AL Bio notes Respiration P.1 RESPIRATION O rganisms maintain themselves by the continual expenditure of energy. Most cells derive this energy from the breakdown of food molecules, a process known as cellular or internal respiration. Substances that are broken down to yield energy are called respiratory substrates. In most organisms glucose is broken down to carbon dioxide and water in a process which requires molecules of oxygen, and is known as aerobic respiration. C6H12O6 glucose + 602 oxygen 6C02 + carbon dioxide 6H2O water + energy In the absence of oxygen molecules, anaerobic respiration takes place in which glucose is partially broken down, to lactic acid in animals, and to carbon dioxide and alcohol in plants. The latter process is also called alcoholic fermentation: anaerobic respiration in plants C6H12O6 C2H5OH + 2CO2 (ethyl alcohol) anaerobic respiration in animals C6H12O6 CH3CHOH.COOH (lactic acid) + C02 + + energy energy The partial breakdown of glucose in anaerobic respiration yields only a fraction of the chemical energy stored in the molecule. Glucose is broken down in a series of carefully controlled reactions, during which energy is made available in relatively small amounts which can be harnessed and made to perform work. A much smaller proportion of the energy in the glucose molecule is dissipated as heat compared with combustion. (at most about forty per cent). The energy made available during the respiratory process is stored in molecules of adenosine triphosphate, usually called ATP. AL Bio notes Respiration I) THE ROLE OF ATP AS A STORE AND SUPPLIER OF ENERGY ATP occurs in all living cells. It is the un___________ energy 'currency'. All energy-spending activities such as synthesis, muscle contraction, transmission of nerve impulses …….., depend on ATP as their im________ source of energy. Energy is supplied by the splitting of the bond that attaches the terminal phosphate groups to the rest of the molecule. The bond is broken by hydrolysis. This reaction yields a large amount of energy (about 33 kJ mol-1). Because of this high energy yield, ATP is often called an 'energy-rich' or 'high-energy' molecule. ATP thus provides a source of energy that is easily and rapidly accessible to any energy-requiring process. Matching : Analogy of Energy availability and Money currency ATP glucose carbohydrate reserves, starch and glycogen fat reserves II) THE FORMATION OF ATP Molecules of ATP are constantly being 'spent' and must be replaced. They are formed from ADP and inorganic phosphate by condensation. This reaction is also a phosphorylation. The regeneration of ATP requires a supply of energy = that which it gives on hydrolysis. This is supplied by respiration. There are two mechanisms that direct the energy yield from respiration into the formation of ATP; namely s_________-linked phosphorylation and o___________ phosphorylation. a) Substrate-linked phosphorylation At 3 steps in the respiratory sequence where p________ is detached from one of the intermediate compounds of glucose breakdown (the respiratory substrate), together with sufficient energy to form a bond with ADP. This process is thus substrate-based or linked and, unlike oxidative phosphorylation, does not need O2. b) Oxidative phosphorylation This process involves oxidation reactions. Such reactions tend to have a high ‘energy yield’, and it is this energy that is used to combine inorganic phosphate and ADP. Investment accounts deposit accounts cash cheque P.2 AL Bio notes Respiration P.3 The following types of reaction are all described as Oxidations: i) The addition of oxygen to a molecule, e.g. C + O2 CO2 (the burning of carbon in air) ii) The removal of hydrogen from a molecule (dehydrogenation). e.g. AH2 + B A + BH 2 iii) The removal of an electron from a charged ion, e.g. Fe ++ - e (an electron) Fe +++ All three types of oxidation are involved in oxidative phosphorylation : 1) Pairs of hydrogen atoms are removed from a number of respiratory intermediates by dehydrogenase enzymes; the respiratory intermediates are thus oxidised. 2) The hydrogen removed is transferred to hydrogen-acceptors, usually nicotinamide adenine dinucleotide (NAD+). These molecules are also described as coenzymes because they are essential for the functioning of the dehydrogenase enzymes. 3) The hydrogen is split into protons and electrons on its way through the series of carriers, the electrons are eventually combined into with oxygen to form water. The whole sequence of carriers, including the initial hydrogen-acceptors, comprises the Electron transport chain or Respiratory chain. c) Electron Transport Chain (ETC) / The Respiratory Chain A series of coupled ox_______/re________ reactions where e________ are passed like hot potatoes from one membrane-bound p______ to another before being finally attached to a terminal electron acceptor (o______ in respiration, N________ in photosynthesis ). ATP is formed in the process. Each of the carriers in the respiratory chain is successively reduced and then oxidised as it first accepts and then hands on the hydrogen. ETS moves both electrons and protons: electrons are passed from carrier to carrier in the membrane, while protons are moved from in_____ to out____ of membrane (inner membrane). In fact, early in the series of protein carriers, it is only the electrons of the hydrogen atoms that are passed along the chain. Hydrogen atoms are split into hydrogen ions and electrons. H H+ + e- (an electron) AL Bio notes Respiration P.4 Each carrier in this chain has a greater affinity for electrons than its predecessor so there is a one-way flow of electrons along the chain. In particular the electron transport chain contains several cytochromes, which are a group of iron-containing proteins (h____ group). The state of the iron is altered from the oxidised ferric (Fe + + +) form to the reduced ferrous (Fe ++) form, and then back to the oxidised form as the cytochrome accepts and then passes on electrons. Fe+++ (oxidized) cytochrome + - electron electron Fe++ (reduced) cytochrome At the final cytochromes in the chain, the electrons combine with hydrogen ions, and then combine with molecules of oxygen to form water. 2H+ + 2e- 2H 2H+ + O2 H2 O Incidentally, cyanide exerts its toxic effects by binding cytochrome oxidase so as to prevent the binding of o______. It thus acts as a respiratory in_________, and in large amounts can rapidly cause death through energy lack. The energy yield is on average 3ATP from each NADH entering the ETC. AL Bio notes Respiration Reference reading: P.5 Proton gradient / Chemiosmotic hypothesis (Peter Mitchell, 1961) As electrons flow through ETC, at certain steps protons (H+) are moved from inside to outside of the membrane. This builds up proton gradient; since + charges are removed from inside of cell, -ve charge remains inside, mainly as OH- ions. pH just outside membrane can reach 5.5, pH just inside membrane can reach 8.5 ---> difference of 3 pH units, or 1000x concentration differential of H+ across membrane. This represents potential energy stored up in proton gradient. 1. Protons are translocated across the membrane, from the matrix to the intermembrane space 2. Electrons are transported along the membrane, through a series of protein carriers 3. Oxygen is the terminal electron acceptor, combining with electrons and H+ ions to produce water 4. As NADH delivers more H+ and electrons into the ETS, the proton gradient increases, with H+ building up outside the inner mitochondrial membrane, and OH - inside the membrane. Membrane is basically impermeable to protons, so gradient doesn't get wasted away by leaky reentry. ATP synthase protein complex contains only channels for proton entry. As protons push in through channel : ADP + Pi ---> ATP. This can be called chemiosmotic phosphorylation or oxidative phosphorylation. III) THE BREAKDOWN OF GLUCOSE Aerobic and anaerobic respiration share a common initial pathway known as glycolysis (which means sugar-splitting), in which the glucose is broken down through a series of reactions to pyruvic acid (2-oxopropanoic). The breakdown of glucose to pyruvic acid takes place in the c_________ and does not require the presence of oxygen molecules. An outline of glycolysis This is the initial pathway shared by both aerobic and anaerobic respiration. The process takes place in the cytoplasm and is not dependent on the presence of oxygen. It comprises two stages; in the first, glucose is phosphorylated to fructose disphosphate. After phosphorylation, the substrates were eventually broken down to two pyruvic acid molecules. Each of the reactions is catalysed by an enzyme. AL Bio notes Respiration P.6 Glucose Glucose is a rather unreactive molecule. Before it can participate in the reactions that follow, it must be supplied with activation energy. This is achieved by two successive phosphorylations to form a hexose diphosphate (fructose diphosphate). 2ATP 2ADP Fructose diphosphate Fructose diphosphate splits into two, interconvertible, 3-carbon compounds known as triose phosphates. Glyceraldehyde phosphate Dihydroxyacetone phosphate NAD+ Triose phosphate is oxidized when Hydrogen is removed by NAD+ and Inorganic phosphate is added to form diphosphoglyceric acid (DPGA). The ‘new’ phosphate is joined to the rest of the molecule by an energy-rich bond. (this bond derived it’s energy from reorganization of energy within the molecule). NADH & H+ inorganic phosphate Diphosphoglyceric acid ADP Diphosphoglyceric acid reacts with ADP to form ATP (substrate linked phosphorylation) and phosphoglyceric acid (PGA). ATP Phosphoglyceric acid The PGA is rearranged to form pyruvic acid. The phosphate is transferred to ADP forming ATP.(substrate linked phosphorylation) ADP ATP Pyruvic acid a) The Energy yield of Glycolysis Summary of glycolysis input output One glucose Two pyruvic acid molecule molecules 2ATP (for phosphorylation of glucose and fructose) 4ATP from substrate level phosphorylation Thus in conditions where oxygen is available the net gain from glycolysis is eight molecules of ATP per glucose molecule. 2ADP and Pi 2 NAD+ Thus, the net yield is two molecules of ATP per glucose molecule. However, if oxygen is present the two pairs of hydrogen atoms removed by NAD+ (one pair from each triose phosphate) can be passed to the ETC and will there yield a further six molecules of ATP (by oxidative phosphorylation). (NADH & H+ ) X 2 The fate of pyruvic acid differs according to whether oxygen is present or not and, in the latter circumstance, according to whether plant or animal cells are being considered. AL Bio notes Respiration P.7 ii) The final anaerobic pathway in animals In animal cells the pyruvic acid is reduced by the NADH, formed during glycolysis, to lactic (2-hydroxypropanoic) acid. This also regenerates the NAD+ (coenzyme for dehydrogenation) needed in glycolysis. i) The final anaerobic pathway in plants In the absence of oxygen pyruvic acid is converted to acetaldehyde (ethana1) and carbon dioxide. The NADH formed during glycolysis is then used to reduce the acetaldehyde to ethanol. The oxidised NAD+ is then regenerated to pick up more hydrogen atoms from glycolysis. b) The Energy yield of Anaerobic respiration The net yield from anaerobic respiration is thus 2 ATP per glucose molecule. Two moles of ATP store about 66kJ. This is a small return from glucose, which stores about 2800 kJmol -1. IV) THE AEROBIC PATHWAY If oxygen is available pyruvic acid enters mitochondria and there embarks on a series of reactions that involves the removal of H atoms to the ETC--and the removal of CO2 molecules. The removal of CO2 is always linked to a dehydrogenation and so is called oxidative decarboxylation. Before entering the cycle, pyruvic acid (3C) is converted to acetyl coenzyme A (2C) (acetyl-CoA) by oxidative decarboxylation. The Krebs cycle may be summarized into 4 major steps : 1. Formation of a 6-C citric acid molecule Acetyl-CoA (2C) combines with oxaloacetic acid (4C), to form citric acid (6C). 2. Oxidation of the 6-C molecule Citric acid (6C) is oxidized to -Ketoglutaric acid (5C) by oxidative decarboxylation. 3. Oxidation of the 5-C molecule -Ketoglutaric acid is oxidized, to a 4-C intermediate by oxidative decarboxylation. 4. Oxidation and regeneration of Oxaloacetic acid Further dehydrogenation occurs and oxaloacetic acid is regenerated. The cycle starts again with another acetyl-CoA produced from pyruvic acid. AL Bio notes Respiration P.8 The Krebs or Tricarboxylic acid cycle (TCA cycle) Take place in the matrix of mitochondria. only operate if oxygen is available to act as the final electron acceptor. 6-carbon citric acid is gradually broken down by four dehydrogenations and two decarboxylations to reform the 4-carbon oxaloacetic acid. Taking into account the oxidative decarboxylation when pyruvic acid are converted into acetyl CoA, three CO2 molecules and 5 pairs of Hydrogens are removed. a) The Energy yield of Aerobic respiration 5 pairs of H atoms are removed as pyruvic acid is broken down in the aerobic pathway, 4 pairs from each turn of the Krebs cycle and one pair from the initial conversion of pyruvic acid into acetyl-CoA. The total yield from this part of the respiratory process is thus ten pairs of hydrogen atoms per glucose molecule (each glucose molecule is broken down to two molecules of pyruvic acid). 8 pairs are removed by NAD+, each pair providing sufficient energy to form 3 ATP molecules. The 2 pairs removed by FAD each provides enough energy to form 2 ATP molecules. To these must be added the 2 pairs of hydrogen atoms removed during glycolysis. These are passed via a carrier/ shuttle in the mitochondrial membrane to either NAD+ or FAD within the mitochondria (varies according to tissue type) resulting in the formation of either 6 or 4 ATP molecules. The total number of ATP molecules formed by oxidative phosphorylation is thus : (8 x 3) + (2 x 2) + 6 or 4 = 32 or 34. To this must be added the 4 ATP molecules formed by substrate-linked phosphorylation ( 2 in glycolysis, 2 in krebs cycle). The total yield from aerobic respiration is thus 36 or 38 molecules of ATP per glucose molecule. ATP yield during aerobic respiration of one molecule of glucose Respiratory process No.of reduced hydrogen carrier molecules formed No. of ATP molecules formed from ETC No. of ATP molecules from substrate-level phosphorylation Total No.of ATP molecules glucose to pyruvate (Glycolysis) pyruvate to acetyl CoA Krebs (TCA) cycle Total ATP = 38 AL Bio notes Respiration P.9 Q. Below is called Warburg apparatus which has been designed to measure the effect of various substances on the rate of respiration. Mung beans seeds were crushed with isotonic buffer enriched with glucose in cold. Its extract was made by a high speed differential centrifugation. The supernatant made by 1,000 g centrifugation was obtained and used in this apparatus. a) Why was it necessary to crush the seeds with isotonic buffer enriched with glucose in cold ? (4m) b) What essential component of the cell must be present in the extract? Why was it necessary? (2m) c) Why should the KOH be separated from the mung bean extract? What was the purpose of using the filter paper? (2m) d) After setting up the apparatus, its was left undisturbed. The manometric reading was recorded at one minute intervals. At ten minutes, the Warburg apparatus was tilted slightly to mix the ADP from the side arm to the mung bean extract. The result was recorded by graph I below. i) Explain the shape of graph I I) before 10 minutes. (2m) II) after 10 minutes. (2m) ii) Identify two possible sources of errors which leaded to inaccuracy of the experiment. which these errors could be minimized. Suggest two ways in (4m) e) Another experiment was performed to demonstrate the effect of substance X on respiration. Substance X was placed in the side arm of the Warburg apparatus instead of ADP. The same experiment was performed and substance X was added to the extract at 20 minutes after commencement of the experiment. The result was recorded in graph II above. Name a possible substance for 'X', explain how it might lead to the effect as shown in graph II. (4m) AL Bio notes Respiration b) P.10 The Site of the Aerobic process Mitochondria tend to be most abundant in cells with a high e________ requirement. They are sausage-shaped organelles comprising two m_______ layers. Within the inner membrane encloses an aqueous solution, the m______. The inner membrane is folded into c______ which project into the matrix. The matrix contains the en______ involved in the Krebs cycle and the breakdown of fatty acids, while the inner mitochondrial membrane contains the en and electron c of the electron transport chain (ETC). There are tiny spheres on the inner membrane which are believed be the sites of ATP synthase, the enzyme that brings about the formation of _____ from ADP and inorganic phosphate.. The cristae presumably serve to increase the surface area for enzyme and carrier attachment. Supporting this, it has been observed that the cristae are more numerous in cells with a high energy requirement. Draw an annotated diagram to show how the structure of a mitochondrion is related to its function (5m) AL Bio notes Respiration V) P.11 RESPIRATION USING OTHER SUBSTRATES Carbohydrates are not the only source of energy. In certain circumstances, fats and proteins can supply part of the organism's energy needs. The breakdown of fats and proteins is largely dependent on the presence of o_______ as they are converted into substances, most of which enter the Krebs cycle where they are oxidised to yield their energy. a) The use of fats as a respiratory substrate Fats form the long-term energy stores in many organisms. They are generally used only when the carbohydrate reserves are exhausted. Fats usually have a much higher energy yield than carbohydrate. Some tissues like liver, and seeds possessing large deposits of fats and are able to use fat directly as a respiratory substrate without first converting it to carbohydrate. The fats are first removed from the fat depots (under the skin and around various internal organs), a process known as mobilisation. They are then transported to the liver where they split into fatty acids and glycerol. The glycerol is phosphorylated to form triose phosphate / glyceraldehydes phosphate which enters the glycolytic sequence. Fatty acid molecules are oxidized by a process called β-oxidation, which involves 2C fragments of acetyl CoA being split off from the acid until the entire fatty acid molecule has been broken up and oxidised. This takes place in the matrix of the mitochondria. Each acetyl CoA formed can enter the Krebs cycle. The longer the fatty acid chain, the greater will be the energy yield. E.g 147 ATP can be obtained from one stearic acid molecules (C17H35COOH). b) Protein as a respiratory substrate Many plant seeds employ protein as an energy storage material. Animals do not generally employ protein as an energy reserve; they do so at the expense of their own tissue proteins. Accordingly, proteins are only respired by animals as a last resort in cases of prolonged starvation. When proteins are respired they are first broken down into amino acids. These are then deaminated, that is, their nitrogen-containing amino group is removed. The products of deamination are ammonia and an α-keto acid AL Bio notes Respiration P.12 Krebs cycle CH3CH.NH2COOH alanine + O2 CH3CO.COOH pyruvic acid + NH3 ammonia The ammonia is toxic and must be removed from the body. The residue (an α-keto acid) enters the respiratory sequence. VI) SURVIVAL IN CONDITIONS OF OXYGEN SHORTAGE The majority of organisms today are so dependent on the aerobic respiration that they will die if deprived of oxygen for more than a few minutes. Such organisms are described as aerobes. At the other extreme, some organisms depend entirely on anaerobic respiration. These organisms are called anaerobes, and include many parasitic flatworms, as well as many of the bacteria that bring about the decay of organic material. Both of these live in conditions where oxygen is not readily available. Some anaerobes are actually poisoned by oxygen. Certain organisms, notably yeast and some other fungi, are described as partial / facultative anaerobes because, although they normally respire aerobically, they can respire anaerobically for extended periods if oxygen is not available. It is sometimes mistakenly supposed that yeast can go on respiring anaerobically indefinitely. This is not the case, yeast is unable to grow and reproduce when respiring anaerobically, and the toxic alcoholic end-product of the process will kill the yeast if it exceeds a certain concentration, usually about eighteen per cent. Returning to aerobic organisms, while it is true that they are rapidly killed by total oxygen deprivation, most have evolved mechanisms which enable them to survive periods of oxygen shortage. One such adaptation is the possession of tissues which can respire anaerobically for limited periods. The muscles of vertebrates provide an example of such a tissue. They can survive anaerobic conditions for many minutes, much longer than, for example, brain cells which suffer irreparable damage if deprived of oxygen for more than one or two minutes. One value of this is that in conditions of oxygen shortage, blood can be diverted away from the muscles to the tissues that need it most. This mechanism is particularly well developed in diving animals such as the seal. During a dive, constriction of the blood vessels supplying the muscles cuts off their blood supply and diverts blood (carrying oxygen) to the heart and brain. The muscles respire anaerobically and the heart rate slows to accommodate the reduced circulation. In this way these animals can remain submerged for up to twenty minutes without harm. Human muscles 'switch over' to anaerobic respiration during strenuous activity, such as running, when the circulatory system cannot supply oxygen to the muscles sufficiently quickly to supply their energy needs. When the muscles respire anaerobically, lactic acid accumulates. The length of time that the muscles can respire anaerobically is largely determined by their tolerance to lactic acid. Regular exertion can build up tolerance to lactic acid. The accumulation of lactic acid creates what is known as the oxygen debt. After the exercise, the lactic acid is converted back to pyruvic acid and enters the aerobic pathway. The debt must then be repaid, that is, extra oxygen must be taken in to complete the breakdown of the pyruvic acid derived from lactic acid. This is usually achieved by panting rapidly in the period immediately after the activity. Some organisms vary their respiratory habits during their life history. For example, many plant seeds respire anaerobically, and the pupal stages of certain insects enter an inactive stage (diapause) during which they respire anaerobically. AL Bio notes Respiration Reference Exercise: P.13 Exercise and training Respiration is generally aerobic. Most muscle tissue, however, also respires anaerobically and produce lactic acid (lactate) as waste products. Lactate is toxic if allowed to accumulate so, after a period of activity has ended, it has to be converted back to glucose or changed to carbon dioxide in aerobic respiration. Obviously, the amount of lactate that is formed during exercise determines the amount of oxygen needed for its conversion after the exercise is over. This oxygen is referred to as the oxygen debt; there is a limit to the oxygen debt which any individual can sustain. Consider a male athlete. The amount of work that he can do in a given time will be determined largely by the amount of oxygen that he can absorb in that time and the extent of the oxygen debt that he is able to sustain. If we assume that for every dm 3 of oxygen available to him, he can generate around 20 kJ, it is possible to calculate how much energy he can make available for doing physical work. For example, suppose he can absorb oxygen at a maximum rate of 2.5 dm 3 minute -1 and can sustain a maximum oxygen debt of 10 dm 3 over a race lasting 3 minutes he can generate [(2.5 x 3) + 10] x 20 = 350kJ. Of course, his muscles are inefficient at converting this into physical work and a figure of around 20 per cent is not unreasonable. So the individual in our example could perform a maximum of 70 kJ of work. When athletes compete, one of the major factors determining who wins is which of them has the largest amount of oxygen available to him or her in terms of consumption and debt. Training must be designed to increase the rate at which an individual can absorb oxygen, and it is this which is used as a measure of physical fitness. (Of course, it is difficult to measure maximum oxygen consumption in field conditions, but there is an almost linear relation between that and heart rate, so maximum heart rate is used instead.) It should be remembered that not all tasks are equally dependent upon aerobic fitness. A trained sprinter, for example, takes less than 10 seconds to run 100 metres. It has been calculated that this requires 6 dm3 of oxygen, of which 0.5 dm 3 or less is inspired during the run, so that an oxygen debt of 5.5 dm 3 or more is built up. So this activity is largely anaerobic and only around 8 per cent aerobic. Virtually all field events javelin, discus, and the jumps are almost entirely anaerobic. A track event lasting a little longer than the sprint, such as a 400-metre race lasting 50 seconds, will be about 80 per cent anaerobic and 20 per cent aerobic and a 2000-metre rowing event lasting 6 minutes will be perhaps 55 per cent aerobic and only 45 per cent anaerobic. This means that an athlete can be advised to choose an event to complement his or her own specific capacities and, more important, to train appropriately for that event. By training, an athlete can modify several other characteristics besides the one mentioned above. The maximum oxygen debt that can be incurred by an untrained individual rarely exceeds 8 to 10 dm 3, but good athletes can double this figure. Efficient use of the muscles is also improved by training and the cardiac output (the volume of blood pumped per minute) can be greatly increased. Measure Heart rate at rest (beats minute -1) 3 Stroke volume at rest (cm ) 3 Stroke volume, maximum (cm ) 3 Heart volume (cm ) -1 Ventilation rate at rest (breaths minute ) Ventilation rate ( maximum breaths minute -1 ) 3 Lung capacity (dm ) VO2 rest 3 -1 (cm kg 3 -1 minute -1 ) minute -1 VO2 max (cm kg ) Lactate at rest (mg 100cm- 3) Lactate, maximum -3 (mg l00cm ) Pre-training 72 64 Post-training 58 79 International 36 128 120 140 200 750 820 1200 14 12 12 40 7.2 45 7.2 55 7.4 3.6 3.8 4.1 40 50 77 20 110 20 125 20 185 Summary of the changes taking place in a variety of cardiovascular and respiratory measures for a normal individual before and as a result of training. The equivalent figures for an international athlete are also given. (VO 2 = oxygen uptake.) a From the data in the table calculate the cardiac output in dm 3 minute -1 of the three athletes at rest. b Which of the rows of data indicates most directly the athletes' ability to sustain oxygen debt ? c Describe the changes that have occurred to the heart as a result of the training in the international athlete compared with the pre-training athlete. AL Bio notes Respiration d P.14 There is a very big difference between the data for the normal individual after training and those for the international athlete. Apart from the duration and extent of the training, what is the main factor determining the features such as maximum cardiac output and maximum oxygen uptake ? Oxygen debt The second aspect of training concerns the most efficient use of the athlete's physical capabilities. A runner who can maintain 85 per cent of a VO2max of 70cm3 kg-1 minute-1 will go faster than one who can only maintain 70 per cent of a higher consumption such as 80 cm 3 kg -1 minute -1. This may seem a small difference but modern competitive sport is decided by much smaller percentage differences than this. What happens if the first runner mentioned above runs at a higher rate of consumption than the 85 per cent quoted? If, in practical terms, 85 per cent of the VO 2max is the maximum he or she can maintain for long periods, then the runner has to obtain the additional energy from anaerobic sources. In other words, for this runner 59.5 cm3 kg-1minute' is the anaerobic threshold. With training this threshold can be elevated to come closer and closer to the VO 2max. If the runner just dips into anaerobic energy sources for a few seconds then there will probably be no apparent penalty; but if the requirement is for more than six seconds or so then anaerobic respiration will be used in earnest and the penalty will be the generation of lactic acid. Lactic acid is one of the main causes of fatigue when muscle is forced to respire anaerobically. a What short-term beneficial effect will the athlete gain from the presence of lactic acid in the blood? Obviously, training for a particular event, such as a 1500-metre track event, will have an important object besides improving general fitness and increasing the capacities mentioned under 'Exercise and training'. It will concentrate upon ensuring that the athlete does not arrive at his or her maximum oxygen debt before reaching the winning tape. In any race, the maximum amount of energy that can be expended is given by the maximum oxygen uptake for the duration of the race plus the athlete's maximum oxygen debt. Suppose that an individual may incur an oxygen debt of 15 dm 3. To run at 8 metres per second he requires oxygen at the rate of 0.2 dm 3 per second. His maximum rate of oxygen absorption is 4.0 dm 3 minute -1. b How far can he run at this speed before becoming completely exhausted? c What proportion of the energy that he has then used is associated with the oxygen debt ? This amount of energy will need to be 'repaid' after the race is finished. Figure on the right represents the demand and usage of oxygen by a runner during a short period of vigorous running. By the end of the race the subject was completely exhausted and could run no further. d From the graph, what is the approximate value of the maximum oxygen debt substainable by this athlete ? 3 3 3 A 12 dm B 21 dm C 33 dm e What is represented by the three areas, L, M, and N on the graph ? AL Bio notes Respiration f P.15 What is the maximum rate of oxygen consumption by this athlete ? Two runners. A and B, can sustain maximum oxygen debts of 15 and 10dm 3 respectively. Additionally A has a VO 2 max of 3 dm3 minute -1 and B has a VO 2max of 4 dm3 minute -1. The two runners are matched in two races. One race is over 400 metres and takes about 50 seconds to complete; the other is over 5 kilometres and takes about 14 minutes to complete. The runners are of the same mass and operate with the same muscular efficiency. g VII) Which runner will win each of these two races ? MEASURING RESPIRATORY RATES The rate of respiration is usually measured in terms of the amount of oxygen consumed or the amount of carbon dioxide produced. The apparatus used to measure respiratory rates is called a respirometer. The respiring organisms are placed in an airtight vessel and changes in gas volume are measured at intervals. If oxygen uptake is to be investigated, sodium hydroxide is placed in the apparatus to absorb carbon dioxide and then any decrease in gas volume can be attributed to oxygen consumption by the respiring tissue. The respiratory rate is generally taken as a measure of the metabolic rate. The two rates are approximately equivalent, as all metabolic processes require energy which is provided by respiration. What are the assumptions taken? Suggest the necessary precautions for the experiment? AL Bio notes Respiration Common misconceptions 14 Anaerobic respiration occurs in the absence of oxygen. This is NOT the case for skeletal muscles, which carry out anaerobic respiration even when oxygen is in abundance. The significance of anaerobic respiration in the skeletal muscle is to provide additional energy to the skeletal muscle so that it can contract more vigorously. 15 Anaerobic respiration produces extra energy for muscle contraction because it can oxidise food faster than aerobic respiration In most cases, aerobic respiration is still a major source of energy for muscle contraction, whereas anaerobic respiration supplies additional energy for promoting maximum exertion It is only under very strenuous muscle activity taking place within a short period, e.g. in a 100m sprint, that lactic acid fermentation will generate most of the energy required for muscle contraction. 16 During exercise the muscle stops aerobic respiration but carries out anaerobic respiration as a result of reduced oxygen supply. In fact, the oxygen supply to the muscle is greatly increased during vigorous muscle contraction so as to maintain a high rate of aerobic respiration. The onset of anaerobic respiration only serves to provide an additional amount of energy for maximum exertion. Also note that anaerobic respiration occurs only in the skeletal muscles, it does NOT take place in all body cells during exercise. 17 “During vigorous exercises, in spite of rapid breathing and strenuous pumping by the heart, oxygen cannot reach the muscles fast enough to supply their needs. The muscles then switch from aerobic respiration to anaerobic respiration, which does not use oxygen.” A better statement should be “During exercise, the muscle cells are carrying out aerobic respiration at a very high rate because of an increased oxygen supply from the blood, but anaerobic respiration is also occurring simultaneously to provide additional energy for promoting maximum exertion.” Explanation for Fig 21: When the skeletal muscle is actively contracting, some glucose molecules are broken down anaerobically into lactic acid (also a 3-carbon compound), while other glucose molecules are broken down aerobically into carbon dioxide and water. While aerobic respiration is occurring in full swing, anaerobic breakdown of glucose provides additional amount of energy for muscular contraction. This additional amount of energy can be very significant in urgent situations, such as when the animal is running away from a predator or when the animal is chasing a prey. 18 Sprints are purely anaerobic and marathons are purely aerobic activity. Most physical exercises involve a mixture of BOTH aerobic and anaerobic muscular activities, but they differ in the proportion of energy contribution from them. P.16 AL Bio notes Respiration 19 Lactic acid is a wasteful by-product of anaerobic respiration to be removed by the liver Lactic acid is an energy-rich by-product of anaerobic respiration in skeletal muscles, its value in providing metabolic energy is recovered by the liver. Anaerobic respiration in the skeletal muscle does not use oxygen. This process produces lactic acid as a waste product. As lactic acid is toxic if allowed to accumulate in the blood, the body can only carry out anaerobic respiration for a brief period of time, after which the lactic acid has to be converted back to glucose or glycogen, or changed to carbon dioxide through aerobic respiration. In the liver, about 20% of the lactic acid in blood is broken down aerobically to produce a large amount of ATP, which is used to convert the remaining 80% of lactic acid to glycogen. The oxygen required to do this is called the oxygen debt. Thus all ATP molecules used by muscle cells come ultimately from aerobic respiration. This explains why one needs to breathe deeply for some time at the end of a sprint race. Try the following questions to consolidate your understanding of oxygen debt: 20 What will happen if you lie down immediately after a 100-metre race? If you are running a 1,600-rn race, what strategy will you use? When should you carry out anaerobic respiration to secure maximum energy output for the body? What kind of training exercise will be useful for an athlete who is preparing for (i) a 100-rn race and (ii) a marathon? The oxygen content of atmospheric air and exhaled air can be compared by the glowing splint test. The glowing splint will be relighted only in the presence of high concentration of oxygen; it will NOT work with normal atmospheric air or exhaled air. A workable way is to compare the time of burning of a candle in the two air samples The End P.17