forests, coral reefs, and algal
                                                                                                       between ecosystem types.
                                2000
                                                                                                       Freshwater wetlands, tropical
                                                                                                       beds show high values on
                                                                                                       average, whereas deserts,
                                1000
                                                                                                       tundra, and the open ocean
                                                                                                       show low values. Data from
                                 500
                                                                                                       Whittaker, R.H., 1975. Communities
                                                                                                       and ecosystems, 2nd ed. NewYork:
                                   0  Open ocean            Boreal forest Savanna  Estuaries           MacMillan.
                                                 Temperate grassland
                                            Lake and stream
                          Desert and semidesert shrub Tundra and alpine Continental shelf Cultivated land Temperate evergreen forest  Swamp and marsh Algal beds and reefs  129
                                                             Temperate deciduous forest
                                                                                     Tropical rainforest
                                                                           Tropical seasonal forest










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