"

3 Bioenergetics and Metabolism

Image showing a variety of foods representing the three primary macronutrients: carbohydrates (e.g., bread, pasta, fruits), proteins (e.g., meat, eggs, legumes), and fats (e.g., nuts, oils, avocados).
The food shown here are those that we consume which are comprised of macronutrients like carbohydrates, proteins, and fats. A healthy diet comprises all of these nutrients as well as minerals, vitamins, and electrolytes. Our body breaks down food in a process called metabolism and the process in which our body transfers energy is called bioenergetics.

Learning Objectives

  • Explain the concept of bioenergetics and its importance in cellular functions.
  • State and explain the first and second laws of bioenergetics.
  • Illustrate how energy is converted and transferred within the body, including examples of exergonic and endergonic reactions.
  • Describe the structure and function of the cell membrane, mitochondria, nucleus, and cytoplasm.
  • Define metabolism and explain the role of metabolic pathways in energy production.
  • Identify the primary substrates used for energy during exercise and describe their metabolic pathways.
  • Classify different types of enzymes and describe their specific roles in metabolic reactions.
  • Explain the concept of enzyme specificity and the “induced fit” model of enzyme action.
  • Define coenzymes and describe their role in assisting enzymatic reactions.
  • Explain the importance of high-energy phosphate molecules like ATP and creatine phosphate in energy transfer.
  • Discuss how an understanding of bioenergetics can inform exercise professionals about muscle energy generation and body responses to exercise.

Bioenergetics

 All living organisms require energy that is generated through chemical reactions. At any given moment, thousands of these reactions are occurring within your cells, enabling your body to produce the energy necessary for sustaining life. This energy can also be stored and used during activities such as physical exercise. Ultimately, all energy on Earth originates from the sun. Plants transform solar energy into carbohydrates, fats, and proteins through photosynthesis. Animals then consume these plants, or other animals, to obtain the energy needed for cellular functions.
        Bioenergetics is a branch of biochemistry that examines the flow of energy from one source to another. It explores how foodstuffs are converted into adenosine triphosphate (ATP), the primary energy currency of cells (Figure 3.1). This process is crucial for understanding how organisms’ harness and utilize energy to maintain life.
Figure 3.1 is a conceptual diagram providing a basic overview of bioenergetics—the study of energy flow through biological systems. The figure illustrates how macronutrients (carbohydrates, fats, and proteins) are broken down through metabolic pathways such as glycolysis, the Krebs cycle, and oxidative phosphorylation. These processes lead to the production of ATP (adenosine triphosphate), the primary energy currency of the cell. Arrows may indicate the direction of energy transfer, with mitochondria highlighted as the central site of aerobic energy production. The diagram also shows chemical waste and byproducts of metabolism such as heat loss, carbon dioxide, and water leaving the body throughout this process. The diagram emphasizes the integration of nutrient metabolism and cellular energy demands.
Figure 3.1 Conceptual diagram providing an overview of bioenergetics.
 Understanding bioenergetics is crucial because it provides the foundational principles by which metabolism operates. Comprehending metabolism is essential for exercise professionals to understand how skeletal muscles generate energy and how the body responds to exercise. Without a thorough grasp of energy flow within the body, it is impossible to fully understand skeletal muscle functions both at rest and during exercise. Metabolism encompasses the chemical reactions responsible for the creation and transfer of energy necessary to sustain exercise.
        There are two fundamental laws of bioenergetics that govern metabolic processes. The first law states that 1) energy cannot be created or destroyed but is converted from one form to another. In the human body, various reactions convert chemical, electrical, mechanical, and thermal energies into different forms. For instance, light entering the eyes is transformed into a chemical signal, which is then converted into electrical action potentials in the brain. Muscles convert chemical energy (ATP) into mechanical energy during muscle contractions, which can be used to exert force on the external environment. The bioenergetic process of energy conversion involves a series of tightly regulated chemical reactions that release energy stored in the chemical bonds of molecules. Ultimately, all energy is eventually transformed into heat.
        The second law of bioenergetics states that 2) energy transfer will proceed in the direction of increased entropy and the release of free energy. Potential energy is stored in the bonds of molecules, referred to as “high energy” bonds. When these bonds are broken, they release a specific amount of energy known as free energy (ΔG), typically measured in calories per mole. For example, the complete oxidation of one mole of glucose releases 686,000 calories[1]. Cells harvest this free energy to perform work, or it can be lost as heat. In thermodynamics, entropy describes the unavailability of a system’s thermal energy for conversion into mechanical work and can also be interpreted as the degree of disorder or randomness in a system. When converting a substrate to a product, by-products such as heat, light, entropy, and free energy are often produced.

Cellular Chemical Reactions

        Chemical reactions within cells are categorized into two major types that work together to sustain energy flow. Exergonic (or exothermic) reactions release free energy, such as those involved in glucose breakdown. The energy liberated from exergonic reactions can drive other reactions. Conversely, endergonic (or endothermic) reactions require an input of energy to proceed. When energy is added to an endergonic reaction, the resulting products contain more free energy than the original substrates.
        Exergonic and endergonic reactions are often linked in a process known as coupled reactions. In coupled reactions, the free energy released from an exergonic reaction is used to drive an endergonic reaction. This systematic linking of reactions ensures the efficient transfer and utilization of energy within cells. An example of coupled reactions is illustrated in figure 3.2 where the same energy that is liberated from reaction 1, is then used to initiate reaction 2.
Figure 3.2 is a schematic diagram illustrating coupled biochemical reactions. Reaction 1 is an exergonic reaction in which substrates A and B are converted into products C and D, releasing free energy. This liberated energy is then used to drive Reaction 2, an endergonic process where substrates E and F are converted into products G and H. Arrows indicate the direction of each reaction, and a connecting line or symbol shows the energy transfer between them.
Figure 3.2 Coupled Reactions. Reaction 1 is an exergonic reaction in which substrates A and B are converted into products C and D, releasing free energy. This liberated energy is then used to drive Reaction 2, an endergonic process where substrates E and F are converted into products G and H. Arrows indicate the direction of each reaction, and a connecting line or symbol shows the energy transfer between them. The figure demonstrates how energy-releasing reactions can power energy-requiring processes, a fundamental principle in cellular metabolism.

Redox reactions

Reduction-oxidation (redox) reactionsinvolve the transfer of electrons between two molecules. These reactions are essential to many fundamental life processes, including photosynthesis, respiration, combustion, and metabolism. When a molecule gains electrons, it undergoes a reduction reaction. Conversely, when a molecule loses electrons, it undergoes an oxidation reaction. A helpful mnemonic to remember this is “LEO the lion says GER,” which stands for “Loss of Electrons is Oxidation” and “Gain of Electrons is Reduction.”
        It is important to note that electrons are not free-floating; they are attached to other molecules by their charge. In many metabolic steps, electrons that are removed are bound to a proton (hydrogen). Figure 3.3 illustrates a hydrogen atom, which consists of a proton and a valence electron. Redox reactions involving hydrogen atoms occur in several stages of catabolism. When hydrogen is removed, a carrier molecule transports the hydrogen (a proton and an electron) to later stages in metabolism. These molecules, known as “electron carriers,” will be discussed further in the co-enzyme section of this chapter.
Figure 3.3 is a simplified atomic diagram illustrating the structure of a hydrogen atom. It shows a single positively charged proton located at the center, representing the nucleus, and a negatively charged electron orbiting around it. The electron is depicted as a small particle in motion, symbolizing its dynamic behavior in the electron cloud.
Figure 3.3 A hydrogen atom is composed of a positively charged proton and a negatively charged electron.​

Energy Transfer and Cell Anatomy

 Energy transfer occurs within the cells of the body, making an understanding of cell anatomy vital to the study of bioenergetics. Cells were first discovered in the seventeenth century by the English natural philosopher Robert Hooke. Over the past 300 years, advancements in cell microscopy have significantly enhanced our understanding of cell structure. We now know that organisms are primarily composed of four elements: oxygen (65%), carbon (18%), hydrogen (10%), and nitrogen (3%)[2]. These elements make up more than 95% of the human body. Additional chemical substances found in the body include sodium, iron, zinc, potassium, magnesium, chloride, and calcium.
        Molecules that contain carbon are known as organic compounds and are often linked with other elements by chemical bonds. For example, glucose (C6H12O6) is an organic molecule and serves as a source of energy classified as a carbohydrate. In contrast, inorganic compounds are molecules that do not contain carbon atoms. Water (H2O), for instance, does not contain carbon atoms and is therefore classified as inorganic.
Figure 3.4 is a labeled diagram of a skeletal muscle fiber, illustrating its basic structure and major organelles. The elongated, cylindrical cell is shown with multiple peripheral nuclei and a surrounding sarcolemma (cell membrane). Inside, longitudinally arranged myofibrils are visible, composed of repeating sarcomeres—the contractile units. Mitochondria are scattered throughout the sarcoplasm, indicating sites of ATP production.
Figure 3.4 The basic structure of a muscle cell (fiber) and its major organelles.​ The elongated, cylindrical cell is shown with multiple peripheral nuclei and a surrounding sarcolemma (cell membrane). Inside, longitudinally arranged myofilaments are visible, composed of repeating sarcomeres—the contractile units. Mitochondria are scattered throughout the sarcoplasm, indicating sites of ATP production. The diagram emphasizes the structural and functional organization of muscle cells in relation to contraction and energy metabolism.
        Cells are highly organized units compartmentalized into smaller organelles that carry out necessary functions. While not all cells have the same function, they share structural similarities. Each cell is surrounded by a semi-permeable membrane composed of a phospholipid bilayer with both hydrophobic and hydrophilic properties. This cell membrane, known as the sarcolemma in skeletal muscle, protects the cell from the external environment and provides structural compartments to house the cell’s inner contents.
        The nucleus is a large, rounded body within the cell that contains the organism’s genetic material in the form of deoxyribonucleic acid (DNA). DNA contains genes that code for proteins, regulating protein synthesis, which determines cell composition and controls cellular activity. Muscle cells, also known as muscle fibers, are unique in that they are multi-nucleated, meaning they have more than one nucleus.
        Another major component of the cell is the cytoplasm, or sarcoplasm in muscle cells. The cytoplasm, also referred to as the cytosol, contains various organelles, including the mitochondria. Mitochondria, often called the powerhouse of the cell, are heavily involved in creating energy from foodstuffs. They are particularly important in skeletal muscle bioenergetics and metabolism due to their role in energy generation. Additionally, the sarcoplasm contains essential proteins such as actin and myosin, which promote organization and prevent structural collapse. Actin and myosin form structures called myofilaments within the muscle cell, providing rigid scaffolding for structure and the ability to produce force. Figure 3.4 illustrates the basic structures of a muscle fiber.

Metabolism

        A significant proportion of chemical reactions in cells occur to create energy from food, which is then used to perform cellular work. These reactions are termed metabolism and are required to maintain life. Metabolism is typically divided into two categories. Catabolism refers to reactions that breakdown molecules to release energy and anabolism are those reactions that synthesize molecules to form larger molecules. Anabolism requires energy to be inputed into the reaction.
      Energy is required for muscle activity, gland secretion, maintenance of nerve and muscle fiber membrane potentials, synthesis of substances in cells, absorption of food from the gastrointestinal tract, and many other functions[3]. A substrate is any substance acted upon by enzymes to create a product molecule. The three forms of usable nutrients in the human body are carbohydrates, proteins, and fats. When consumed, these nutrients are metabolized and used as substrates to create usable cellular energy. A metabolic pathway is a sequence of enzyme-mediated chemical reactions that convert substrates to products.
        During exercise, the primary substances used for energy generation are fats and carbohydrates, with minimal energy contribution from protein metabolism. Therefore, carbohydrate and fat metabolism will be highlighted in this text, along with an explanation of the relationship between carbohydrate, fat, and protein metabolism.

Substrates Used for Energy

        Carbohydrates  are synthesized during photosynthesis by plants from the interaction of CO2, water, and solar energy. Composed of carbon (C), hydrogen (H), and oxygen (O2), carbohydrates exist as monosaccharides, disaccharides, or polysaccharides. Monosaccharides are simple sugars such as fructose and glucose. All carbohydrates are ultimately converted to glucose, which is transported through the blood to all body tissues. Glucose is the preferred carbohydrate for muscles due to its high availability in the blood and the substantial energy yield it provides. Glucose yields approximately 4 kcal of energy per gram and has the chemical formula C6H12O6.
        Muscle cells can store small amounts of glucose in the form of glycogen, a polysaccharide also stored in the liver. Glycogen is synthesized within cells by linking single glucose molecules together with the enzyme glycogen synthase. Glycogen molecules can consist of hundreds to thousands of glucose molecules. Carbohydrate stores in the liver and skeletal muscle are limited to about 2,500 to 2,600 kcal of energy, equivalent to the energy needed for approximately 25 miles (40 km) of running[4].
Figure 3.5 is a structural diagram of a glucose molecule, a six-carbon monosaccharide with the molecular formula C₆H₁₂O₆.
Figure 3.5 The molecular structure of glucose.​ A glucose molecule is a six-carbon monosaccharide with the molecular formula C₆H₁₂O₆. The image shows glucose in its cyclic (pyranose) form, with a six-membered ring composed of five carbon atoms and one oxygen atom. Each carbon is bonded to hydrogen and hydroxyl (–OH) groups in a specific orientation, distinguishing D-glucose as a biologically active isomer. The diagram includes labels for carbon atoms (C1–C6), and hydroxyl groups, highlighting the molecule’s role in energy metabolism and cellular respiration.
Fructose, another monosaccharide, is the sweetest of the carbohydrates and is found in fruits and honey. Disaccharides are formed when two monosaccharides combine. A common example is table sugar, or sucrose, which consists of glucose and fructose. Sucrose is the most common dietary disaccharide in the United States, accounting for nearly 25% of the total caloric intake for most Americans[5]. Polysaccharides, which are complex carbohydrates, contain three or more monosaccharides. Cellulose and starch are two forms of polysaccharides; however, humans lack the enzymes to digest cellulose, which is excreted as waste. Starch, found in corn, beans, grains, and peas, is easily digested and constitutes a significant portion of the human diet[6]. It is recommended that adults obtain 45% to 65% of their calories from carbohydrates[7].
        Fats are also a preferred substrate for energy production during exercise, providing a substantial portion of the energy used during prolonged, less intense activities. Fats contain the same chemical elements as carbohydrates, but with a higher ratio of carbon to oxygen. This higher ratio allows fats to yield more energy per gram than carbohydrates, with 1 gram of fat providing approximately 9 kcal of energy, roughly double that of carbohydrates. Fats are generally classified into three groups: triglycerides, phospholipids, and steroids.
        Triglycerides consist of three fatty acid chains and one glycerol molecule. Fatty acids, the primary fats used by muscles for metabolism, are stored in the body as triglycerides. They are released into the bloodstream through a process called lipolysis, which is regulated by enzymes known as lipases. The glycerol molecule can be used by the liver to synthesize glucose if necessary. Phospholipids, which make up the cell membrane’s phospholipid bilayer, provide structural integrity for cells but are not used as an energy source. Steroids, derived from dietary cholesterol, are components of cell membranes and are necessary for synthesizing sex hormones. Nutritional guidelines recommend that adults obtain 20% to 35% of their calories from fat[8].
        Proteins are not a major energy source during exercise but can be used under certain conditions. To be used for energy, proteins must be converted to glucose through gluconeogenesis or to free fatty acids through lipogenesis in cases of severe energy depletion or starvation. Proteins are composed of amino acids, which can be broken down through deamination. There are 20 amino acids needed by the body to synthesize proteins, tissues, and enzymes, nine of which are essential and must be obtained from food. Proteins can also become intermediates in metabolism to help generate ATP. It is recommended that adults obtain 10% to 35% of their calories from protein[9].
        Enzymes are biological catalysts that increase the speed of metabolic pathways without becoming part of the final product (Figure 3.6). Chemical reactions occur only when the reacting molecules have sufficient initial free energy, or activation energy, to start the reaction. Enzymes lower the activation energy required, thereby conserving energy and improving reaction time. Many enzymes facilitate the breakdown (catabolism) of chemical compounds. All enzymes are proteins that act upon a substrate to create a product. They are highly specific, interacting only with their designated substrate to form an enzyme-substrate (E-S) complex. This complex temporarily changes shape to facilitate the reaction, after which the enzyme returns to its original shape, remaining virtually unaltered. This process is known as the “induced fit” model of enzyme action. Enzymes rely on maintaining their correct conformation, which can be affected by temperature and pH. The major characteristics of enzymes are listed in Table 3.1.
Table 3.1 A summary of the major characteristics of enzymes
Major characteristics of enzymes
Proteins
Specific
Unaltered
Affected by temperature
Affected by pH
Facilitate the reaction
        Other cellular constituents, such as ATP, can regulate enzyme activity. Enzymes can be inhibited through negative feedback, slowing the overall rate of the reaction or pathway. Rate-limiting enzymes, typically found early in the pathway, can be inhibited or stimulated. Allosteric enzymes, which bind to effectors, can either stimulate or inhibit enzyme activity at the active site. These enzymes are major regulators of metabolic pathways.
Schematic diagram illustrating the enzyme-substrate complex mechanism. The image shows an enzyme with a specifically shaped active site binding to a complementary substrate molecule, forming the enzyme-substrate complex. The substrate undergoes a chemical transformation at the active site, resulting in the release of product(s) and regeneration of the enzyme. Arrows indicate the sequence: enzyme + substrate → enzyme-substrate complex → enzyme + product.
Figure 3.6 The enzyme-substrate complex mechanism of enzymes.

Classification of Enzymes

        Enzymes play a crucial role in energy transfer and control the rate of free-energy release. Metabolic pathways that produce a product from a substrate typically involve multiple steps, each catalyzed by a specific enzyme with a specific function. Enzymes are named based on their role in rearranging, adding, or cleaving sub-molecules during a reaction.
  • Kinases: These enzymes add a phosphate group to a molecule.
  • Dehydrogenases: These enzymes remove hydrogen atoms, such as lactate dehydrogenase.
  • Oxidases: These enzymes catalyze redox reactions involving oxygen.
  • Isomerases: These enzymes rearrange molecular substances.
        Most enzymes have names ending in “-ase.” Additionally, many enzymes require other molecules called coenzymes or cofactors to function.
        Coenzymes, also known as cofactors, are non-protein organic substances that assist enzymes. They act as temporary carriers of products and are often considered “helper molecules” in biochemical transformations. The availability of coenzymes can affect enzymatic function and the rate of metabolic reactions. Dietary coenzymes are derived from vitamins. Two important coenzymes in metabolism are Nicotinamide Adenine Dinucleotide (NAD+) and Flavin Adenine Dinucleotide (FAD). NAD+ is derived from niacin (vitamin B3), while FAD comes from riboflavin (vitamin B2). Both NAD+ and FAD are electron carriers essential for ATP production.

High-Energy Phosphates

        High-energy phosphate molecules store potential energy within their chemical bonds, making them vital for energy use in the body. The most immediate source of energy for skeletal muscle contraction is adenosine triphosphate (ATP), known as the universal energy donor. ATP can be broken down when energy is needed for cellular activities. ATP’s middle-range energy potential allows other molecules to donate energy to create ATP from adenosine diphosphate (ADP) and inorganic phosphate (Pi). ATP consists of an adenine molecule combined with a ribose (sugar) and three linked phosphates.
        When ATP is combined with water (hydrolysis) and acted upon by the enzyme ATPase, the last phosphate group is cleaved, releasing approximately 7.3 kcal per mole of ATP under standard conditions [2]. This process reduces ATP to ADP and Pi. To regenerate ATP, a phosphate group is added to ADP in a process called phosphorylation, which requires a considerable amount of energy. Some ATP is generated independently of oxygen availability through substrate-level phosphorylation, while other ATP-producing reactions occur with the aid of oxygen through oxidative phosphorylation. ATP must be continuously synthesized as it is only stored for about 10 seconds in the body, necessitating various metabolic pathways to synthesize ATP from other molecules.
        Creatine phosphate (CrP), also known as phosphocreatine (PCr), is another high-energy phosphate molecule stored in muscles in small amounts and used to quickly generate ATP. Creatine phosphate is depleted in less than 15 seconds of exercise but is resynthesized and stored during rest and recovery.

Chapter Summary

        In this chapter, we explored the fundamental principles of bioenergetics, emphasizing its critical role in understanding how energy flows through living organisms. We discussed the two primary laws of bioenergetics, which govern energy conversion and transfer, and highlighted the importance of cellular structures such as the cell membrane, nucleus, and mitochondria in energy production. We delved into the metabolic pathways that convert carbohydrates, fats, and proteins into usable energy, focusing on the processes of glycolysis, lipolysis, and gluconeogenesis. The chapter also covered the classification and function of enzymes, including the roles of kinases, dehydrogenases, oxidases, and isomerases, as well as the significance of coenzymes like NAD+and FAD in facilitating biochemical reactions.
        Additionally, we examined the role of high-energy phosphate molecules, particularly ATP and creatine phosphate, in storing and transferring energy within cells. Understanding these concepts is essential for comprehending how the body generates and utilizes energy, especially during physical exercise. By integrating these principles, we gain a deeper insight into the intricate processes that sustain life and enable physical activity, providing a solid foundation for further study in bioenergetics and exercise physiology.

Scholarly Questions

  1. What is a substrate?
  2. List 6 characteristics of enzymes.
  3. What are the 3 substrates we use for fuel?
  4. Basic Terms to Know with this section: bioenergetics, glycogen, glucose, metabolism, metabolic pathway, oxidation, reduction, redox reactions, creatine kinase, ATPase, enzyme, co-enzyme, mitochondria, and substrate.
  5. What is the difference between anabolism and catabolism?
  6. Know the basic structure of a muscle cell. What is the cell membrane called? The fluid portion of the cell? Which organelle is the most energy made? Where is the genetic material (DNA) of the cell located?
  7. Can you name some high energy phosphates? Where is the potential energy stored?
  8. Name two co-enzymes. How are they different than enzymes? How do they assist enzymes in biochemical reactions?
  9. What is the chemical formula for glucose?
  10. Approximately how long can ATP be stored?
  11. Describe the roles of kinases, dehydrogenases, oxidases, and isomerases in metabolic pathways.

  1. Guyton AC, Hall JE. Textbook of medical physiology. 11th ed. Philadelphia, PA: Elsevier Saunders; 2006.
  2. Powers SK, Howley ET. Exercise physiology (theory and application to fitness and performance). 9th Edition ed. New York, NY: McGraw-Hill; 2015.
  3. Guyton AC, Hall JE. Textbook of medical physiology. 11th ed. Philadelphia, PA: Elsevier Saunders; 2006.
  4. Kenney LK, Wilmore JH, Costil DL. Physiology of sport and exercise. In. Champaign, IL: Human Kinetics; 2012.
  5. McArdle WD, Katch FI, Katch VL. Exercise physiology: Nutrition, energy, and human performance. In. Exercise physiology: Nutrition, energy, and human performance: LWW; 2014, p 1088.
  6. McArdle WD, Katch FI, Katch VL. Exercise physiology: Nutrition, energy, and human performance. In. Exercise physiology: Nutrition, energy, and human performance: LWW; 2014, p 1088.
  7. Powers SK, Howley ET. Exercise physiology (theory and application to fitness and performance). 9th Edition ed. New York, NY: McGraw-Hill; 2015.
  8. Powers SK, Howley ET. Exercise physiology (theory and application to fitness and performance). 9th Edition ed. New York, NY: McGraw-Hill; 2015.
  9. Powers SK, Howley ET. Exercise physiology (theory and application to fitness and performance). 9th Edition ed. New York, NY: McGraw-Hill; 2015.
definition

License

Icon for the Creative Commons Attribution-NonCommercial 4.0 International License

The Physiology of Exercise Copyright © 2025 by Rosie Lanphere is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License, except where otherwise noted.