Chapter 3: Ecosystem Ecology Chapter PDF

Summary

This chapter introduces ecosystem ecology, covering the concept of ecosystems, food webs, and biogeochemical cycles. It explores different types of ecosystems, including terrestrial and aquatic biomes, and their interactions with the environment. The chapter also explains how energy transfers and nutrient cycles in ecosystems work.

Full Transcript

**Chapter 3: Ecosystem Ecology** **Overview:** **Ecology** is the study of how living things interact with each other and with their environment. It is a major branch of biology, but has areas of overlap with geography, geology, climatology, and other sciences. The study of ecology begins with two...

**Chapter 3: Ecosystem Ecology** **Overview:** **Ecology** is the study of how living things interact with each other and with their environment. It is a major branch of biology, but has areas of overlap with geography, geology, climatology, and other sciences. The study of ecology begins with two fundamental concepts in ecology: the ecosystem and their organisms. **Objectives:** At the end of the chapter the students should be able to: 1. describe the concept of ecosystem. 2. discuss food web, its energy transfer and productivity within ecosystem. 3. trace and explain the steps of biogeochemical cycles. 4. describe the types and general features of terrestrial and aquatic biomes. 5. explain the ecological and economic value of different biomes. **Lesson 3.1: Ecology of the Ecosystem** **Overview:** An **ecosystem** is a unit of nature and the focus of study in ecology. It consists of all the biotic and abiotic factors in an area and their interactions. The abiotic and biotic parts of an ecosystem are linked together by flows of energy and cycles of nutrients through the system. An Ecosystems can vary in size. **Objectives:** **At the end of the lesson the students should be able to:** - Describe the basic types of ecosystems on Earth - Explain the methods that ecologists use to study ecosystem structure and dynamics - Identify the different methods of ecosystem modeling - Differentiate between food chains and food webs and recognize the importance of each **Content:** There is no widely agreed upon way to delineate a specific ecosystem. Theoretically, ecosystems can vary tremendously in size. Consider a forest as an example. It might cover hundreds or even thousands of acres, forming a large ecosystem in which an individual tree is of little consequence. However, an individual tree can also be considered an ecosystem, with millions of organisms living in and on it, ranging from microbes to small mammals. Even a single leaf can be considered an ecosystem. Several generations of an aphid population can exist over the lifespan of the leaf, as in the photo below. Each of the aphids, in turn, supports a diverse community of bacteria. Figure 3.1.1: Tiny insects called aphids live on and feed off this leaf. Along with bacteria, they form a miniature leaf ecosystem. (CC BY-NC 3.0; Thomas Bresson via [[Wikimedia.org]](https://commons.wikimedia.org/wiki/File:2013-10-03_17-58-42-Aphidoidea.JPG)) Life in an ecosystem is often about competition for limited resources, a characteristic of the theory of natural selection. Competition in communities (all living things within specific habitats) is observed both within species and among different species. The resources for which organisms compete include organic material from living or previously living organisms, sunlight, and mineral nutrients, which provide the energy for living processes and the matter to make up organisms' physical structures. Other critical factors influencing community dynamics are the components of its physical and geographic environment: a habitat's latitude, amount of rainfall, topography (elevation), and available species. These are all important environmental variables that determine which organisms can exist within a particular area. An ecosystem is a community of living organisms and their interactions with their abiotic (non-living) environment. Ecosystems can be small, such as in the leaf surface(figure above) or large, such as the Amazon Rainforest in Brazil (Figure below) ![Left photo shows a rocky tide pool with seaweed and snails. Right photo shows the Amazon Rainforest.](media/image2.jpeg)Figure 3.1.2 A (a) tidal pool ecosystem in Matinicus Island in Maine is a small ecosystem, while the (b) Amazon Rainforest in Brazil is a large ecosystem. (credit a: modification of work by "takomabibelot"/Flickr; credit b: modification of work by Ivan Mlinaric) There are three broad categories of ecosystems based on their general environment: freshwater, ocean water, and terrestrial. Within these broad categories are individual ecosystem types based on the organisms present and the type of environmental habitat. Ocean ecosystems are the most common, comprising 75 percent of the Earth\'s surface and consisting of three basic types: shallow ocean, deep ocean water, and deep ocean surfaces (the low depth areas of the deep oceans). The shallow ocean ecosystems include extremely biodiverse coral reef ecosystems, and the deep ocean surface is known for its large numbers of plankton and krill (small crustaceans) that support it. These two environments are especially important to aerobic respirators worldwide as the phytoplankton perform 40 percent of all photosynthesis on Earth. Although not as diverse as the other two, deep ocean ecosystems contain a wide variety of marine organisms. Such ecosystems exist even at the bottom of the ocean where light is unable to penetrate through the water. Freshwater ecosystems are the rarest, occurring on only 1.8 percent of the Earth\'s surface. Lakes, rivers, streams, and springs comprise these systems; they are quite diverse, and they support a variety of fish, amphibians, reptiles, insects, phytoplankton, fungi, and bacteria. Terrestrial ecosystems, also known for their diversity, are grouped into large categories called biomes, such as tropical rain forests, savannas, deserts, coniferous forests, deciduous forests, and tundra. Grouping these ecosystems into just a few biome categories obscures the great diversity of the individual ecosystems within them. For example, there is great variation in desert vegetation: the saguaro cacti and other plant life in the Sonoran Desert, in the United States, are relatively abundant compared to the desolate rocky desert of Boa Vista, an island off the coast of Western Africa. Photo (a) shows saguaro cacti that look like telephone poles with arms extended from them. Photo (b) shows a barren plain of red soil littered with rocks.Figure 3.1.3: Desert ecosystems, like all ecosystems, can vary greatly. The desert in (a) Saguaro National Park, Arizona, has abundant plant life, while the rocky desert of (b) Boa Vista island, Cape Verde, Africa, is devoid of plant life. (credit a: modification of work by Jay Galvin; credit b: modification of work by Ingo Wölbern) Ecosystems are complex with many interacting parts. They are routinely exposed to various disturbances, or changes in the environment that effect their compositions: yearly variations in rainfall and temperature and the slower processes of plant growth, which may take several years. Many of these disturbances are a result of natural processes. For example, when lightning causes a forest fire and destroys part of a forest ecosystem, the ground is eventually populated by grasses, then by bushes and shrubs, and later by mature trees, restoring the forest to its former state. The impact of environmental disturbances caused by human activities is as important as the changes wrought by natural processes. Human agricultural practices, air pollution, acid rain, global deforestation, overfishing, eutrophication, oil spills, and illegal dumping on land and into the ocean are all issues of concern to conservationists. Equilibrium is the steady state of an ecosystem where all organisms are in balance with their environment and with each other. In ecology, two parameters are used to measure changes in ecosystems: resistance and resilience. The ability of an ecosystem to remain at equilibrium in spite of disturbances is called resistance. The speed at which an ecosystem recovers equilibrium after being disturbed, called its resilience. Ecosystem resistance and resilience are especially important when considering human impact. The nature of an ecosystem may change to such a degree that it can lose its resilience entirely. This process can lead to the complete destruction or irreversible altering of the ecosystem. **Food Chains and Food Webs** ![](media/image4.jpeg)The term "food chain" is sometimes used metaphorically to describe human social situations. In this sense, food chains are thought of as a competition for survival, such as "who eats whom?" Someone eats and someone is eaten. Therefore, it is not surprising that in our competitive "dog-eat-dog" society, individuals who are considered successful are seen as being at the top of the food chain, consuming all others for their benefit, whereas the less successful are seen as being at the bottom. The scientific understanding of a food chain is more precise than in its everyday usage. In ecology, a food chain is a linear sequence of organisms through which nutrients and energy pass: primary producers, primary consumers, and higher-level consumers are used to describe ecosystem structure and dynamics. There is a single path through the chain. Each organism in a food chain occupies what is called a trophic level. Depending on their role as producers or consumers, species or groups of species can be assigned to various trophic levels. In many ecosystems, the bottom of the food chain consists of photosynthetic organisms (plants and/or phytoplankton), which are called primary producers. The organisms that consume the primary producers are herbivores: the primary consumers. Secondary consumers are usually carnivores that eat the primary consumers. Tertiary consumers are carnivores that eat other carnivores. Higher-level consumers feed on the next lower tropic levels, and so on, up to the organisms at the top of the food chain: the apex consumers. In the Lake Ontario food chain shown in Figure above, the Chinook salmon is the apex consumer at the top of this food chain. These are the trophic levels of a food chain in Lake Ontario at the United States-Canada border. Energy and nutrients flow from photosynthetic green algae at the bottom to the top of the food chain: the Chinook salmon. One major factor that limits the length of food chains is energy. Energy is lost as heat between each trophic level due to the second law of thermodynamics. Thus, after a limited number of trophic energy transfers, the amount of energy remaining in the food chain may not be great enough to support viable populations at yet a higher trophic level. The loss of energy between trophic levels is illustrated by the pioneering studies of Howard T. Odum in the Silver Springs, Florida, ecosystem in the 1940s (Figure 3.1.4). The primary producers generated 20,819 kcal/m^2^/yr (kilocalories per square meter per year), the primary consumers generated 3368 kcal/m^2^/yr, the secondary consumers generated 383 kcal/m^2^/yr, and the tertiary consumers only generated 21 kcal/m^2^/yr. Thus, there is little energy remaining for another level of consumers in this ecosystem. Graph shows energy content in different trophic levels. The energy content of primary producers is over 20,000 kilocalories per meter squared per year. The energy content of primary consumers is much smaller, about 3,400 kilocalories per meter squared per year. The energy content of secondary consumers is 383 kilocalories per meter squared per year, and the energy content of tertiary consumers is only 21 kilocalories per meter squared per year. Figure 3.1.4: The relative energy in trophic levels in a Silver Springs, Florida, ecosystem is shown. Each trophic level has less energy available and supports fewer organisms at the next level. ![](media/image6.jpeg)There is a one problem when using food chains to accurately describe most ecosystems. Even when all organisms are grouped into appropriate trophic levels, some of these organisms can feed on species from more than one trophic level; likewise, some of these organisms can be eaten by species from multiple trophic levels. In other words, the linear model of ecosystems, the food chain, is not completely descriptive of ecosystem structure. A holistic model---which accounts for all the interactions between different species and their complex interconnected relationships with each other and with the environment---is a more accurate and descriptive model for ecosystems. A food web is a graphic representation of a holistic, non-linear web of primary producers, primary consumers, and higher-level consumers used to describe ecosystem structure and dynamics. Figure 3.1.5: This food web shows the interactions between organisms across trophic levels in the Lake Ontario ecosystem. Primary producers are outlined in green, primary consumers in orange, secondary consumers in blue, and tertiary (apex) consumers in purple. Arrows point from an organism that is consumed to the organism that consumes it. Notice how some lines point to more than one trophic level. For example, the opossum shrimp eats both primary producers and primary consumers. (credit: NOAA, GLERL) A comparison of the two types of structural ecosystem models shows strength in both. Food chains are more flexible for analytical modeling, are easier to follow, and are easier to experiment with, whereas food web models more accurately represent ecosystem structure and dynamics, and data can be directly used as input for simulation modeling. *.* Two general types of food webs are often shown interacting within a single ecosystem. A grazing food web (such as the Lake Ontario food web in Figure 46.1.5) has plants or other photosynthetic organisms at its base, followed by herbivores and various carnivores. A detrital food web consists of a base of organisms that feed on decaying organic matter (dead organisms), called decomposers or detritivores. These organisms are usually bacteria or fungi that recycle organic material back into the biotic part of the ecosystem as they themselves are consumed by other organisms. As all ecosystems require a method to recycle material from dead organisms, most grazing food webs have an associated detrital food web. For example, in a meadow ecosystem, plants may support a grazing food web of different organisms, primary and other levels of consumers, while at the same time supporting a detrital food web of bacteria, fungi, and detrivorous invertebrates feeding off dead plants and animals. **Summary:** Equilibrium is the steady state of an ecosystem where all organisms are in balance with their environment and with each other. In ecology, two parameters are used to measure changes in ecosystems: resistance and resilience. Life in an ecosystem is often about competition for limited resources, a characteristic of the theory of natural selection. Competition in communities (all living things within specific habitats) is observed both within species and among different species. A food chain is a linear sequence of organisms through which nutrients and energy pass: primary producers, primary consumers, and higher-level consumers are used to describe ecosystem structure and dynamics **Enrichment:** **Multiple choice** 1. All organism in an ecosystem are in balance with their environment. a.resilience b\. resistance c\. equilibrium 2\. The ability of an ecosystem to remain at equilibrium after being disturbed. a.resilience b\. resistance c\. equilibrium 3\. The speed at which an ecosystem recover after disturbance. a.resilience b\. resistance c\. equilibrium 4.The linear sequence of organisms through nutrients and energy a\. food web b\. food chain c\. energy pyramid 5\. The primary producers in an ecosystem a\. photosynthetic organisms b\. herbivores c\. Carnivores **Assignment:** 1\..Construct a food chain in an agricultural ecosystem. 2.Construct a grazing and detrimental ecosystem in an African oil Palm Plantation. 3.Prepare resistance and resiliency Activity plan of your community inorder to attain the new normal (equilibrium) after the pandemic. Sample table +-----------------------+-----------------------+-----------------------+ | Resistance | Activity | Person / Agency | +=======================+=======================+=======================+ | | 1.Observe Safety | Everybody | | | protocol: | | | | | | | | a\. hand washing... | | +-----------------------+-----------------------+-----------------------+ | Resilience | Activity | Person / Agency | +-----------------------+-----------------------+-----------------------+ | | Be obedient to the | DOH, LGU and all | | | DOH and LGU... | populace | +-----------------------+-----------------------+-----------------------+ **Lesson 3.2:** **Energy Flow through** **Ecosystems** **Overview:** All living things require energy in one form or another. Energy is required by most complex metabolic pathways (often in the form of adenosine triphosphate, ATP), especially those responsible for building large molecules from smaller compounds, and life itself is an energy-driven process. Living organisms would not be able to assemble macromolecules (proteins, lipids, nucleic acids, and complex carbohydrates) from their monomeric subunits without a constant energy input. It is important to understand how organisms acquire energy and how that energy is passed from one organism to another through food webs and their constituent food chains. Food webs illustrate how energy flows directionally through ecosystems, including how efficiently organisms acquire it, use it, and how much remains for use by other organisms of the food web. **Objectives:** **At the end of the lesson the students should be able to :** - Describe how organisms acquire energy in a food web and in associated food chains - Explain how the efficiency of energy transfers between trophic levels affects ecosystem structure and dynamics - Discuss trophic levels and how ecological pyramids are used to model them **Content:** How Organisms Acquire Energy in a Food Web Energy is acquired by living things in three ways: photosynthesis, chemosynthesis, and the consumption and digestion of other living or previously living organisms by heterotrophs. Photosynthetic and chemosynthetic organisms are both grouped into a category known as autotrophs: organisms capable of synthesizing their own food (more specifically, capable of using inorganic carbon as a carbon source). Photosynthetic autotrophs (photoautotrophs) use sunlight as an energy source, whereas chemosynthetic autotrophs (chemoautotrophs) use inorganic molecules as an energy source. Autotrophs are critical for all ecosystems. Without these organisms, energy would not be available to other living organisms and life itself would not be possible. Photoautotrophs, such as plants, algae, and photosynthetic bacteria, serve as the energy source for a majority of the world's ecosystems. These ecosystems are often described by grazing food webs. Photoautotrophs harness the solar energy of the sun by converting it to chemical energy in the form of ATP (and NADP). The energy stored in ATP is used to synthesize complex organic molecules, such as glucose. Chemoautotrophs are primarily bacteria that are found in rare ecosystems where sunlight is not available, such as in those associated with dark caves or hydrothermal vents at the bottom of the ocean (Figure 3.2.1). Many chemoautotrophs in hydrothermal vents use hydrogen sulfide (H~2~S), which is released from the vents as a source of chemical energy. This allows chemoautotrophs to synthesize complex organic molecules, such as glucose, for their own energy and in turn supplies energy to the rest of the ecosystem. Photo shows shrimp, lobster, and white crabs crawling on a rocky ocean floor littered with mussels. Figure 3.2.1: Swimming shrimp, a few squat lobsters, and hundreds of vent mussels are seen at a hydrothermal vent at the bottom of the ocean. As no sunlight penetrates to this depth, the ecosystem is supported by chemoautotrophic bacteria and organic material that sinks from the ocean's surface. This picture was taken in 2006 at the submerged NW Eifuku volcano off the coast of Japan by the National Oceanic and Atmospheric Administration (NOAA). The summit of this highly active volcano lies 1535 m below the surface. Productivity within Trophic Levels Productivity within an ecosystem can be defined as the percentage of energy entering the ecosystem incorporated into biomass in a particular trophic level. Biomass is the total mass, in a unit area at the time of measurement, of living or previously living organisms within a trophic level. Ecosystems have characteristic amounts of biomass at each trophic level. For example, in the English Channel ecosystem the primary producers account for a biomass of 4 g/m^2^ (grams per meter squared), while the primary consumers exhibit a biomass of 21 g/m^2^. The productivity of the primary producers is especially important in any ecosystem because these organisms bring energy to other living organisms by photoautotrophy or chemoautotrophy. The rate at which photosynthetic primary producers incorporate energy from the sun is called gross primary productivity. An example of gross primary productivity is shown in the compartment diagram of energy flow within the Silver Springs aquatic ecosystem as shown (\[link\]). In this ecosystem, the total energy accumulated by the primary producers (gross primary productivity) was shown to be 20,810 kcal/m^2^/yr. Because all organisms need to use some of this energy for their own functions (like respiration and resulting metabolic heat loss) scientists often refer to the net primary productivity of an ecosystem. Net primary productivity is the energy that remains in the primary producers after accounting for the organisms' respiration and heat loss. The net productivity is then available to the primary consumers at the next trophic level. In our Silver Spring example, 13,187 of the 20,810 kcal/m^2^/yr were used for respiration or were lost as heat, leaving 7,632 kcal/m^2^/yr of energy for use by the primary consumers. Ecological Efficiency: The Transfer of Energy between Trophic Levels As illustrated in Figure 46.2.346.2.3, large amounts of energy are lost from the ecosystem from one trophic level to the next level as energy flows from the primary producers through the various trophic levels of consumers and decomposers. The main reason for this loss is the second law of thermodynamics, which states that whenever energy is converted from one form to another, there is a tendency toward disorder (entropy) in the system. In biologic systems, this means a great deal of energy is lost as metabolic heat when the organisms from one trophic level consume the next level. In the Silver Springs ecosystem example (Figure 46.1.1), we see that the primary consumers produced 1103 kcal/m^2^/yr from the 7618 kcal/m^2^/yr of energy available to them from the primary producers. The measurement of energy transfer efficiency between two successive trophic levels is termed the trophic level transfer efficiency (TLTE) and is defined by the formula: TLTE=production at present trophic levelproduction at previous trophic level∗100(46.2.1)(46.2.1)TLTE=production at present trophic levelproduction at previous trophic level∗100 In Silver Springs, the TLTE between the first two trophic levels was approximately 14.8 percent. The low efficiency of energy transfer between trophic levels is usually the major factor that limits the length of food chains observed in a food web. The fact is, after four to six energy transfers, there is not enough energy left to support another trophic level. In the Lake Ontario example shown in Figure 46.2.346.2.3, only three energy transfers occurred between the primary producer, (green algae), and the apex consumer (Chinook salmon). Ecologists have many different methods of measuring energy transfers within ecosystems. Some transfers are easier or more difficult to measure depending on the complexity of the ecosystem and how much access scientists have to observe the ecosystem. In other words, some ecosystems are more difficult to study than others, and sometimes the quantification of energy transfers has to be estimated. Another main parameter that is important in characterizing energy flow within an ecosystem is the net production efficiency. Net production efficiency (NPE) allows ecologists to quantify how efficiently organisms of a particular trophic level incorporate the energy they receive into biomass; it is calculated using the following formula: NPE=net consumer productivityassimilation∗100(46.2.2)(46.2.2)NPE=net consumer productivityassimilation∗100 Net consumer productivity is the energy content available to the organisms of the next trophic level. Assimilation is the biomass (energy content generated per unit area) of the present trophic level after accounting for the energy lost due to incomplete ingestion of food, energy used for respiration, and energy lost as waste. Incomplete ingestion refers to the fact that some consumers eat only a part of their food. For example, when a lion kills an antelope, it will eat everything except the hide and bones. The lion is missing the energy-rich bone marrow inside the bone, so the lion does not make use of all the calories its prey could provide. Thus, NPE measures how efficiently each trophic level uses and incorporates the energy from its food into biomass to fuel the next trophic level. In general, cold-blooded animals (ectotherms), such as invertebrates, fish, amphibians, and reptiles, use less of the energy they obtain for respiration and heat than warm-blooded animals (endotherms), such as birds and mammals. The extra heat generated in endotherms, although an advantage in terms of the activity of these organisms in colder environments, is a major disadvantage in terms of NPE. Therefore, many endotherms have to eat more often than ectotherms to get the energy they need for survival. In general, NPE for ectotherms is an order of magnitude (10x) higher than for endotherms. For example, the NPE for a caterpillar eating leaves has been measured at 18 percent, whereas the NPE for a squirrel eating acorns may be as low as 1.6 percent. The inefficiency of energy use by warm-blooded animals has broad implications for the world\'s food supply. It is widely accepted that the meat industry uses large amounts of crops to feed livestock, and because the NPE is low, much of the energy from animal feed is lost. For example, it costs about 1¢ to produce 1000 dietary calories (kcal) of corn or soybeans, but approximately \$0.19 to produce a similar number of calories growing cattle for beef consumption. The same energy content of milk from cattle is also costly, at approximately \$0.16 per 1000 kcal. Much of this difference is due to the low NPE of cattle. Thus, there has been a growing movement worldwide to promote the consumption of non-meat and non-dairy foods so that less energy is wasted feeding animals for the meat industry. Modeling Ecosystems Energy Flow: Ecological Pyramids The structure of ecosystems can be visualized with ecological pyramids, which were first described by the pioneering studies of Charles Elton in the 1920s. Ecological pyramids show the relative amounts of various parameters (such as number of organisms, energy, and biomass) across trophic levels. Pyramids of numbers can be either upright or inverted, depending on the ecosystem. As shown in \[link\], typical grassland during the summer has a base of many plants and the numbers of organisms decrease at each trophic level. However, during the summer in a temperate forest, the base of the pyramid consists of few trees compared with the number of primary consumers, mostly insects. Because trees are large, they have great photosynthetic capability, and dominate other plants in this ecosystem to obtain sunlight. Even in smaller numbers, primary producers in forests are still capable of supporting other trophic levels. Another way to visualize ecosystem structure is with pyramids of biomass. This pyramid measures the amount of energy converted into living tissue at the different trophic levels. Using the Silver Springs ecosystem example, this data exhibits an upright biomass pyramid (Figure 3.2..2), whereas the pyramid from the English Channel example is inverted. The plants (primary producers) of the Silver Springs ecosystem make up a large percentage of the biomass found there. However, the phytoplankton in the English Channel example make up less biomass than the primary consumers, the zooplankton. As with inverted pyramids of numbers, this inverted pyramid is not due to a lack of productivity from the primary producers, but results from the high turnover rate of the phytoplankton. The phytoplankton are consumed rapidly by the primary consumers, thus, minimizing their biomass at any particular point in time. However, phytoplankton reproduce quickly, thus they are able to support the rest of the ecosystem. ![](media/image8.png)Pyramid ecosystem modeling can also be used to show energy flow through the trophic levels. Notice that these numbers are the same as those used in the energy flow compartment diagram in Figure 3.2.2. Pyramids of energy are always upright, and an ecosystem without sufficient primary productivity cannot be supported. All types of ecological pyramids are useful for characterizing ecosystem structure. However, in the study of energy flow through the ecosystem, pyramids of energy are the most consistent and representative models of ecosystem structure. Figure 3..2.2: Ecological pyramids depict the (a) biomass, (b) number of organisms, and (c) energy in each trophic level. Pyramids depicting the number of organisms or biomass may be inverted, upright, or even diamond-shaped. Energy pyramids, however, are always upright. Why? Consequences of Food Webs: Biological Magnification One of the most important environmental consequences of ecosystem dynamics is biomagnification. Biomagnification is the increasing concentration of persistent, toxic substances in organisms at each trophic level, from the primary producers to the apex consumers. Many substances have been shown to bioaccumulate, including classical studies with the pesticide **d**ichloro**d**iphenyl**t**richloroethane (DDT), which was published in the 1960s bestseller, *Silent Spring*, by Rachel Carson. DDT was a commonly used pesticide before its dangers became known. In some aquatic ecosystems, organisms from each trophic level consumed many organisms of the lower level, which caused DDT to increase in birds (apex consumers) that ate fish. Thus, the birds accumulated sufficient amounts of DDT to cause fragility in their eggshells. This effect increased egg breakage during nesting and was shown to have adverse effects on these bird populations. The use of DDT was banned in the United States in the 1970s.Other substances that biomagnify are polychlorinated biphenyls (PCBs), which were used in coolant liquids in the United States until their use was banned in 1979, and heavy metals, such as mercury, lead, and cadmium. These substances were best studied in aquatic ecosystems, where fish species at different trophic levels accumulate toxic substances brought through the ecosystem by the primary producers. As illustrated in a study performed by the National Oceanic and Atmospheric Administration (NOAA) in the Saginaw Bay of Lake Huron , PCB concentrations increased from the ecosystem's primary producers (phytoplankton) through the different trophic levels of fish species. The apex consumer (walleye) has more than four times the amount of PCBs compared to phytoplankton. Also, based on results from other studies, birds that eat these fish may have PCB levels at least one order of magnitude higher than those found in the lake fish. Figure 3.2.3: This chart shows the PCB concentrations found at the various trophic levels in the Saginaw Bay ecosystem of Lake Huron. Numbers on the x-axis reflect enrichment with heavy isotopes of nitrogen (^15^N), which is a marker for increasing trophic level. Notice that the fish in the higher trophic levels accumulate more PCBs than those in lower trophic levels. (credit: Patricia Van Hoof, NOAA, GLERL) Other cncerns have been raised by the accumulation of heavy metals, such as mercury and cadmium, in certain types of seafood. The United States Environmental Protection Agency (EPA) recommends that pregnant women and young children should not consume any swordfish, shark, king mackerel, or tilefish because of their high mercury content. These individuals are advised to eat fish low in mercury: salmon, tilapia, shrimp, pollock, and catfish. Biomagnification is a good example of how ecosystem dynamics can affect our everyday lives, even influencing the food we eat. Summary Organisms in an ecosystem acquire energy in a variety of ways, which is transferred between trophic levels as the energy flows from the bottom to the top of the food web, with energy being lost at each transfer. The efficiency of these transfers is important for understanding the different behaviors and eating habits of warm-blooded versus cold-blooded animals. Modeling of ecosystem energy is best done with ecological pyramids of energy, although other ecological pyramids provide other vital information about ecosystem structure. **Enrichment:** A. Multiple choice: The weight of living organisms in an ecosystem at a particular point in time is called: 1. energy 2. production 3. entropy 4. biomass Which term describes the process whereby toxic substances increase along trophic levels of an ecosystem? 1. biomassification 2. biomagnification 3. bioentropy 4. heterotrophy Organisms that can make their own food using inorganic molecules are called: 1. autotrophs 2. heterotrophs 3. photoautotrophs 4. chemoautotrophs In the English Channel ecosystem, the number of primary producers is smaller than the number of primary consumers because\_\_\_\_\_\_\_\_. 1. the apex consumers have a low turnover rate 2. the primary producers have a low turnover rate 3. the primary producers have a high turnover rate 4. the primary consumers have a high turnover rate What law of chemistry determines how much energy can be transferred when it is converted from one form to another? 1. the first law of thermodynamics 2. the second law of thermodynamics 3. the conservation of matter 4. the conservation of energy B.1.Compare the three types of ecological pyramids and how well they describe ecosystem structure. Identify which ones can be inverted and give an example of an inverted pyramid for each. 2.How does the amount of food a warm blooded-animal (endotherm) eats relate to its net production efficiency (NPE)? 3\. Pyramids depicting the number of organisms or biomass may be inverted, upright, or even diamond-shaped. Energy pyramids, however, are always upright. Why?. **\ ** **Lesson 3.3 Biogeochemical Cycles** **Overview:** Mineral nutrients are cycled through ecosystems and their environment. Of particular importance are water, carbon, nitrogen, phosphorus, and sulfur. All of these cycles have major impacts on ecosystem structure and function.. The health of Earth depends on understanding these cycles and how to protect the environment from irreversible damage. **Objective** - Discuss the biogeochemical cycles of water, carbon, nitrogen, phosphorus, and sulfur - Explain how human activities have impacted these cycles and the potential consequences for Earth - Suggest some activities to protect the biogeochemical cycle of water, carbon, nitrogen, phosphorus and sulfur. **Content:** Energy flows directionally through ecosystems, entering as sunlight (or inorganic molecules for chemoautotrophs) and leaving as heat during the many transfers between trophic levels. However, the matter that makes up living organisms is conserved and recycled. The six most common elements associated with organic molecules---carbon, nitrogen, hydrogen, oxygen, phosphorus, and sulfur---take a variety of chemical forms and may exist for long periods in the atmosphere, on land, in water, or beneath the Earth's surface. Geologic processes, such as weathering, erosion, water drainage, and the subduction of the continental plates, all play a role in this recycling of materials. Because geology and chemistry have major roles in the study of this process, the recycling of inorganic matter between living organisms and their environment is called a biogeochemical cycle. Water contains hydrogen and oxygen, which is essential to all living processes. The hydrosphere is the area of the Earth where water movement and storage occurs: as liquid water on the surface and beneath the surface or frozen (rivers, lakes, oceans, groundwater, polar ice caps, and glaciers), and as water vapor in the atmosphere. Carbon is found in all organic macromolecules and is an important constituent of fossil fuels. Nitrogen is a major component of our nucleic acids and proteins and is critical to human agriculture. Phosphorus, a major component of nucleic acid (along with nitrogen), is one of the main ingredients in artificial fertilizers used in agriculture and their associated environmental impacts on our surface water. Sulfur, critical to the 3--D folding of proteins (as in disulfide binding), is released into the atmosphere by the burning of fossil fuels, such as coal.The cycling of these elements is interconnected. For example, the movement of water is critical for the leaching of nitrogen and phosphate into rivers, lakes, and oceans. Furthermore, the ocean itself is a major reservoir for carbon. Thus, mineral nutrients are cycled, either rapidly or slowly, through the entire biosphere, from one living organism to another, and between the biotic and abiotic world. The Water (Hydrologic) Cycle Water is the basis of all living processes. The human body is more than 1/2 water and human cells are more than 70 percent water. Thus, most land animals need a supply of fresh water to survive. However, when examining the stores of water on Earth, 97.5 percent of it is non-potable salt water (Figure 3.3.1). Of the remaining water, 99 percent is locked underground as water or as ice. Thus, less than 1 percent of fresh water is easily accessible from lakes and rivers. Many living things, such as plants, animals, and fungi, are dependent on the small amount of fresh surface water supply, a lack of which can have massive effects on ecosystem dynamics. Humans, of course, have developed technologies to increase water availability, such as digging wells to harvest groundwater, storing rainwater, and using desalination to obtain drinkable water from the ocean. Although this pursuit of drinkable water has been ongoing throughout human history, the supply of fresh water is still a major issue in modern times. ![](media/image10.jpeg)Figure 3.3.1: Only 2.5 percent of water on Earth is fresh water, and less than 1 percent of fresh water is easily accessible to living things. Water cycling is extremely important to ecosystem dynamics. Water has a major influence on climate and, thus, on the environments of ecosystems, some located on distant parts of the Earth. Most of the water on Earth is stored for long periods in the oceans, underground, and as ice. Figure 46.3.246.3.2 illustrates the average time that an individual water molecule may spend in the Earth's major water reservoirs. Residence time is a measure of the average time an individual water molecule stays in a particular reservoir. A large amount of the Earth's water is locked in place in these reservoirs as ice, beneath the ground, and in the ocean, and, thus, is unavailable for short-term cycling (only surface water can evaporate). Figure 3.3.2: This graph shows the average residence time for water molecules in the Earth's water reservoirs. There are various processes that occur during the cycling of water, shown in Figure 46.3.346.3.3. These processes include the following: - evaporation/sublimation - condensation/precipitation - subsurface water flow - surface runoff/snowmelt - streamflow The water cycle is driven by the sun's energy as it warms the oceans and other surface waters. This leads to the evaporation (water to water vapor) of liquid surface water and the sublimation (ice to water vapor) of frozen water, which deposits large amounts of water vapor into the atmosphere. Over time, this water vapor condenses into clouds as liquid or frozen droplets and is eventually followed by precipitation (rain or snow), which returns water to the Earth's surface. Rain eventually permeates into the ground, where it may evaporate again if it is near the surface, flow beneath the surface, or be stored for long periods. More easily observed is surface runoff: the flow of fresh water either from rain or melting ice. Runoff can then make its way through streams and lakes to the oceans or flow directly to the oceans themselves. *.* ![](media/image12.jpeg)Rain and surface runoff are major ways in which minerals, including carbon, nitrogen, phosphorus, and sulfur, are cycled from land to water. The environmental effects of runoff will be discussed later as these cycles are described. Figure 3.3.3: Water from the land and oceans enters the atmosphere by evaporation or sublimation, where it condenses into clouds and falls as rain or snow. Precipitated water may enter freshwater bodies or infiltrate the soil. The cycle is complete when surface or groundwater reenters the ocean. (credit: modification of work by John M. Evans and Howard Perlman, USGS) The Carbon Cycle Carbon is the second most abundant element in living organisms. Carbon is present in all organic molecules, and its role in the structure of macromolecules is of primary importance to living organisms. Carbon compounds contain especially high energy, particularly those derived from fossilized organisms, mainly plants, which humans use as fuel. Since the 1800s, the number of countries using massive amounts of fossil fuels has increased. Since the beginning of the Industrial Revolution, global demand for the Earth's limited fossil fuel supplies has risen; therefore, the amount of carbon dioxide in our atmosphere has increased. This increase in carbon dioxide has been associated with climate change and other disturbances of the Earth's ecosystems and is a major environmental concern worldwide. Thus, the "carbon footprint" is based on how much carbon dioxide is produced and how much fossil fuel countries consume. The carbon cycle is most easily studied as two interconnected sub-cycles: one dealing with rapid carbon exchange among living organisms and the other dealing with the long-term cycling of carbon through geologic processes. The entire carbon cycle is shown in Figure 3.3.4. Figure 3.3.4: Carbon dioxide gas exists in the atmosphere and is dissolved in water. Photosynthesis converts carbon dioxide gas to organic carbon, and respiration cycles the organic carbon back into carbon dioxide gas. Long-term storage of organic carbon occurs when matter from living organisms is buried deep underground and becomes fossilized. Volcanic activity and, more recently, human emissions, bring this stored carbon back into the carbon cycle. (credit: modification of work by John M. Evans and Howard Perlman, USGS) The Biological Carbon Cycle Living organisms are connected in many ways, even between ecosystems. A good example of this connection is the exchange of carbon between autotrophs and heterotrophs within and between ecosystems by way of atmospheric carbon dioxide. Carbon dioxide is the basic building block that most autotrophs use to build multi-carbon, high energy compounds, such as glucose. The energy harnessed from the sun is used by these organisms to form the covalent bonds that link carbon atoms together. These chemical bonds thereby store this energy for later use in the process of respiration. Most terrestrial autotrophs obtain their carbon dioxide directly from the atmosphere, while marine autotrophs acquire it in the dissolved form (carbonic acid, H~2~CO~3~^−^). However carbon dioxide is acquired, a by-product of the process is oxygen. The photosynthetic organisms are responsible for depositing approximately 21 percent oxygen content of the atmosphere that we observe today. Heterotrophs and autotrophs are partners in biological carbon exchange (especially the primary consumers, largely herbivores). Heterotrophs acquire the high-energy carbon compounds from the autotrophs by consuming them, and breaking them down by respiration to obtain cellular energy, such as ATP. The most efficient type of respiration, aerobic respiration, requires oxygen obtained from the atmosphere or dissolved in water. Thus, there is a constant exchange of oxygen and carbon dioxide between the autotrophs (which need the carbon) and the heterotrophs (which need the oxygen). Gas exchange through the atmosphere and water is one way that the carbon cycle connects all living organisms on Earth. The Biogeochemical Carbon Cycle The movement of carbon through the land, water, and air is complex, and in many cases, it occurs much more slowly geologically than as seen between living organisms. Carbon is stored for long periods in what are known as carbon reservoirs, which include the atmosphere, bodies of liquid water (mostly oceans), ocean sediment, soil, land sediments (including fossil fuels), and the Earth's interior. ![](media/image14.jpeg)As stated, the atmosphere is a major reservoir of carbon in the form of carbon dioxide and is essential to the process of photosynthesis. The level of carbon dioxide in the atmosphere is greatly influenced by the reservoir of carbon in the oceans. The exchange of carbon between the atmosphere and water reservoirs influences how much carbon is found in each location, and each one affects the other reciprocally. Carbon dioxide (CO~2~) from the atmosphere dissolves in water and combines with water molecules to form carbonic acid, and then it ionizes to carbonate and bicarbonate ions (Figure 3.3.5) Figure 3.3.5: Carbon dioxide reacts with water to form bicarbonate and carbonate ions. The equilibrium coefficients are such that more than 90 percent of the carbon in the ocean is found as bicarbonate ions. Some of these ions combine with seawater calcium to form calcium carbonate (CaCO~3~), a major component of marine organism shells. These organisms eventually form sediments on the ocean floor. Over geologic time, the calcium carbonate forms limestone, which comprises the largest carbon reservoir on Earth. On land, carbon is stored in soil as a result of the decomposition of living organisms (by decomposers) or from weathering of terrestrial rock and minerals. This carbon can be leached into the water reservoirs by surface runoff. Deeper underground, on land and at sea, are fossil fuels: the anaerobically decomposed remains of plants that take millions of years to form. Fossil fuels are considered a non-renewable resource because their use far exceeds their rate of formation. A non-renewable resource, such as fossil fuel, is either regenerated very slowly or not at all. Another way for carbon to enter the atmosphere is from land (including land beneath the surface of the ocean) by the eruption of volcanoes and other geothermal systems. Carbon sediments from the ocean floor are taken deep within the Earth by the process of subduction: the movement of one tectonic plate beneath another. Carbon is released as carbon dioxide when a volcano erupts or from volcanic hydrothermal vents. Carbon dioxide is also added to the atmosphere by the animal husbandry practices of humans. The large numbers of land animals raised to feed the Earth's growing population results in increased carbon dioxide levels in the atmosphere due to farming practices and the respiration and methane production. This is another example of how human activity indirectly affects biogeochemical cycles in a significant way. Although much of the debate about the future effects of increasing atmospheric carbon on climate change focuses on fossils fuels, scientists take natural processes, such as volcanoes and respiration, into account as they model and predict the future impact of this increase. The Nitrogen Cycle Getting nitrogen into the living world is difficult. Plants and phytoplankton are not equipped to incorporate nitrogen from the atmosphere (which exists as tightly bonded, triple covalent N~2~) even though this molecule comprises approximately 78 percent of the atmosphere. Nitrogen enters the living world via free-living and symbiotic bacteria, which incorporate nitrogen into their macromolecules through nitrogen fixation (conversion of N~2~). Cyanobacteria live in most aquatic ecosystems where sunlight is present; they play a key role in nitrogen fixation. Cyanobacteria are able to use inorganic sources of nitrogen to "fix" nitrogen. *Rhizobium* bacteria live symbiotically in the root nodules of legumes (such as peas, beans, and peanuts) and provide them with the organic nitrogen they need. Free-living bacteria, such as *Azotobacter*, are also important nitrogen fixers. Organic nitrogen is especially important to the study of ecosystem dynamics since many ecosystem processes, such as primary production and decomposition, are limited by the available supply of nitrogen. As shown in Figure 46.3.646.3.6, the nitrogen that enters living systems by nitrogen fixation is successively converted from organic nitrogen back into nitrogen gas by bacteria. This process occurs in three steps in terrestrial systems: ammonification, nitrification, and denitrification. First, the ammonification process converts nitrogenous waste from living animals or from the remains of dead animals into ammonium (NH~4~^+^) by certain bacteria and fungi. Second, the ammonium is converted to nitrites (NO~2~^−^) by nitrifying bacteria, such as *Nitrosomonas*, through nitrification. Subsequently, nitrites are converted to nitrates (NO~3~^−^) by similar organisms. Third, the process of denitrification occurs, whereby bacteria, such as *Pseudomonas* and *Clostridium*, convert the nitrates into nitrogen gas, allowing it to re-enter the atmosphere. Figure 3.3.6: Nitrogen enters the living world from the atmosphere via nitrogen-fixing bacteria. This nitrogen and nitrogenous waste from animals is then processed back into gaseous nitrogen by soil bacteria, which also supply terrestrial food webs with the organic nitrogen they need. (credit: modification of work by John M. Evans and Howard Perlman, USGS) Which of the following statements about the nitrogen cycle is false? 1. Ammonification converts organic nitrogenous matter from living organisms into ammonium (NH4+). 2. Denitrification by bacteria converts nitrates (NO3−) to nitrogen gas (N2). 3. Nitrification by bacteria converts nitrates (NO3−) to nitrites (NO2−). 4. Nitrogen fixing bacteria convert nitrogen gas (N2) into organic compounds. Human activity can release nitrogen into the environment by two primary means: the combustion of fossil fuels, which releases different nitrogen oxides, and by the use of artificial fertilizers in agriculture, which are then washed into lakes, streams, and rivers by surface runoff. Atmospheric nitrogen is associated with several effects on Earth's ecosystems including the production of acid rain (as nitric acid, HNO~3~) and greenhouse gas (as nitrous oxide, N~2~O) potentially causing climate change. A major effect from fertilizer runoff is saltwater and freshwater eutrophication, a process whereby nutrient runoff causes the excess growth of microorganisms, depleting dissolved oxygen levels and killing ecosystem fauna. A similar process occurs in the marine nitrogen cycle, where the ammonification, nitrification, and denitrification processes are performed by marine bacteria. Some of this nitrogen falls to the ocean floor as sediment, which can then be moved to land in geologic time by uplift of the Earth's surface and thereby incorporated into terrestrial rock. Although the movement of nitrogen from rock directly into living systems has been traditionally seen as insignificant compared with nitrogen fixed from the atmosphere, a recent study showed that this process may indeed be significant and should be included in any study of the global nitrogen cycle.^1^ The Phosphorus Cycle Phosphorus is an essential nutrient for living processes; it is a major component of nucleic acid and phospholipids, and, as calcium phosphate, makes up the supportive components of our bones. Phosphorus is often the limiting nutrient (necessary for growth) in aquatic ecosystems (Figure 3.3.7). Phosphorus occurs in nature as the phosphate ion (PO~4~^3−^). In addition to phosphate runoff as a result of human activity, natural surface runoff occurs when it is leached from phosphate-containing rock by weathering, thus sending phosphates into rivers, lakes, and the ocean. This rock has its origins in the ocean. Phosphate-containing ocean sediments form primarily from the bodies of ocean organisms and from their excretions. However, in remote regions, volcanic ash, aerosols, and mineral dust may also be significant phosphate sources. This sediment then is moved to land over geologic time by the uplifting of areas of the Earth's surface. ![](media/image16.jpeg)Phosphorus is also reciprocally exchanged between phosphate dissolved in the ocean and marine ecosystems. The movement of phosphate from the ocean to the land and through the soil is extremely slow, with the average phosphate ion having an oceanic residence time between 20,000 and 100,000 years. Figure 3.3.7: In nature, phosphorus exists as the phos[phate] ion (PO~4~^3−^). Weathering of rocks and volcanic activity releases phosphate into the soil, water, and air, where it becomes available to terrestrial food webs. Phosphate enters the oceans via surface runoff, groundwater flow, and river flow. Phosphate dissolved in ocean water cycles into marine food webs. Some phosphate from the marine food webs falls to the ocean floor, where it forms sediment. (credit: modification of work by John M. Evans and Howard Perlman, USGS) Excess phosphorus and nitrogen that enters these ecosystems from fertilizer runoff and from sewage causes excessive growth of microorganisms and depletes the dissolved oxygen, which leads to the death of many ecosystem fauna, such as shellfish and finfish. This process is responsible for dead zones in lakes and at the mouths of many major rivers (Figure 3.3.8). Figure 3.3.8: Dead zones occur when phosphorus and nitrogen from fertilizers cause excessive growth of microorganisms, which depletes oxygen and kills fauna. Worldwide, large dead zones are found in coastal areas of high population density. (credit: NASA Earth Observatory) A dead zone is an area within a freshwater or marine ecosystem where large areas are depleted of their normal flora and fauna; these zones can be caused by eutrophication, oil spills, dumping of toxic chemicals, and other human activities. The number of dead zones has been increasing for several years, and more than 400 of these zones were present as of 2008. One of the worst dead zones is off the coast of the United States in the Gulf of Mexico, where fertilizer runoff from the Mississippi River basin has created a dead zone of over 8463 square miles. Phosphate and nitrate runoff from fertilizers also negatively affect several lake and bay ecosystems including the Chesapeake Bay in the eastern United States. ![](media/image18.png)Chesapeake Bay Figure 3.3.9: This (a) satellite image shows the Chesapeake Bay, an ecosystem affected by phosphate and nitrate runoff. A (b) member of the Army Corps of Engineers holds a clump of oysters being used as a part of the oyster restoration effort in the bay. (credit a: modification of work by NASA/MODIS; credit b: modification of work by U.S. Army) The Chesapeake Bay has long been valued as one of the most scenic areas on Earth; it is now in distress and is recognized as a declining ecosystem. In the 1970s, the Chesapeake Bay was one of the first ecosystems to have identified dead zones, which continue to kill many fish and bottom-dwelling species, such as clams, oysters, and worms. Several species have declined in the Chesapeake Bay due to surface water runoff containing excess nutrients from artificial fertilizer used on land. The source of the fertilizers (with high nitrogen and phosphate content) is not limited to agricultural practices. There are many nearby urban areas and more than 150 rivers and streams empty into the bay that are carrying fertilizer runoff from lawns and gardens. Thus, the decline of the Chesapeake Bay is a complex issue and requires the cooperation of industry, agriculture, and everyday homeowners. Of particular interest to conservationists is the oyster population; it is estimated that more than 200,000 acres of oyster reefs existed in the bay in the 1700s, but that number has now declined to only 36,000 acres. Oyster harvesting was once a major industry for Chesapeake Bay, but it declined 88 percent between 1982 and 2007. This decline was due not only to fertilizer runoff and dead zones but also to overharvesting. Oysters require a certain minimum population density because they must be in close proximity to reproduce. Human activity has altered the oyster population and locations, greatly disrupting the ecosystem. The restoration of the oyster population in the Chesapeake Bay has been ongoing for several years with mixed success. Not only do many people find oysters good to eat, but they also clean up the bay. Oysters are filter feeders, and as they eat, they clean the water around them. In the 1700s, it was estimated that it took only a few days for the oyster population to filter the entire volume of the bay. Today, with changed water conditions, it is estimated that the present population would take nearly a year to do the same job. Restoration efforts have been ongoing for several years by non-profit organizations, such as the Chesapeake Bay Foundation. The restoration goal is to find a way to increase population density so the oysters can reproduce more efficiently. Many disease-resistant varieties (developed at the Virginia Institute of Marine Science for the College of William and Mary) are now available and have been used in the construction of experimental oyster reefs. Efforts to clean and restore the bay by Virginia and Delaware have been hampered because much of the pollution entering the bay comes from other states, which stresses the need for inter-state cooperation to gain successful restoration.The new, hearty oyster strains have also spawned a new and economically viable industry---oyster aquaculture---which not only supplies oysters for food and profit, but also has the added benefit of cleaning the bay. The Sulfur Cycle Sulfur is an essential element for the macromolecules of living things. As a part of the amino acid cysteine, it is involved in the formation of disulfide bonds within proteins, which help to determine their 3-D folding patterns, and hence their functions. As shown in Figure 46.3.1046.3.10, sulfur cycles between the oceans, land, and atmosphere. Atmospheric sulfur is found in the form of sulfur dioxide (SO~2~) and enters the atmosphere in three ways: from the decomposition of organic molecules, from volcanic activity and geothermal vents, and from the burning of fossil fuels by humans. Figure 3.3.10: Sulfur dioxide from the atmosphere becomes available to terrestrial and marine ecosystems when it is dissolved in precipitation as weak sulfuric acid or when it falls directly to the Earth as fallout. Weathering of rocks also makes sulfates available to terrestrial ecosystems. Decomposition of living organisms returns sulfates to the ocean, soil and atmosphere. (credit: modification of work by John M. Evans and Howard Perlman, USGS) On land, sulfur is deposited in four major ways: precipitation, direct fallout from the atmosphere, rock weathering, and geothermal vents. Atmospheric sulfur is found in the form of sulfur dioxide (SO~2~), and as rain falls through the atmosphere, sulfur is dissolved in the form of weak sulfuric acid (H~2~SO~4~). Sulfur can also fall directly from the atmosphere in a process called fallout. Also, the weathering of sulfur-containing rocks releases sulfur into the soil. These rocks originate from ocean sediments that are moved to land by the geologic uplifting of ocean sediments. Terrestrial ecosystems can then make use of these soil sulfates (SO−4SO4−), and upon the death and decomposition of these organisms, release the sulfur back into the atmosphere as hydrogen sulfide (H~2~S) gas. ![ This photo shows a white pyramid-shaped mound with gray steam escaping from it.](media/image20.jpeg) Figure 3.3.11: At this sulfur vent in Lassen Volcanic National Park in northeastern California, the yellowish sulfur deposits are visible near the mouth of the vent. Sulfur enters the ocean via runoff from land, from atmospheric fallout, and from underwater geothermal vents. Some ecosystems rely on chemoautotrophs using sulfur as a biological energy source. This sulfur then supports marine ecosystems in the form of sulfates. Human activities have played a major role in altering the balance of the global sulfur cycle. The burning of large quantities of fossil fuels, especially from coal, releases larger amounts of hydrogen sulfide gas into the atmosphere. As rain falls through this gas, it creates the phenomenon known as acid rain. Acid rain is corrosive rain caused by rainwater falling to the ground through sulfur dioxide gas, turning it into weak sulfuric acid, which causes damage to aquatic ecosystems. Acid rain damages the natural environment by lowering the pH of lakes, which kills many of the resident fauna; it also affects the man-made environment through the chemical degradation of buildings. For example, many marble monuments, such as the Lincoln Memorial in Washington, DC, have suffered significant damage from acid rain over the years. These examples show the wide-ranging effects of human activities on our environment and the challenges that remain for our future. Mineral nutrients are cycled through ecosystems and their environment. Of particular importance are water, carbon, nitrogen, phosphorus, and sulfur. All of these cycles have major impacts on ecosystem structure and function. As human activities have caused major disturbances to these cycles, their study and modeling is especially important. A variety of human activities, such as pollution, oil spills, and events) have damaged ecosystems, potentially causing global climate change. The health of Earth depends on understanding these cycles and how to protect the environment from irreversible damage. Enrichment: A. Multiple choice 1Which of the following statements about the nitrogen cycle is false? 1. Ammonification converts organic nitrogenous matter from living organisms into ammonium (NH~4~^+^). 2. Denitrification by bacteria converts nitrates (NO~3~^−^) to nitrogen gas (N~2~). 3. Nitrification by bacteria converts nitrates (NO~3~^−^) to nitrites (NO~2~^−^). 4. Nitrogen fixing bacteria convert nitrogen gas (N~2~) into organic compound The movement of mineral nutrients through organisms and their environment is called a \_\_\_\_\_\_\_\_ cycle. 1. biological 2. bioaccumulation 3. biogeochemical 4. biochemical Carbon is present in the atmosphere as \_\_\_\_\_\_\_\_. 1. carbon dioxide 2. carbonate ion 3. carbon dust 4. carbon monoxide The majority of water found on Earth is: 1. ice 2. water vapor 3. fresh water 4. salt water The average time a molecule spends in its reservoir is known as \_\_\_\_\_\_\_\_. 1. residence time 2. restriction time 3. resilience time 4. storage time The process whereby oxygen is depleted by the growth of microorganisms due to excess nutrients in aquatic systems is called \_\_\_\_\_\_\_\_. 1. dead zoning 2. eutrophication 3. retrofication 4. depletion The process whereby nitrogen is brought into organic molecules is called \_\_\_\_\_\_\_\_. 1. nitrification 2. denitrification 3. nitrogen fixation 4. nitrogen cycling B. 1..Describe nitrogen fixation and why it is important to agriculture. 2..What are the factors that cause dead zones? Describe eutrophication, in particular, as a cause. 3\. Why are drinking water supplies still a major concern for many countries? Lesson 3.4: Biomes Overview: The Earth's biomes are categorized into two major groups: terrestrial and aquatic. Terrestrial biomes are based on land, while aquatic biomes include both ocean and freshwater biomes. The eight major terrestrial biomes on Earth are each distinguished by characteristic temperatures and amount of precipitation. Comparing the annual totals of precipitation and fluctuations in precipitation from one biome to another provides clues as to the importance of abiotic factors in the distribution of biomes. Temperature variation on a daily and seasonal basis is also important for predicting the geographic distribution of the biome and the vegetation type in the biome: Objectives: At the end of the lesson the students should be able to: - Identify the two major abiotic factors that determine terrestrial biomes - Recognize distinguishing characteristics of each of the eight major terrestrial biomes - Describe the effects of abiotic factors on the composition of plant and animal communities in aquatic biomes - Compare and contrast the characteristics of the ocean zones - Summarize the characteristics of standing water and flowing water freshwater biomes Content:. The distribution of biomes shows that the same biome can occur in geographically distinct areas with similar climates (Figure 3.4.11). This world map shows the eight major biomes, polar ice, and mountains. Tropical forests, deserts and savannas are found primarily in South America, Africa, and Australia. Tropical forests also dominate Southeast Asia. Deserts dominate the Middle East and are found in the southwestern United States. Temperate forests dominate the eastern United States, Europe, and Eastern Asia. Temperate grasslands dominate the midwestern United States and parts of Asia, and are also found in South America. The boreal forest is found in northern Canada, Europe, and Asia, and tundra exists to the north of the boreal forest. Mountainous regions run the length of North and South America, and are found in northern India, Africa, and parts of Europe. Polar ice covers Greenland and Antarctica, which the latter is not shown on the map.Figure 3.4..1: Each of the world's major biomes is distinguished by characteristic temperatures and amounts of precipitation. Polar ice and mountains are also shown.. Tropical Wet Forest Tropical wet forests are also referred to as tropical rainforests. This biome is found in equatorial regions (\[link\]). The vegetation is characterized by plants with broad leaves that fall off throughout the year. Unlike the trees of deciduous forests, the trees in this biome do not have a seasonal loss of leaves associated with variations in temperature and sunlight; these forests are "evergreen" year-round. The temperature and sunlight profiles of tropical wet forests are very stable in comparison to that of other terrestrial biomes, with the temperatures ranging from 20 °C to 34 °C (68 °F to 93 °F). When one compares the annual temperature variation of tropical wet forests with that of other forest biomes, the lack of seasonal temperature variation in the tropical wet forest becomes apparent. This lack of seasonality leads to year-round plant growth, rather than the seasonal (spring, summer, and fall) growth seen in other biomes. In contrast to other ecosystems, tropical ecosystems do not have long days and short days during the yearly cycle. Instead, a constant daily amount of sunlight (11--12 hrs per day) provides more solar radiation, thereby, a longer period of time for plant growth. The annual rainfall in tropical wet forests ranges from 125 to 660 cm (50--200 in) with some monthly variation. While sunlight and temperature remain fairly consistent, annual rainfall is highly variable. Tropical wet forests have wet months in which there can be more than 30 cm (11--12 in) of precipitation, as well as dry months in which there are fewer than 10 cm (3.5 in) of rainfall. However, the driest month of a tropical wet forest still exceeds the *annual* rainfall of some other biomes, such as deserts. Tropical wet forests have high net primary productivity because the annual temperatures and precipitation values in these areas are ideal for plant growth. Therefore, the extensive biomass present in the tropical wet forest leads to plant communities with very high species diversities (Figure 3.4.2). Tropical wet forests have more species of trees than any other biome; on average between 100 and 300 species of trees are present in a single hectare (2.5 acres) of South America. One way to visualize this is to compare the distinctive horizontal layers within the tropical wet forest biome. On the forest floor is a sparse layer of plants and decaying plant matter. Above that is an understory of short shrubby foliage. A layer of trees rises above this understory and is topped by a closed upper canopy---the uppermost overhead layer of branches and leaves. Some additional trees emerge through this closed upper canopy. These layers provide diverse and complex habitats for the variety of plants, fungi, animals, and other organisms within the tropical wet forests. For instance, epiphytes are plants that grow on other plants, which typically are not harmed. Epiphytes are found throughout tropical wet forest biomes. Many species of animals use the variety of plants and the complex structure of the tropical wet forests for food and shelter. Some organisms live several meters above ground and have adapted to this arboreal lifestyle. ![ This photo depicts a section of the Amazon River, which is brown with mud. Trees line the edge of the river.](media/image22.jpeg)Figure 3.4.2: Tropical wet forests, such as these forests of Madre de Dios, Peru, near the Amazon River, have high species diversity. (credit: Roosevelt Garcia) Savannas Savannas are grasslands with scattered trees, and they are located in Africa, South America, and northern Australia (Figure 44.3.344.3.3). Savannas are hot, tropical areas with temperatures averaging from 24 °C to 29 °C (75 °F to 84 °F) and an annual rainfall of 10--40 cm (3.9--15.7 in). Savannas have an extensive dry season; for this reason, forest trees do not grow as well as they do in the tropical wet forest (or other forest biomes). As a result, within the grasses and forbs (herbaceous flowering plants) that dominate the savanna, there are relatively few trees (Figure 3.4.3). Since fire is an important source of disturbance in this biome, plants have evolved well-developed root systems that allow them to quickly re-sprout after a fire. A grassy slope plain is dotted with small trees.Figure 3.4..3: Savannas, like this one in Taita Hills Wildlife Sanctuary in Kenya, are dominated by grasses. (credit: Christopher T. Cooper) Subtropical Deserts Subtropical deserts exist between 15 ° and 30 ° north and south latitude and are centered on the Tropics of Cancer and Capricorn (Figure 44.3.444.3.4). This biome is very dry; in some years, evaporation exceeds precipitation. Subtropical hot deserts can have daytime soil surface temperatures above 60 °C (140 °F) and nighttime temperatures approaching 0 °C (32 °F). In cold deserts, temperatures can be as high as 25 °C and can drop below -30 °C (-22 °F). Subtropical deserts are characterized by low annual precipitation of fewer than 30 cm (12 in) with little monthly variation and lack of predictability in rainfall. In some cases, the annual rainfall can be as low as 2 cm (0.8 in) in subtropical deserts located in central Australia ("the Outback") and northern Africa. The vegetation and low animal diversity of this biome is closely related to this low and unpredictable precipitation. Very dry deserts lack perennial vegetation that lives from one year to the next; instead, many plants are annuals that grow quickly and reproduce when rainfall does occur, then they die. Many other plants in these areas are characterized by having a number of adaptations that conserve water, such as deep roots, reduced foliage, and water-storing stems (Figure 44.3.444.3.4). Seed plants in the desert produce seeds that can be in dormancy for extended periods between rains. Adaptations in desert animals include nocturnal behavior and burrowing. ![ This photo shows a sandy desert dotted with scrubby bushes. An ocotillo plant dominates the picture. It has long, thin unbranched stems that grow straight up from the base of the plant and radiate out slightly. The plant has no leaves.](media/image24.jpeg) Figure 3.4..4: To reduce water loss, many desert plants have tiny leaves or no leaves at all. The leaves of ocotillo (Fouquieria splendens), shown here in the Sonora Desert near Gila Bend, Arizona, appear only after rainfall, and then are shed. Chaparral The chaparral is also called the scrub forest and is found in California, along the Mediterranean Sea, and along the southern coast of Australia (Figure 44.3.544.3.5). The annual rainfall in this biome ranges from 65 cm to 75 cm (25.6--29.5 in), and the majority of the rain falls in the winter. Summers are very dry and many chaparral plants are dormant during the summertime. The chaparral vegetation, shown in Figure 3.4.5, is dominated by shrubs and is adapted to periodic fires, with some plants producing seeds that only germinate after a hot fire. The ashes left behind after a fire are rich in nutrients like nitrogen that fertilize the soil and promote plant regrowth. Photo depicts a landscape with many shrubs, dormant grass, a few trees, and mountains in the background. Figure 3.4..5: The chaparral is dominated by shrubs. (credit: Miguel Vieira) Temperate Grasslands Temperate grasslands are found throughout central North America, where they are also known as prairies; they are also in Eurasia, where they are known as steppes (Figure.3.4.6). Temperate grasslands have pronounced annual fluctuations in temperature with hot summers and cold winters. The annual temperature variation produces specific growing seasons for plants. Plant growth is possible when temperatures are warm enough to sustain plant growth and when ample water is available, which occurs in the spring, summer, and fall. During much of the winter, temperatures are low, and water, which is stored in the form of ice, is not available for plant growth. Annual precipitation ranges from 25 cm to 75 cm (9.8--29.5 in). Because of relatively lower annual precipitation in temperate grasslands, there are few trees except for those found growing along rivers or streams. The dominant vegetation tends to consist of grasses and some prairies sustain populations of grazing animals Figure .3.4.6. The vegetation is very dense and the soils are fertile because the subsurface of the soil is packed with the roots and rhizomes (underground stems) of these grasses. The roots and rhizomes act to anchor plants into the ground and replenish the organic material (humus) in the soil when they die and decay. ![ This photo shows a bison, which is dark brown in color with an even darker head. The hind part of the animal has short fur, and the front of the animal has longer, curly fur.](media/image26.jpeg) Figure 3.4..6: The American bison (Bison bison), more commonly called the buffalo, is a grazing mammal that once populated American prairies in huge numbers. (credit: Jack Dykinga, USDA Agricultural Research Service) Fires, mainly caused by lightning, are a natural disturbance in temperate grasslands. When fire is suppressed in temperate grasslands, the vegetation eventually converts to scrub and dense forests. Often, the restoration or management of temperate grasslands requires the use of controlled burns to suppress the growth of trees and maintain the grasses. Temperate Forests Temperate forests are the most common biome in eastern North America, Western Europe, Eastern Asia, Chile, and New Zealand (Figure 44.3.744.3.7). This biome is found throughout mid-latitude regions. Temperatures range between -30 °C and 30 °C (-22 °F to 86 °F) and drop to below freezing on an annual basis. These temperatures mean that temperate forests have defined growing seasons during the spring, summer, and early fall. Precipitation is relatively constant throughout the year and ranges between 75 cm and 150 cm (29.5--59 in). Because of the moderate annual rainfall and temperatures, deciduous trees are the dominant plant in this biome (Figure 3.4.7). Deciduous trees lose their leaves each fall and remain leafless in the winter. Thus, no photosynthesis occurs in the deciduous trees during the dormant winter period. Each spring, new leaves appear as the temperature increases. Because of the dormant period, the net primary productivity of temperate forests is less than that of tropical wet forests. In addition, temperate forests show less diversity of tree species than tropical wet forest biomes. Photo shows a deciduous forest with many tall trees, some smaller trees and grass, and lots of dead leaves on the forest floor. Sunlight filters down to the forest floor.Figure 3.4.7: Deciduous trees are the dominant plant in the temperate forest. (credit: Oliver Herold) The trees of the temperate forests leaf out and shade much of the ground; however, this biome is more open than tropical wet forests because trees in the temperate forests do not grow as tall as the trees in tropical wet forests. The soils of the temperate forests are rich in inorganic and organic nutrients. This is due to the thick layer of leaf litter on forest floors. As this leaf litter decays, nutrients are returned to the soil. The leaf litter also protects soil from erosion, insulates the ground, and provides habitats for invertebrates (such as the pill bug or roly-poly, *Armadillidium vulgare*) and their predators, such as the red-backed salamander (*Plethodon cinereus*). Boreal Forests The boreal forest, also known as taiga or coniferous forest, is found south of the Arctic Circle and across most of Canada, Alaska, Russia, and northern Europe (Figure 44.3.844.3.8). This biome has cold, dry winters and short, cool, wet summers. The annual precipitation is from 40 cm to 100 cm (15.7--39 in) and usually takes the form of snow. Little evaporation occurs because of the cold temperatures. The long and cold winters in the boreal forest have led to the predominance of cold-tolerant cone-bearing plants. These are evergreen coniferous trees like pines, spruce, and fir, which retain their needle-shaped leaves year-round. Evergreen trees can photosynthesize earlier in the spring than deciduous trees because less energy from the sun is required to warm a needle-like leaf than a broad leaf. This benefits evergreen trees, which grow faster than deciduous trees in the boreal forest. In addition, soils in boreal forest regions tend to be acidic with little available nitrogen. Leaves are a nitrogen-rich structure and deciduous trees must produce a new set of these nitrogen-rich structures each year. Therefore, coniferous trees that retain nitrogen-rich needles may have a competitive advantage over the broad-leafed deciduous trees. The net primary productivity of boreal forests is lower than that of temperate forests and tropical wet forests. The aboveground biomass of boreal forests is high because these slow-growing tree species are long lived and accumulate standing biomass over time. Plant species diversity is less than that seen in temperate forests and tropical wet forests. Boreal forests lack the pronounced elements of the layered forest structure seen in tropical wet forests. The structure of a boreal forest is often only a tree layer and a ground layer (Figure 3.4..8). When conifer needles are dropped, they decompose more slowly than broad leaves; therefore, fewer nutrients are returned to the soil to fuel plant growth. ![ The photo shows a boreal forest with a uniform low layer of plants and tall conifers scattered throughout the landscape. The snowcapped mountains of the Alaska Range are in the background.](media/image28.jpeg)Figure 3.4.8: The boreal forest (taiga) has low lying plants and conifer trees. (credit: L.B. Brubaker) Arctic Tundra The Arctic tundra lies north of the subarctic boreal forest and is located throughout the Arctic regions of the northern hemisphere (Figure 44.3.944.3.9). The average winter temperature is -34 °C (-34 °F) and the average summer temperature is from 3 °C to 12 °C (37 °F--52 °F). Plants in the arctic tundra have a very short growing season of approximately 10--12 weeks. However, during this time, there are almost 24 hours of daylight and plant growth is rapid. The annual precipitation of the Arctic tundra is very low with little annual variation in precipitation. And, as in the boreal forests, there is little evaporation due to the cold temperatures. Plants in the Arctic tundra are generally low to the ground (Figure 3.4..9). There is little species diversity, low net primary productivity, and low aboveground biomass. The soils of the Arctic tundra may remain in a perennially frozen state referred to as permafrost. The permafrost makes it impossible for roots to penetrate deep into the soil and slows the decay of organic matter, which inhibits the release of nutrients from organic matter. During the growing season, the ground of the Arctic tundra can be completely covered with plants or lichens. This photo shows a flat plain covered with shrub. Many of the shrubs are covered in pink flowers.Figure 3.4.9**:** Low-growing plants such as shrub willow dominate the tundra landscape, shown here in the Arctic National Wildlife Refuge. (credit: USFWS Arctic National Wildlife Refuge) **Abiotic Factors Influencing Aquatic Biomes** Like terrestrial biomes, aquatic biomes are influenced by a series of abiotic factors. The aquatic medium---water--- has different physical and chemical properties than air, however. Even if the water in a pond or other body of water is perfectly clear (there are no suspended particles), water, on its own, absorbs light. As one descends into a deep body of water, there will eventually be a depth which the sunlight cannot reach. While there are some abiotic and biotic factors in a terrestrial ecosystem that might obscure light (like fog, dust, or insect swarms), usually these are not permanent features of the environment. The importance of light in aquatic biomes is central to the communities of organisms found in both freshwater and marine ecosystems. In freshwater systems, stratification due to differences in density is perhaps the most critical abiotic factor and is related to the energy aspects of light. The thermal properties of water (rates of heating and cooling) are significant to the function of marine systems and have major impacts on global climate and weather patterns. Marine systems are also influenced by large-scale physical water movements, such as currents; these are less important in most freshwater lakes. The ocean is categorized by several areas or zones (Figure 44.4.144.4.1). All of the ocean's open water is referred to as the pelagic realm (or zone). The benthic realm (or zone) extends along the ocean bottom from the shoreline to the deepest parts of the ocean floor. Within the pelagic realm is the photic zone, which is the portion of the ocean that light can penetrate (approximately 200 m or 650 ft). At depths greater than 200 m, light cannot penetrate; thus, this is referred to as the aphotic zone. The majority of the ocean is aphotic and lacks sufficient light for photosynthesis. The deepest part of the ocean, the Challenger Deep (in the Mariana Trench, located in the western Pacific Ocean), is about 11,000 m (about 6.8 mi) deep. To give some perspective on the depth of this trench, the ocean is, on average, 4267 m or 14,000 ft deep. These realms and zones are relevant to freshwater lakes as well. ![ The illustration divides the ocean into different zones based on depth. The top layer, called the photic zone, extends from the surface to 200 m. The aphotic zone extends from 200 to 4,000 m. They abyssal zone extends from 4,000 m to the ocean bottom. The ocean is also divided into zones based on distance from the shore. The intertidal zone extends from high to low tide. The neritic zone extends from the intertidal zone to the point at which ocean depth is about 200 m. At about this depth, the continental shelf ends in a steep slope to the ocean bottom. The oceanic zone is the area of open ocean. A thin section of the oceanic zone extending from top to bottom and adjacent to the continental shelf is labeled the benthic realm. All of the ocean's open water is referred to as the pelagic realm, which is labeled on the left.](media/image30.png)Figure 3.4.10.: The ocean is divided into different zones based on water depth and distance from the shoreline. In which of the following regions would you expect to find photosynthetic organisms? 1. the aphotic zone, the neritic zone, the oceanic zone, and the benthic realm 2. the photic zone, the intertidal zone, the neritic zone, and the oceanic zone 3. the photic zone, the abyssal zone, the neritic zone, and the oceanic zone 4. the pelagic realm, the aphotic zone, the neritic zone, and the oceanic zone **Marine Biomes** The ocean is the largest marine biome. It is a continuous body of salt water that is relatively uniform in chemical composition; it is a weak solution of mineral salts and decayed biological matter. Within the ocean, coral reefs are a second kind of marine biome. Estuaries, coastal areas where salt water and fresh water mix, form a third unique marine biome. Ocean The physical diversity of the ocean is a significant influence on plants, animals, and other organisms. The ocean is categorized into different zones based on how far light reaches into the water. Each zone has a distinct group of species adapted to the biotic and abiotic conditions particular to that zone. The intertidal zone, which is the zone between high and low tide, is the oceanic region that is closest to land (Figure 44.4.144.4.1). Generally, most people think of this portion of the ocean as a sandy beach. In some cases, the intertidal zone is indeed a sandy beach, but it can also be rocky or muddy. The intertidal zone is an extremely variable environment because of tides. Organisms are exposed to air and sunlight at low tide and are underwater most of the time, especially during high tide. Therefore, living things that thrive in the intertidal zone are adapted to being dry for long periods of time. The shore of the intertidal zone is also repeatedly struck by waves, and the organisms found there are adapted to withstand damage from the pounding action of the waves (Figure 3.4.11). The exoskeletons of shoreline crustaceans (such as the shore crab, *Carcinus maenas*) are tough and protect them from desiccation (drying out) and wave damage. Another consequence of the pounding waves is that few algae and plants establish themselves in the constantly moving rocks, sand, or mud. Figure 3.4.11. Sea urchins, mussel shells, and starfish are often found in the intertidal zone, shown here in Kachemak Bay, Alaska. (credit: NOAA) The neritic zone (Figure 3.4.10) extends from the intertidal zone to depths of about 200 m (or 650 ft) at the edge of the continental shelf. Since light can penetrate this depth, photosynthesis can occur in the neritic zone. The water here contains silt and is well-oxygenated, low in pressure, and stable in temperature. Phytoplankton and floating *Sargassum* (a type of free-floating marine seaweed) provide a habitat for some sea life found in the neritic zone. Zooplankton, protists, small fishes, and shrimp are found in the neritic zone and are the base of the food chain for most of the world's fisheries. Beyond the neritic zone is the open ocean area known as the oceanic zone . Within the oceanic zone there is thermal stratification where warm and cold waters mix because of ocean currents. Abundant plankton serve as the base of the food chain for larger animals such as whales and dolphins. Nutrients are scarce and this is a relatively less productive part of the marine biome. When photosynthetic organisms and the protists and animals that feed on them die, their bodies fall to the bottom of the ocean where they remain; unlike freshwater lakes, the open ocean lacks a process for bringing the organic nutrients back up to the surface. The majority of organisms in the aphotic zone include sea cucumbers (phylum Echinodermata) and other organisms that survive on the nutrients contained in the dead bodies of organisms in the photic zone. Beneath the pelagic zone is the benthic realm, the deepwater region beyond the continental shelf. The bottom of the benthic realm is comprised of sand, silt, and dead organisms. Temperature decreases, remaining above freezing, as water depth increases. This is a nutrient-rich portion of the ocean because of the dead organisms that fall from the upper layers of the ocean. Because of this high level of nutrients, a diversity of fungi, sponges, sea anemones, marine worms, sea stars, fishes, and bacteria exist. The deepest part of the ocean is the abyssal zone, which is at depths of 4000 m or greater. The abyssal zone (Figure 3.4.10) is very cold and has very high pressure, high oxygen content, and low nutrient content. There are a variety of invertebrates and fishes found in this zone, but the abyssal zone does not have plants because of the lack of light. Hydrothermal vents are found primarily in the abyssal zone; chemosynthetic bacteria utilize the hydrogen sulfide and other minerals emitted from the vents. These chemosynthetic bacteria use the hydrogen sulfide as an energy source and serve as the base of the food chain found in the abyssal zone. Coral Reefs Coral reefs are ocean ridges formed by marine invertebrates living in warm shallow waters within the photic zone of the ocean. They are found within 30˚ north and south of the equator. The Great Barrier Reef is a well-known reef system located several miles off the northeastern coast of Australia. Other coral reef systems are fringing islands, which are directly adjacent to land, or atolls, which are circular reef systems surrounding a former landmass that is now underwater. The coral organisms (members of phylum Cnidaria) are colonies of saltwater polyps that secrete a calcium carbonate skeleton. These calcium-rich skeletons slowly accumulate, forming the underwater reef (Figure 44.4.344.4.3). Corals found in shallower waters (at a depth of approximately 60 m or about 200 ft) have a mutualistic relationship with photosynthetic unicellular algae. The relationship provides corals with the majority of the nutrition and the energy they require. The waters in which these corals live are nutritionally poor and, without this mutualism, it would not be possible for large corals to grow. Some corals living in deeper and colder water do not have a mutualistic relationship with algae; these corals attain energy and nutrients using stinging cells on their tentacles to capture prey. ![](media/image32.jpeg)It is estimated that more than 4,000 fish species inhabit coral reefs. These fishes can feed on coral, the cryptofauna (invertebrates found within the calcium carbonate substrate of the coral reefs), or the seaweed and algae that are associated with the coral. In addition, some fish species inhabit the boundaries of a coral reef; these species include predators, herbivores, or planktivores. Predators are animal species that hunt and are carnivores or "flesh eaters." Herbivores eat plant material, and planktivores eat plankton. Figure 3.4.12: Coral reefs are formed by the calcium carbonate skeletons of coral organisms, which are marine invertebrates in the phylum Cnidaria. (credit: Terry Hughes) It takes a long time to build a coral reef. The animals that create coral reefs have evolved over millions of years, continuing to slowly deposit the calcium carbonate that forms their characteristic ocean homes. Bathed in warm tropical waters, the coral animals and their symbiotic algal partners evolved to survive at the upper limit of ocean water temperature. Together, climate change and human activity pose dual threats to the long-term survival of the world's coral reefs. As global warming due to fossil fuel emissions raises ocean temperatures, coral reefs are suffering. The excessive warmth causes the reefs to expel their symbiotic, food-producing algae, resulting in a phenomenon known as bleaching. When bleaching occurs, the reefs lose much of their characteristic color as the algae and the coral animals die if loss of the symbiotic zooxanthellae is prolonged. Rising levels of atmospheric carbon dioxide further threaten the corals in other ways; as CO2 dissolves in ocean waters, it lowers the pH and increases ocean acidity. As acidity increases, it interferes with the calcification that normally occurs as coral animals build their calcium carbonate homes. When a coral reef begins to die, species diversity plummets as animals lose food and shelter. Coral reefs are also economically important tourist destinations, so the decline of coral reefs poses a serious threat to coastal economies. Human population growth has damaged corals in other ways, too. As human coastal populations increase, the runoff of sediment and agricultural chemicals has increased, too, causing some of the once-clear tropical waters to become cloudy. At the same time, overfishing of popular fish species has allowed the predator species that eat corals to go unchecked. Although a rise in global temperatures of 1--2˚C (a conservative scientific projection) in the coming decades may not seem large, it is very significant to this biome. When change occurs rapidly, species can become extinct before evolution leads to new adaptations. Many scientists believe that global warming, with its rapid (in terms of evolutionary time) and inexorable increases in temperature, is tipping the balance beyond the point at which many of the world's coral reefs can recover. Estuaries: Where the Ocean Meets Fresh Water Estuaries are biomes that occur where a source of fresh water, such as a river, meets the ocean. Therefore, both fresh water and salt water are found in the same vicinity; mixing results in a diluted (brackish) saltwater. Estuaries form protected areas where many of the young offspring of crustaceans, mollusks, and fish begin their lives. Salinity is a very important factor that influences the organisms and the adaptations of the organisms found in estuaries. The salinity of estuaries varies and is based on the rate of flow of its freshwater sources. Once or twice a day, high tides bring salt water into the estuary. Low tides occurring at the same frequency reverse the current of salt water. The short-term and rapid variation in salinity due to the mixing of fresh water and salt water is a difficult physiological challenge for the plants and animals that inhabit estuaries. Many estuarine plant species are halophytes: plants that can tolerate salty conditions. Halophytic plants are adapted to deal with the salinity resulting from saltwater on their roots or from sea spray. In some halophytes, filters in the roots remove the salt from the water that the plant absorbs. Other plants are able to pump oxygen into their roots. Animals, such as mussels and clams (phylum Mollusca), have developed behavioral adaptations that expend a lot of energy to function in this rapidly changing environment. When these animals are exposed to low salinity, they stop feeding, close their shells, and switch from aerobic respiration (in which they use gills) to anaerobic respiration (a process that does not require oxygen). When high tide returns to the estuary, the salinity and oxygen content of the water increases, and these animals open their shells, begin feeding, and return to aerobic respiration. **Freshwater Biomes** Freshwater biomes include lakes and ponds (standing water) as well as rivers and streams (flowing water). They also include wetlands, which will be discussed later. Humans rely on freshwater biomes to provide aquatic resources for drinking water, crop irrigation, sanitation, and industry. These various roles and human benefits are referred to as ecosystem services. Lakes and ponds are found in terrestrial landscapes and are, therefore, connected with abiotic and biotic factors influencing these terrestrial biomes. Lakes and Ponds Lakes and ponds can range in area from a few square meters to thousands of square kilometers. Temperature is an important abiotic factor affecting living things found in lakes and ponds. In the summer, thermal stratification of lakes and ponds occurs when the upper layer of water is warmed by the sun and does not mix with deeper, cooler water. Light can penetrate within the photic zone of the lake or pond. Phytoplankton (algae and cyanobacteria) are found here and carry out photosynthesis, providing the base of the food web of lakes and ponds. Zooplankton, such as rotifers and small crustaceans, consume these phytoplankton. At the bottom of lakes and ponds, bacteria in the aphotic zone break down dead organisms that sink to the bottom. Nitrogen and phosphorus are important limiting nutrients in lakes and ponds. Because of this, they are determining factors in the amount of phytoplankton growth in lakes and ponds. When there is a large input of nitrogen and phosphorus (from sewage and runoff from fertilized lawns and farms, for example), the growth of algae skyrockets, resulting in a large accumulation of algae called an algal bloom. Algal blooms can become so extensive that they reduce light penetration in water. As a result, the lake or pond becomes aphotic and photosynthetic plants cannot survive. When the algae die and decompose, severe oxygen depletion of the water occurs. Fishes and other organisms that require oxygen are then more likely to die, and resulting dead zones are found across the globe. Lake Erie and the Gulf of Mexico represent freshwater and marine habitats where phosphorus control and storm water runoff pose significant environmental challenges. Figure 3.4.13: The uncontrolled growth of algae in this lake has resulted in an algal bloom. (credit: Jeremy Nettleton) Rivers and Streams Rivers and streams are continuously moving bodies of water that carry large amounts of water from the source, or headwater, to a lake or ocean. The largest rivers include the Nile River in Africa, the Amazon River in South America, and the Mississippi River in North America. Abiotic features of rivers and streams vary along the length of the river or stream. Streams begin at a point of origin referred to as source water. The source

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