Chapter 6 How Cells Harvest Chemical Energy Overview All living organisms require energy to survive, grow and reproduce. In this chapter we will examine how cells produce ATP from carbohydrates and other nutrients. First, we will explore the relationship between cellular respiration (and glycolysis) and photosynthesis to further understand the "web of life". The second part of this unit will examine the stages of cellular respiration by following the path of a glucose molecule through the cell. From glycolysis, to the Kreb's cycle and finally the electron transport chain, we will learn the major components of each pathway. Finally, we will explore alternative to cellular respiration in environments where oxygen is limited, such as those present in the early stages of out planet's development. All of this can look very confusing at first, and is far more complicated than we will be covering. Take each step slowly and individually, and don’t get scared by looking at the figures. This does make sense, but will require some careful understanding, and yes, some rote memorization. You need to focus on the major happenings in each pathway, including: Number of ATP produced and the method by which they are produced Cellular location of the pathway Inputs and outputs of the pathway (which chemical compounds go in and which come out) Key molecules and terms Assigned Reading Text, Pages 89-103 PowerPoint Presentation Chapter Review, Page 104 Testing Your Knowledge, Page 105 Key Terms glycolysis Citric acid (Krebs) cycle electron transport systems pyruvate mitochondria cytoplasm NADH, FADH ATP, ADP fermentation chemiosmosis phosphorylation oxygen acetyl-CoA aerobic and anaerobic Introduction In this unit we will examine how organisms can release this energy for use within the cell. Energy may be released from storage molecules such as glucose by pathways that involve oxygen (aerobic pathways) and pathways that are independent of oxygen (anaerobic pathways). Since oxygen was almost nonexistent on the early Earth, the anaerobic pathways represent the more ancient methods of releasing the energy in chemical bonds. A key concept here is that oxygen is utilized within the mitochondria of eukaryotic cells. So a good concept is to remember that aerobic pathways will occur in the mitochondria, while anaerobic will occur in the cytoplasm. Glycolysis The term glycolysis means "sugar breaking". When we break chemical bonds, energy and electrons are released. We will use the energy to create ATP from ADP and will harvest the electrons to create energy as well. Energy reactions often require an input of energy and glycolysis is not an exception. Glycolysis has both an energy investment and energy releasing phase. Examine Figure 6.7c—notice at the top that we need to invest 2 ATP to start the reaction, while at the end of the reaction we get 4 ATP out for a net production of 2 ATP. In glycolysis we break the 6 carbon glucose to form two 3-carbon molecules called pyruvate. Like photosynthesis, we need to have a molecule to attract electrons. In cellular respiration, the molecule is NAD+, which when combined with an electron, forms NADH. Notice how many steps this simple process requires. Each step has enzymes which must be regulated. You do not need to know every step, but you do need to recognize the inputs/outputs of each stage. Formation of Acetyl-CoA Literally for billions of years the glycolysis pathway described above was the primary method of getting energy from biomolecules. However, after the levels of oxygen on the planet rise to significant levels, cells begin to use the power to oxygen to "burn" biomolecules. However, this requires a specialized area of the cell, the mitochondria. The pyruvate leaving glycolysis must be converted to a compound called acetyl-CoA before it may be utilized in the next pathway. This is essentially a rearrangement step - but an important one. While we will be talking about glucose in this lecture, you already know that your body can use both proteins and fats as energy as well. Many of these biomolecules may be converted to acetyl-CoA. So this molecule acts as a key point in the use of biomolecules to produce ATP. Note that since each glucose is split to form 2 pyruvates in glycolysis, 2 acetyl-CoA molecules enter the next cycle. Figure 6.8 (preparatory steps) shows that this process releases electrons which are captured by NAD+ to form NADH, and some carbon dioxide as well. Citric Acid Cycle This is also called the Krebs cycle (in honor of the man who first described it), though the text refers to it as the citric acid cycle. Be aware of both names, and that they mean the same process. Finally we are in the mitochondria and ready to "burn" our food for energy. But as you can see by Figure 6.9B, this is a very complicated pathway. The first thing that you should notice is that the pathway is circular. The outputs of the pathway (oxaloacetate) are used to start the pathway over again. With the exception of acetyl-CoA and oxaloacetate, we will not learn the other intermediate molecules. This process effectively uses oxygen to break the C-C bonds found in the acetyl-CoA (which was pyruvate). As the bonds are broken, energy may be harvested (to form ATP), but usually electrons are released and then captured by two molecules. One of these is NAD+ (same as the previous 2 steps), but the other is FAD+. For our purposes these molecules will be treated the same, but there are some important differences. So what are the outputs of this pathway. Well you will notice that 2 carbons enter the cycle (one of the carbons from pyruvate was lost in the acetyl-CoA step) and that 2 CO2 molecules depart. Thus after the Krebs cycle all of the carbons leave the cycle as CO2. We also produce 1 ATP, 3 NADH and 1 FADH2. But for each glucose that entered the cycle, 2 acetyl-CoA molecules are eventually produced. Thus we need to double the outputs of this pathway. So the Krebs cycle produces 2 ATP, the same as glycolysis! Since we use oxygen to generate the ATP, this is often called oxidative phosphorylation. Oxidative Phosphorylation The processes above have only given us 4 ATP for each glucose entering the system. This is simply not efficient. But what has been generated is 10 NADH and 2 FADH2 molecules. These molecules have captured electrons and we can use these electrons. The process is much the same as above. We will hand the electrons over to a series of proteins located in a membrane and then generate an electrical potential. Yet instead of using this electrical potential to make C-C bonds, we will use them to generate large amounts of ATP. Take a good look at Figure 6.10. Notice that the NADH and FADH2 molecules enter the inner compartment of the mitochondria and release electrons. Once the electrons are handed off to the integral proteins, we regenerate NAD+ and FAD+, which will then return to the pathways above (recycled). We can then use the electrons to generate ATP. This process produces between 28 and 32 ATP (depending on the cell). All told, from glycolysis on we generate about 36 ATP per glucose for an efficiency of about 39%. Not bad for a biological organism! Notice that the end of the ETS uses oxygen to accept the electrons to form water. Anaerobic Pathways Many prokaryotic organisms do not require oxygen to extract energy from glucose. However, in the aerobic pathways the ETS uses oxygen to recycle NAD+ and FAD+. In anaerobic pathways, another method must be found. Figure 6.13A and Figure 6.13B show two different methods by which this may occur. While we may think that these pathways are not important, in actuality they are. Lactic acid fermentation may be used to make cheese and is also used by our muscles when our circulatory system does not deliver enough oxygen to the tissues. Alcohol fermentation is used to produce alcoholic beverages and other alcohol-related compounds. Lactate Formation Ethanol Formation Concepts Recognize the relationship between energy-releasing and energy-acquiring pathways Know the difference between aerobic and anaerobic respiration For glycolysis, the citric acid cycle and the electron transport chain, know the following: cellular location, inputs to the stage (chemical compounds), outputs (chemical compounds), number of ATP produced. Understand how certain poisons work by affecting steps of oxidative phosphorylation. Understand lactic acid and alcohol fermentation. Understand how fats, carbohydrates, and proteins are broken down to enter the aerobic respiration process. Review Material MyBiology.com—Study guides and resource for this text. Specifically look at MP3 Tutor and all Web Activities