4.1: Energy Flow through Ecosystems - Geosciences
An ecosystem is a community of living organisms and their abiotic (non-living) environment. Ecosystems can be small, such as the tide pools found near the rocky shores of many oceans, or large, such as those found in the tropical rainforest of the Amazon in Brazil (Figure below).
There are three broad categories of ecosystems based on their general environment: freshwater, marine, and terrestrial. Within these three categories are individual ecosystem types based on the environmental habitat and organisms present.
Ecology of Ecosystems
Life in an ecosystem often involves competition for limited resources, which occurs both within a single species and between different species. Organisms compete for food, water, sunlight, space, and mineral nutrients. These resources provide the energy for metabolic processes and the matter to make up organisms' physical structures. Other critical factors influencing community dynamics are the components of its physical environment: a habitat's climate (seasons, sunlight, and rainfall), elevation, and geology. These can all be important environmental variables that determine which organisms can exist within a particular area.
Freshwater ecosystems are the least common, occurring on only 1.8 percent of Earth's surface. These systems comprise lakes, rivers, streams, and springs; they are quite diverse, and support a variety of animals, plants, fungi, protists and prokaryotes.
Marine ecosystems are the most common, comprising 75 percent of Earth's surface and consisting of three basic types: shallow ocean, deep ocean water, and deep ocean bottom. Shallow ocean ecosystems include extremely biodiverse coral reef ecosystems, yet the deep ocean water is known for 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 bottom ecosystems contain a wide variety of marine organisms. Such ecosystems exist even at depths where light is unable to penetrate through the water.
Terrestrial ecosystems, also known for their diversity, are grouped into large categories called biomes. A biome is a large-scale community of organisms, primarily defined on land by the dominant plant types that exist in geographic regions of the planet with similar climatic conditions. Examples of biomes include tropical rainforests, savannas, deserts, grasslands, temperate forests, and tundras. Grouping these ecosystems into just a few biome categories obscures the great diversity of the individual ecosystems within them. For example, the saguaro cacti (Carnegiea gigantean) and other plant life in the Sonoran Desert, in the United States, are relatively diverse compared with the desolate rocky desert of Boa Vista, an island off the coast of Western Africa (Figure below).
Ecosystems and Disturbance
Ecosystems are complex with many interacting parts. They are routinely exposed to various disturbances: changes in the environment that affect their compositions, such as yearly variations in rainfall and temperature. Many 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 with grasses, followed by bushes and shrubs, and later mature trees: thus, the forest is restored to its former state. This process is so universal that ecologists have given it a name—succession. The impact of environmental disturbances caused by human activities is now as significant as the changes wrought by natural processes. Human agricultural practices, air pollution, acid rain, global deforestation, overfishing, oil spills, and illegal dumping on land and into the ocean all have impacts on ecosystems.
Equilibrium is a dynamic state of an ecosystem in which, despite changes in species numbers and occurrence, biodiversity remains somewhat constant. 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 is called 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
A food chain is a linear sequence of organisms through which nutrients and energy pass as one organism eats another; the levels in the food chain are producers, primary consumers, higher-level consumers, and finally decomposers. These levels are used to describe ecosystem structure and dynamics. There is a single path through a food chain. Each organism in a food chain occupies a specific trophic level (energy level), its position in the food chain or food web.
In many ecosystems, the base, or foundation, of the food chain consists of photosynthetic organisms (plants or phytoplankton), which are called producers. The organisms that consume the 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 trophic 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 below, the Chinook salmon is the apex consumer at the top of this food chain.
One major factor that limits the number of steps in a food chain is energy. Energy is lost at each trophic level and between trophic levels as heat and in the transfer to decomposers (Figure below). 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.
There is a one problem when using food chains to describe most ecosystems. Even when all organisms are grouped into appropriate trophic levels, some of these organisms can feed on more than one trophic level; likewise, some of these organisms can also be fed on from multiple trophic levels. In addition, species feed on and are eaten by more than one species. In other words, the linear model of ecosystems, the food chain, is a hypothetical, overly simplistic representation of ecosystem structure. A holistic model—which includes 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 concept that accounts for the multiple trophic (feeding) interactions between each species and the many species it may feed on, or that feed on it. In a food web, the several trophic connections between each species and the other species that interact with it may cross multiple trophic levels. The matter and energy movements of virtually all ecosystems are more accurately described by food webs (Figure below).
Two general types of food webs are often shown interacting within a single ecosystem. A grazing food web 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), including decomposers (which break down dead and decaying organisms) and detritivores (which consume organic detritus). These organisms are usually bacteria, fungi, and invertebrate animals that recycle organic material back into the biotic part of the ecosystem as they themselves are consumed by other organisms. As ecosystems require a method to recycle material from dead organisms, 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 and fungi feeding off dead plants and animals. Simultaneously, a detrital food web can contribute energy to a grazing food web, as when a robin eats an earthworm.
How Organisms Acquire Energy in a Food Web
All living things require energy in one form or another. Energy is used by most complex metabolic pathways (usually in the form of ATP), especially those responsible for building large molecules from smaller compounds. Living organisms would not be able to assemble macromolecules (proteins, lipids, nucleic acids, and complex carbohydrates) from their monomers without a constant energy input.
Food-web diagrams illustrate how energy flows directionally through ecosystems. They can also indicate how efficiently organisms acquire energy, use it, and how much remains for use by other organisms of the food web. Energy is acquired by living things in two ways: autotrophs harness light or chemical energy and heterotrophs acquire energy through the consumption and digestion of other living or previously living organisms.
Photosynthetic and chemosynthetic organisms are autotrophs, which are 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, and chemosynthetic autotrophs (chemoautotrophs) use inorganic molecules as an energy source. Autotrophs are critical for most ecosystems: they are the producer trophic level. 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, are the energy source for a majority of the world's ecosystems. These ecosystems are often described by grazing and detrital food webs. Photoautotrophs harness the Sun's solar energy 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. The rate at which photosynthetic producers incorporate energy from the Sun is called gross primary productivity. However, not all of the energy incorporated by producers is available to the other organisms in the food web because producers must also grow and reproduce, which consumes energy. Net primary productivity is the energy that remains in the producers after accounting for these organisms' respiration and heat loss. The net productivity is then available to the primary consumers at the next trophic level.
Chemoautotrophs are primarily bacteria and archaea that are found in rare ecosystems where sunlight is not available, such as those associated with dark caves or hydrothermal vents at the bottom of the ocean (Figure below). Many chemoautotrophs in hydrothermal vents use hydrogen sulfide (H2S), which is released from the vents as a source of chemical energy; this allows them to synthesize complex organic molecules, such as glucose, for their own energy and, in turn, supplies energy to the rest of the ecosystem.
Consequences of Food Webs: Biological Magnification
One of the most important consequences of ecosystem dynamics in terms of human impact is biomagnification. Biomagnification is the increasing concentration of persistent, toxic substances in organisms at each successive trophic level. These are substances that are fat soluble, not water soluble, and are stored in the fat reserves of each organism. Many substances have been shown to biomagnify, including classical studies with the pesticide dichlorodiphenyltrichloroethane (DDT), which were described in the 1960s bestseller, Silent Spring by Rachel Carson. DDT was a commonly used pesticide before its dangers to apex consumers, such as the bald eagle, became known. In aquatic ecosystems, organisms from each trophic level consumed many organisms in 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 devastating 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 (PCB), which were used as coolant liquids in the United States until their use was banned in 1979, and heavy metals, such as mercury, lead, and cadmium. These substances are best studied in aquatic ecosystems, where predatory fish species accumulate very high concentrations of toxic substances that are at quite low concentrations in the environment and in producers. As illustrated in a study performed by the NOAA in the Saginaw Bay of Lake Huron of the North American Great Lakes (Figure below), PCB concentrations increased from the producers of the ecosystem (phytoplankton) through the different trophic levels of fish species. The apex consumer, the 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.
Other concerns have been raised by the biomagnification of heavy metals, such as mercury and cadmium, in certain types of seafood. The United States Environmental Protection Agency 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, shrimp, pollock, and catfish. Biomagnification is a good example of how ecosystem dynamics can affect our everyday lives, even influencing the food we eat.
Groundwater is a critical resource for human life, as well as for the maintenance of ecosystems. Understanding contaminated groundwater with human health is an emerging area of interest and study. Groundwater contributes to the sustainability of ecosystems for harmonizing human beings with nature. On the other hand, environmental changes affect surface and ground water resources more and more. This Special Issue will be published in Geosciences and provide topics regarding groundwater and human health, geohydrology in connection with water and life, geomicrobial characterization of subsurface environments, groundwater resource management, environmental drivers changing hydrologic system and consequences, groundwater quantity and quality issues related to environmental changes, groundwater&ndashsurface water interaction, biogeochemistry and ecohydraulics of the riparian and hyporheic zone, groundwater-dependent ecosystems, and so on. The Guest Editors are encouraging papers that contains novel approaches and technologies on the abovementioned topics.
Prof. Se-Yeong Hamm
Prof. Tianming Huang
Prof. Min-Ho Koo
Prof. Jung-Hwi Kihm
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The Multiple Levels Of Energy Flow
The flow of energy through our ecosystem happens through multiple levels. The first trophic level is made up of producers that use solar energy and the process of photosynthesis to create organic material. Those producers are plants, and they play a part in the second trophic level. What happens in the second trophic level is that the herbivores use those plants as food, which then provides them with energy. This energy is mostly used for maintaining essential bodily functions. Those functions include food digestion, breathing, growth of tissues, and body temperature, and blood circulation maintenance.
The flow of energy through our ecosystem happens through multiple levels.
The next trophic level of the flow of energy through an ecosystem includes carnivores. They feed on the herbivores from the previous level and use them for energy that maintains their growth and sustenance. It doesn’t stop there, because smaller carnivores can be eaten by larger predators, which makes this level more complicated in some cases. But the important thing to note is how different animal species and plants are linked by being a part of the food chain. Once the animal dies, it is decomposed by bacteria, fungi, and different insects, thereby returning the nutrients to the ground. Those nutrients are taken up by the plants again. The energy gets released during the process of decomposition it cannot be recycled.
4.1 Energy and Metabolism
Scientists use the term bioenergetics to describe the concept of energy flow (Figure 4.2) through living systems, such as cells. Cellular processes such as the building and breaking down of complex molecules occur through stepwise chemical reactions. Some of these chemical reactions are spontaneous and release energy, whereas others require energy to proceed. Just as living things must continually consume food to replenish their energy supplies, cells must continually obtain more energy to replenish that used by the many energy-requiring chemical reactions that constantly take place. Together, all of the chemical reactions that take place inside cells, including those that consume or generate energy, are referred to as the cell’s metabolism .
Consider the metabolism of sugar. This is a classic example of one of the many cellular processes that use and produce energy. Living things consume sugars as a major energy source, because sugar molecules have a great deal of energy stored within their bonds. For the most part, photosynthesizing organisms like plants produce these sugars. During photosynthesis, plants use energy (originally from sunlight) to convert carbon dioxide gas (CO2) into sugar molecules (like glucose: C6H12O6). They consume carbon dioxide and produce oxygen as a waste product. This reaction is summarized as:
Because this process involves synthesizing an energy-storing molecule, it requires energy input to proceed. During the light reactions of photosynthesis, energy is provided by a molecule called adenosine triphosphate (ATP), which is the primary energy currency of all cells. Just as the dollar is used as currency to buy goods, cells use molecules of ATP as energy currency to perform immediate work. In contrast, energy-storage molecules such as glucose are consumed only to be broken down to use their energy. The reaction that harvests the energy of a sugar molecule in cells requiring oxygen to survive can be summarized by the reverse reaction to photosynthesis. In this reaction, oxygen is consumed and carbon dioxide is released as a waste product. The reaction is summarized as:
Both of these reactions involve many steps.
The processes of making and breaking down sugar molecules illustrate two examples of metabolic pathways. A metabolic pathway is a series of chemical reactions that takes a starting molecule and modifies it, step-by-step, through a series of metabolic intermediates, eventually yielding a final product. In the example of sugar metabolism, the first metabolic pathway synthesized sugar from smaller molecules, and the other pathway broke sugar down into smaller molecules. These two opposite processes—the first requiring energy and the second producing energy—are referred to as anabolic pathways (building polymers) and catabolic pathways (breaking down polymers into their monomers), respectively. Consequently, metabolism is composed of synthesis (anabolism) and degradation (catabolism) (Figure 4.3).
It is important to know that the chemical reactions of metabolic pathways do not take place on their own. Each reaction step is facilitated, or catalyzed, by a protein called an enzyme. Enzymes are important for catalyzing all types of biological reactions—those that require energy as well as those that release energy.
Thermodynamics refers to the study of energy and energy transfer involving physical matter. The matter relevant to a particular case of energy transfer is called a system, and everything outside of that matter is called the surroundings. For instance, when heating a pot of water on the stove, the system includes the stove, the pot, and the water. Energy is transferred within the system (between the stove, pot, and water). There are two types of systems: open and closed. In an open system, energy can be exchanged with its surroundings. The stovetop system is open because heat can be lost to the air. A closed system cannot exchange energy with its surroundings.
Biological organisms are open systems. Energy is exchanged between them and their surroundings as they use energy from the sun to perform photosynthesis or consume energy-storing molecules and release energy to the environment by doing work and releasing heat. Like all things in the physical world, energy is subject to physical laws. The laws of thermodynamics govern the transfer of energy in and among all systems in the universe.
In general, energy is defined as the ability to do work, or to create some kind of change. Energy exists in different forms. For example, electrical energy, light energy, and heat energy are all different types of energy. To appreciate the way energy flows into and out of biological systems, it is important to understand two of the physical laws that govern energy.
The first law of thermodynamics states that the total amount of energy in the universe is constant and conserved. In other words, there has always been, and always will be, exactly the same amount of energy in the universe. Energy exists in many different forms. According to the first law of thermodynamics, energy may be transferred from place to place or transformed into different forms, but it cannot be created or destroyed. The transfers and transformations of energy take place around us all the time. Light bulbs transform electrical energy into light and heat energy. Gas stoves transform chemical energy from natural gas into heat energy. Plants perform one of the most biologically useful energy transformations on earth: that of converting the energy of sunlight to chemical energy stored within organic molecules (Figure 4.2). Some examples of energy transformations are shown in Figure 4.4.
The challenge for all living organisms is to obtain energy from their surroundings in forms that they can transfer or transform into usable energy to do work. Living cells have evolved to meet this challenge. Chemical energy stored within organic molecules such as sugars and fats is transferred and transformed through a series of cellular chemical reactions into energy within molecules of ATP. Energy in ATP molecules is easily accessible to do work. Examples of the types of work that cells need to do include building complex molecules, transporting materials, powering the motion of cilia or flagella, and contracting muscle fibers to create movement.
A living cell’s primary tasks of obtaining, transforming, and using energy to do work may seem simple. However, the second law of thermodynamics explains why these tasks are harder than they appear. All energy transfers and transformations are never completely efficient. In every energy transfer, some amount of energy is lost in a form that is unusable. In most cases, this form is heat energy. Thermodynamically, heat energy is defined as the energy transferred from one system to another that is not work. For example, when a light bulb is turned on, some of the energy being converted from electrical energy into light energy is lost as heat energy. Likewise, some energy is lost as heat energy during cellular metabolic reactions.
An important concept in physical systems is that of order and disorder. The more energy that is lost by a system to its surroundings, the less ordered and more random the system is. Scientists refer to the measure of randomness or disorder within a system as entropy. High entropy means high disorder and low energy. Molecules and chemical reactions have varying entropy as well. For example, entropy increases as molecules at a high concentration in one place diffuse and spread out. The second law of thermodynamics says that energy will always be lost as heat in energy transfers or transformations.
Living things are highly ordered, requiring constant energy input to be maintained in a state of low entropy.
Potential and Kinetic Energy
When an object is in motion, there is energy associated with that object. Think of a wrecking ball. Even a slow-moving wrecking ball can do a great deal of damage to other objects. Energy associated with objects in motion is called kinetic energy (Figure 4.5). A speeding bullet, a walking person, and the rapid movement of molecules in the air (which produces heat) all have kinetic energy.
Now what if that same motionless wrecking ball is lifted two stories above ground with a crane? If the suspended wrecking ball is unmoving, is there energy associated with it? The answer is yes. The energy that was required to lift the wrecking ball did not disappear, but is now stored in the wrecking ball by virtue of its position and the force of gravity acting on it. This type of energy is called potential energy (Figure 4.5). If the ball were to fall, the potential energy would be transformed into kinetic energy until all of the potential energy was exhausted when the ball rested on the ground. Wrecking balls also swing like a pendulum through the swing, there is a constant change of potential energy (highest at the top of the swing) to kinetic energy (highest at the bottom of the swing). Other examples of potential energy include the energy of water held behind a dam or a person about to skydive out of an airplane.
Potential energy is not only associated with the location of matter, but also with the structure of matter. Even a spring on the ground has potential energy if it is compressed so does a rubber band that is pulled taut. On a molecular level, the bonds that hold the atoms of molecules together exist in a particular structure that has potential energy. Remember that anabolic cellular pathways require energy to synthesize complex molecules from simpler ones and catabolic pathways release energy when complex molecules are broken down. The fact that energy can be released by the breakdown of certain chemical bonds implies that those bonds have potential energy. In fact, there is potential energy stored within the bonds of all the food molecules we eat, which is eventually harnessed for use. This is because these bonds can release energy when broken. The type of potential energy that exists within chemical bonds, and is released when those bonds are broken, is called chemical energy. Chemical energy is responsible for providing living cells with energy from food. The release of energy occurs when the molecular bonds within food molecules are broken.
Concepts in Action
Visit the site and select “Pendulum” from the “Work and Energy” menu to see the shifting kinetic and potential energy of a pendulum in motion.
Free and Activation Energy
After learning that chemical reactions release energy when energy-storing bonds are broken, an important next question is the following: How is the energy associated with these chemical reactions quantified and expressed? How can the energy released from one reaction be compared to that of another reaction? A measurement of free energy is used to quantify these energy transfers. Recall that according to the second law of thermodynamics, all energy transfers involve the loss of some amount of energy in an unusable form such as heat. Free energy specifically refers to the energy associated with a chemical reaction that is available after the losses are accounted for. In other words, free energy is usable energy, or energy that is available to do work.
If energy is released during a chemical reaction, then the change in free energy, signified as ∆G (delta G) will be a negative number. A negative change in free energy also means that the products of the reaction have less free energy than the reactants, because they release some free energy during the reaction. Reactions that have a negative change in free energy and consequently release free energy are called exergonic reactions . Think: exergonic means energy is exiting the system. These reactions are also referred to as spontaneous reactions, and their products have less stored energy than the reactants. An important distinction must be drawn between the term spontaneous and the idea of a chemical reaction occurring immediately. Contrary to the everyday use of the term, a spontaneous reaction is not one that suddenly or quickly occurs. The rusting of iron is an example of a spontaneous reaction that occurs slowly, little by little, over time.
If a chemical reaction absorbs energy rather than releases energy on balance, then the ∆G for that reaction will be a positive value. In this case, the products have more free energy than the reactants. Thus, the products of these reactions can be thought of as energy-storing molecules. These chemical reactions are called endergonic reactions and they are non-spontaneous. An endergonic reaction will not take place on its own without the addition of free energy.
Look at each of the processes shown and decide if it is endergonic or exergonic.
There is another important concept that must be considered regarding endergonic and exergonic reactions. Exergonic reactions require a small amount of energy input to get going, before they can proceed with their energy-releasing steps. These reactions have a net release of energy, but still require some energy input in the beginning. This small amount of energy input necessary for all chemical reactions to occur is called the activation energy .
Concepts in Action
Watch an animation of the move from free energy to transition state of the reaction.
A substance that helps a chemical reaction to occur is called a catalyst, and the molecules that catalyze biochemical reactions are called enzymes . Most enzymes are proteins and perform the critical task of lowering the activation energies of chemical reactions inside the cell. Most of the reactions critical to a living cell happen too slowly at normal temperatures to be of any use to the cell. Without enzymes to speed up these reactions, life could not persist. Enzymes do this by binding to the reactant molecules and holding them in such a way as to make the chemical bond-breaking and -forming processes take place more easily. It is important to remember that enzymes do not change whether a reaction is exergonic (spontaneous) or endergonic. This is because they do not change the free energy of the reactants or products. They only reduce the activation energy required for the reaction to go forward (Figure 4.7). In addition, an enzyme itself is unchanged by the reaction it catalyzes. Once one reaction has been catalyzed, the enzyme is able to participate in other reactions.
The chemical reactants to which an enzyme binds are called the enzyme’s substrates . There may be one or more substrates, depending on the particular chemical reaction. In some reactions, a single reactant substrate is broken down into multiple products. In others, two substrates may come together to create one larger molecule. Two reactants might also enter a reaction and both become modified, but they leave the reaction as two products. The location within the enzyme where the substrate binds is called the enzyme’s active site . The active site is where the “action” happens. Since enzymes are proteins, there is a unique combination of amino acid side chains within the active site. Each side chain is characterized by different properties. They can be large or small, weakly acidic or basic, hydrophilic or hydrophobic, positively or negatively charged, or neutral. The unique combination of side chains creates a very specific chemical environment within the active site. This specific environment is suited to bind to one specific chemical substrate (or substrates).
Active sites are subject to influences of the local environment. Increasing the environmental temperature generally increases reaction rates, enzyme-catalyzed or otherwise. However, temperatures outside of an optimal range reduce the rate at which an enzyme catalyzes a reaction. Hot temperatures will eventually cause enzymes to denature, an irreversible change in the three-dimensional shape and therefore the function of the enzyme. Enzymes are also suited to function best within a certain pH and salt concentration range, and, as with temperature, extreme pH, and salt concentrations can cause enzymes to denature.
For many years, scientists thought that enzyme-substrate binding took place in a simple “lock and key” fashion. This model asserted that the enzyme and substrate fit together perfectly in one instantaneous step. However, current research supports a model called induced fit (Figure 4.8). The induced-fit model expands on the lock-and-key model by describing a more dynamic binding between enzyme and substrate. As the enzyme and substrate come together, their interaction causes a mild shift in the enzyme’s structure that forms an ideal binding arrangement between enzyme and substrate.
Concepts in Action
When an enzyme binds its substrate, an enzyme-substrate complex is formed. This complex lowers the activation energy of the reaction and promotes its rapid progression in one of multiple possible ways. On a basic level, enzymes promote chemical reactions that involve more than one substrate by bringing the substrates together in an optimal orientation for reaction. Another way in which enzymes promote the reaction of their substrates is by creating an optimal environment within the active site for the reaction to occur. The chemical properties that emerge from the particular arrangement of amino acid R groups within an active site create the perfect environment for an enzyme’s specific substrates to react.
The enzyme-substrate complex can also lower activation energy by compromising the bond structure so that it is easier to break. Finally, enzymes can also lower activation energies by taking part in the chemical reaction itself. In these cases, it is important to remember that the enzyme will always return to its original state by the completion of the reaction. One of the hallmark properties of enzymes is that they remain ultimately unchanged by the reactions they catalyze. After an enzyme has catalyzed a reaction, it releases its product(s) and can catalyze a new reaction.
It would seem ideal to have a scenario in which all of an organism's enzymes existed in abundant supply and functioned optimally under all cellular conditions, in all cells, at all times. However, a variety of mechanisms ensures that this does not happen. Cellular needs and conditions constantly vary from cell to cell, and change within individual cells over time. The required enzymes of stomach cells differ from those of fat storage cells, skin cells, blood cells, and nerve cells. Furthermore, a digestive organ cell works much harder to process and break down nutrients during the time that closely follows a meal compared with many hours after a meal. As these cellular demands and conditions vary, so must the amounts and functionality of different enzymes.
Since the rates of biochemical reactions are controlled by activation energy, and enzymes lower and determine activation energies for chemical reactions, the relative amounts and functioning of the variety of enzymes within a cell ultimately determine which reactions will proceed and at what rates. This determination is tightly controlled in cells. In certain cellular environments, enzyme activity is partly controlled by environmental factors like pH, temperature, salt concentration, and, in some cases, cofactors or coenzymes.
Enzymes can also be regulated in ways that either promote or reduce enzyme activity. There are many kinds of molecules that inhibit or promote enzyme function, and various mechanisms by which they do so. In some cases of enzyme inhibition, an inhibitor molecule is similar enough to a substrate that it can bind to the active site and simply block the substrate from binding. When this happens, the enzyme is inhibited through competitive inhibition , because an inhibitor molecule competes with the substrate for binding to the active site.
On the other hand, in noncompetitive inhibition , an inhibitor molecule binds to the enzyme in a location other than the active site, called an allosteric site, but still manages to prevent substrate binding to the active site. Some inhibitor molecules bind to enzymes in a location where their binding induces a conformational change that reduces the enzyme activity as it no longer effectively catalyzes the conversion of the substrate to product. This type of inhibition is called allosteric inhibition (Figure 4.9). Most allosterically regulated enzymes are made up of more than one polypeptide, meaning that they have more than one protein subunit. When an allosteric inhibitor binds to a region on an enzyme, all active sites on the protein subunits are changed slightly such that they bind their substrates with less efficiency. There are allosteric activators as well as inhibitors. Allosteric activators bind to locations on an enzyme away from the active site, inducing a conformational change that increases the affinity of the enzyme’s active site(s) for its substrate(s) (Figure 4.9).
Pharmaceutical Drug Developer
Enzymes are key components of metabolic pathways. Understanding how enzymes work and how they can be regulated are key principles behind the development of many of the pharmaceutical drugs on the market today. Biologists working in this field collaborate with other scientists to design drugs (Figure 4.10).
Consider statins for example—statins is the name given to one class of drugs that can reduce cholesterol levels. These compounds are inhibitors of the enzyme HMG-CoA reductase, which is the enzyme that synthesizes cholesterol from lipids in the body. By inhibiting this enzyme, the level of cholesterol synthesized in the body can be reduced. Similarly, acetaminophen, popularly marketed under the brand name Tylenol, is an inhibitor of the enzyme cyclooxygenase. While it is used to provide relief from fever and inflammation (pain), its mechanism of action is still not completely understood.
How are drugs discovered? One of the biggest challenges in drug discovery is identifying a drug target. A drug target is a molecule that is literally the target of the drug. In the case of statins, HMG-CoA reductase is the drug target. Drug targets are identified through painstaking research in the laboratory. Identifying the target alone is not enough scientists also need to know how the target acts inside the cell and which reactions go awry in the case of disease. Once the target and the pathway are identified, then the actual process of drug design begins. In this stage, chemists and biologists work together to design and synthesize molecules that can block or activate a particular reaction. However, this is only the beginning: If and when a drug prototype is successful in performing its function, then it is subjected to many tests from in vitro experiments to clinical trials before it can get approval from the U.S. Food and Drug Administration to be on the market.
Many enzymes do not work optimally, or even at all, unless bound to other specific non-protein helper molecules. They may bond either temporarily through ionic or hydrogen bonds, or permanently through stronger covalent bonds. Binding to these molecules promotes optimal shape and function of their respective enzymes. Two examples of these types of helper molecules are cofactors and coenzymes. Cofactors are inorganic ions such as ions of iron and magnesium. Coenzymes are organic helper molecules, those with a basic atomic structure made up of carbon and hydrogen. Like enzymes, these molecules participate in reactions without being changed themselves and are ultimately recycled and reused. Vitamins are the source of coenzymes. Some vitamins are the precursors of coenzymes and others act directly as coenzymes. Vitamin C is a direct coenzyme for multiple enzymes that take part in building the important connective tissue, collagen. Therefore, enzyme function is, in part, regulated by the abundance of various cofactors and coenzymes, which may be supplied by an organism’s diet or, in some cases, produced by the organism.
Feedback Inhibition in Metabolic Pathways
Molecules can regulate enzyme function in many ways. The major question remains, however: What are these molecules and where do they come from? Some are cofactors and coenzymes, as you have learned. What other molecules in the cell provide enzymatic regulation such as allosteric modulation, and competitive and non-competitive inhibition? Perhaps the most relevant sources of regulatory molecules, with respect to enzymatic cellular metabolism, are the products of the cellular metabolic reactions themselves. In a most efficient and elegant way, cells have evolved to use the products of their own reactions for feedback inhibition of enzyme activity. Feedback inhibition involves the use of a reaction product to regulate its own further production (Figure 4.11). The cell responds to an abundance of the products by slowing down production during anabolic or catabolic reactions. Such reaction products may inhibit the enzymes that catalyzed their production through the mechanisms described above.
The production of both amino acids and nucleotides is controlled through feedback inhibition. Additionally, ATP is an allosteric regulator of some of the enzymes involved in the catabolic breakdown of sugar, the process that creates ATP. In this way, when ATP is in abundant supply, the cell can prevent the production of ATP. On the other hand, ADP serves as a positive allosteric regulator (an allosteric activator) for some of the same enzymes that are inhibited by ATP. Thus, when relative levels of ADP are high compared to ATP, the cell is triggered to produce more ATP through sugar catabolism.
4.1: Energy Flow through Ecosystems - Geosciences
An ecosystem consists of a community of living organisms interacting with each other and the environment. The source of energy that fuels most ecosystems is the Sun. Plants use the Sun's energy to produce food in a process called photosynthesis. Organisms that use energy from the Sun or energy stored in chemical compounds to produce their own nutrients are called autotrophs. They are also called producers because most other organisms depend on autotrophs for nutrients and energy. Heterotrophic organisms that cannot make their own food may obtain nutrients by eating other organisms. A heterotroph that feeds only on plants is called an herbivore. Herbivores are also called first order heterotrophs. Carnivores that feed on herbivores are called second order heterotrophs. Carnivores that feed on other carnivores are called third order heterotrophs. A food chain is a simple model of how energy and matter move through an ecosystem.
Each level of production and consumption in a food chain is a trophic level. The autotrophs form the first trophic level, first order heterotrophs (herbivores) constitute the second trophic level, second order heterotrophs the third trophic level, and third order heterotrophs are layered on top.
In the pyramid of energy, the energy moves in only one direction and decreases at each succeeding trophic level. The total energy transfer from one trophic level to the next is, in general, only about ten percent or less. This is called the energy conversion efficiency. Organisms fail to capture and eat all the food available at the trophic level below them. The food consumers ingest is used to metabolize and build body tissues. Some food is given off as waste. The energy lost at each trophic level enters the environment as heat.
A pyramid of biomass expresses the weight of living material at each trophic level. Biomass is calculated by finding the average weight of each species at that trophic level and multiplying the weight by the estimated number of organisms in each population. In terrestrial ecosystems, biomass decreases as the trophic level increases. In contrast to terrestrial ecosystems, freshwater and marine ecosystems have less primary producer biomass than biomass present at higher trophic levels, leading to an inverted biomass pyramid. This is because algae and phytoplankton have a shorter lifespan, are more edible than terrestrial plants, and are more rapidly grazed. Their biomass does not accumulate.
In this exploration, you will study and analyze five simplified model ecosystems: a deciduous forest, a hot desert, a freshwater lake, grassland, and an Antarctic ocean shore. Many more plant and animal species would be involved in a real-world ecosystem. The field notes for each model ecosystem present a profile of the plant and animal inventory for each ecosystem.
1. Select the icon next to the animals' and plants' names from the Field Notes tablet and drag and drop them to the appropriate trophic level in the ecosystem pyramid.
2. Click the Check button when all the names have been placed in the ecosystem pyramid to verify the accuracy of animal and plant assignments to the appropriate trophic level. For each accurate placement, the names of the animals are replaced with pictures and the number of each kind of animal is displayed beneath it.
3. Click the Pyramid of Energy button to display numbers indicating amounts of energy.
4. Click the Pyramid of Numbers button to display numbers indicating numbers of plants and animals.
5. Analyze this data by calculating the conversion efficiency for each trophic level for each of the five ecosystems and record the results in the Data Table. The energy conversion efficiency is calculated by dividing the energy at the higher trophic level by the energy at the lower level to obtain a ratio. Enter the value as a decimal number.
6. When all the data are recorded in the Data Table and analyzed, answer the Journal Questions.
4.1 Ecosystems and energy flow
When certain chemicals enters the food chains, they may accumulate in the fatty tissues or bones and increase in amount for every step in the food chain. Persistent Organic Pollutants (POPs) are regarded as particularly dangerous accumulating chemicals.
Ecosystems can be disrupted
Eosystems may change suddenly or over time. If the change happens over a long period of time, most species can adapt to the new conditions.
Puffin (Alca arctica), Noss, Shetland Islands. Vulnerable species due to changes in the environment and overfishing. Photo: P. Prokosch
Ecosystems may be disrupted through natural causes, such as a natural change in climate, or natural disasters like earthquakes, volcanic eruptions and tsunamies. Human activities may also disrupt ecosystems, locally as well as globally. Human activities may enhance existing natural processes or cycles, such as the greenhouse effect, or create completely new conditions, such as depleting the ozone layer with ozone depleting chemicals, or add POPs to the energy flow in the food chains.
Pollution through high concentrations of waste and secreta may affect water bodies and cause eutrophication.
Ecosystem services are the benefits people obtain from ecosystems. These include provisioning services such as food, water, timber, and fiber regulating services that affect climate, floods, disease, wastes, and water quality cultural services that provide recreational, aesthetic, and spiritual benefits and supporting services such as soil formation, photosynthesis, and nutrient cycling
Millennium Ecosystem Assessment – Synthesis
Ecosystems exist underground, on land, at sea, and in the air. Organisms in an ecosystem acquire energy in a variety of ways, which is transferred between trophic levels as the energy flows from the base to the top of the food web, with energy being lost at each transfer. There is energy lost at each trophic level, so the lengths of food chains are limited because there is a point where not enough energy remains to support a population of consumers. Fat soluble compounds biomagnify up a food chain causing damage to top consumers. even when environmental concentrations of a toxin are low.