A2 Biology Notes - PP & MS - Energy and Respiration - 2019

Code: A2B-Unit 12: Energy & Respiration
Mohammad Hussham Arshad, MD
Energy &
Respiration

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ADVANCED LEVEL
BIOLOGY 9700
UNIT 12: ENERGY AND
RESPIRATION

Guidance Notes
Biology Department

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SYLLABUS OUTLINE

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DEFINITIONS

Cellular Respiration refers to oxidation of food moleculesin a series of enzyme controlled metabolic
reactions to release energyin the form of ATP.

Dehydrogenation is a form of oxidation resulting in removal of H from organic substrates. The process is
catalysed by enzymes referred to as dehydrogenases.These enzymes are therefore essential for forming
NADH and FADH2.
Decarboxylation refers to removal of carbon dioxide/ carboxyl group from organic molecules.
Glycolysis -is defined as the oxidation of glucose to pyruvate.
Kreb’s Cycle- is a cyclic enzyme controlled process involving a series of decarboxylation and
dehydrogenation reactions with little production of ATP.
Substrate Level Phosphorylationis a type of metabolic reaction that results in the formation of
adenosine triphosphate (ATP) by the direct transfer and donation of a phosphate group to adenosine
diphosphate (ADP) from a phosphorylated reactive intermediate. It refers to formation of ATP from ADP
and Pi catalyzed by enzymes in reactions that do not depend on a proton-motive force. Glycolysis and
Krebs Cycle involves Substrate Level Phosphorylation.
Oxidative Phosphorylation is the enzymatic phosphorylation of ADP to ATP coupled to electron

transport chain in the mitochondria. Electrons derived from TCA cycle intermediates flow down a series
of electron-carrier proteins (i.e., cytochromes) until finally being transferred to molecular oxygen. It
refers to synthesis of ATP involving a membrane-associated electron-transport chain and the creation of
a proton-motive force. Also called electron-transport chain phosphorylation.
Chemiosmosis-- The process whereby a proton gradient is generated by electron transport and then
used to drive ATP synthesis by oxidative phosphorylation.
Respiratory quotient, RQ – the volume of carbon dioxide produced divided by the volume of oxygen
used during respiration.
RQ = CO2 produced/ O2 used

Oxygen Deficit-- defined as the difference in O2 volume between an ideal, hypothetical O2 uptake and
the actual uptake as it occurs in real life, for eg during exercise.
Oxygen Debt—Excess post exercise oxygen uptake to repay the oxygen deficit incurred during exercise
and restore the bodys’ metabolic state.

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WHY WE NEED ENERGY?

Humans need energy for carrying out activities of daily living. While some activities need more energy
than the rest, the exact mechanism of energy production stays the same. Following highlight just few of
the very important energy consuming activities.
• Active transport (ref. AS syllabus: against the [gradient]/ATP/Carrier proteins/shape change)

• Locomotion (eg of each in all five kingdoms)
• Transmission of nerve impulses
• Metabolic Reactions (Anabolism and Catabolism_know examples of each)
• Maintenance of a constant body temperature (homeostasis)
ADENOSINE TRIPHOSPHATE (ATP)

Most of the energy needed by the human body is derived from large organic substrates like
carbohydrates, fats and proteins. These molecules contain energy locked up in strong C-H bonds. Thus a
mechanism is required to release and convert this locked energy into readily usable form. The process
responsible for doing that is respiration, enabling provision of energy in the form of Adenosine
TriPhosphate or ATP.
Structure
ATP is derived from the nucleotide adenosine monophosphate (AMP) or adenylic acid, to which two
additional phosphate groups are attached through pyrophosphate bonds (~P). These two bonds are
energy rich in the sense that their hydrolysis yields a great deal more energy than a corresponding
covalent bond.Thus, it’s a phosphorylated nucleotide with energy rich pyrophosphate
(phosphoanhydride) bonds.

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ATP- Energy Currency
Adenosine triphosphate (ATP) is considered by biologists to be the energy currency of cells. It is the
high-energy molecule that carries the energy we need to do just about everything we do. It is present in
the cytoplasm and nucleoplasm of every cell, and essentially all the physiological mechanisms that
require energy for operation obtain it directly from the stored ATP.
Its utility as the best to carry out its function stems from various factors including:
• Small size
• Water soluble nature
• Rapid diffusion
• Phosphate groups are negatively charged so ATP doesn’t cross the membranes easily
• ATP ‘fits into’ many parts of the cell or different cellular proteins.
• Ease of hydrolysis to release energy- releases 30.5 kJ/ mol of bond hydrolysed
• Immediate energy donor
• Rapid turnover (ATP turnover is enormous. It is estimated that a resting human uses about 40 kg
of ATP in 24 hours, but contains only 5g of ATP at any one time. ATP is henceforth the immediate
energy donor getting hydrolysed so that ADP produced is used to regenerate more ATP).
The importance of ATP centers on the storage of about 7 kcal (30.6kJ)/mole of energy in the
phosphorus-oxygen bond between the first and second phosphate group. The relationship of energy and
the formation and hydrolysis of ATP is illustrated in the following equations: ADP = adenine
diphosphate.
a) Hydrolysis: ATP + H2O --> ADP + Pi + energy
b)Condensation: ADP + Pi+ energy --> ATP + H2O
Under certain conditions ATP may be hydrolyzed directly to AMP (adenine monophosphate).
ATP + H2O --> AMP + PP + energy
MITOCHONDRION

Mitochondria (singular – mitochondrion) are rod-shaped or kidney-shaped, membrane-enclosed
organelles, ranging in size from 1 to 10 micrometers, that are found in the cytoplasm of most eukaryotic
cells. Depending on organism, tissue type and level of cellular metabolic activity, a cellmay contain just
one mitochondrion or several thousand mitochondria; human cells normally contain 3000-5000
mitochondria.
Mitochondrion- Structure
The mitochondrion consists of four major sections – the outer membrane, the intermembranous space,
the inner membrane, and the matrix.

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The Outer Membrane
This membrane contains a great number of large transport proteins, which allows for large molecules to
enter with ease.
The Intermembrane Space
This space lies between the inner and the outer membrane and serves as the site where protons are
pumped via electron transfer chain to create a proton motive force essential for making ATP from ADP
through chemiosmosis.
The Inner Membrane
This membrane is highly convoluted, forming many folds called cristae. The inner membrane is highly
specialised to carry out its function in cellular respiration.
The Matrix
The Link reaction and Kreb Cycle takes place here. It also contains several copies of the mitochondrial
DNA genome, special mitochondrial ribosomes (70 S), tRNAs, and various enzymes required for the
expression of the mitochondrial gene.

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Mitochondrion- Functions
• Production of ATP via cellular respiration (see below)

• Programmed cell death (referred to as apoptosis)
• Monitoring cell differentiation, growth and development
• Cholesterol metabolism
• Detoxification of ammonia in urea cycle
• Steroid synthesis
Structural adaptations of mitochondrion for energy production
Structure Function Structural Adaptation

Intermembranous Space Proton motive force (pmf) Allows accumulation of H+
Inner Membrane- Features § Electron Transport Chain • Cristae maximises the
o Cristae (ETC) and surface area for ETC
o Selectively permeable § Oxidative • Being impermeable to H+
nature Phosphorylation helps in creating a
o Linearly arranged ETC proton pump gradient
o Presence of stalked • Linear arrangement of
particles ETC increases the overall
efficiency of the process
• Stalked particles for ATP
synthesis
Matrix § Link Reaction • Contains enzymes vital
§ Krebs Cycle for link reaction and
Krebs Cycle.

1. List three structural features that indicate the prokaryotic origin of mitochondria.
2. List two physiological features which indicate the prokaryotic origin of mitochondria.
3. Highlight the significance of mitochondrial DNA. (Attempt at the end of this chapter)

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Respiration

Respiration is of two types:
1. Aerobic
2. Anaerobic
Aerobic respiration involves the followingfour stages:
a) Glycolysis
b) Link Reaction
c) Kreb’s Cycle
d) Oxidative phosphorylation
Anaerobic respration is of two types:
a) Lactate fermentation (glycolysis followed by lactate formation)

b) Alcoholic fermentation (glycolysis followed by ethanol formation)
Glycolysis is the common pathway in aerobic and anaerobic respiration.
Dehydrogenation & Decarboxylation

The entire process of respiration involves multiple dehydrogenation and decarboxylation steps. Refer to
class lectures for details.

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CELLULAR RESPIRATION- AEROBIC
GLYCOLYSIS

Glycolysis is defined as the oxidation of glucose to pyruvate.
During glycolysis, glucose (C6) is broken down to two molecules of pyruvate (C3). Glycolysis occurs in the
cytoplasm (cytosol) and does not require oxygen. Cytoplasm contains the enzymes required for the
process of glycolysis.
Each step in the glycolytic pathway is catalyzed by a specific enzyme. A brief summary of these reactions
is presented here. For ease in remembrance and strict relevance to your course, I have divided the
process into THREE simple steps (the exact details are beyond the scope of your
course):(1)Phosphorylation, (2) Splitting/Lysis&(3) Dehydrogenation.
1. 2 ATP molecules are used to phosphorylate glucose that will eventually become converted to
pyruvate (or pyruvic acid) (see diagram below). The advantages of the initial phosphorylation
step are two-fold:
• Activates glucose to participate actively in the process (gains activation energy)

• Maintains a concentration gradient to enable glucose to continually enter the cell via facilitated
diffusion.
Explain why glucose needs to undergo the process of phosphorylation at the start of glycolysis?
• Phosphorylated glucose gets entrapped within the cell to maintain the concentration gradient
• Initial phosphorylation provides the activation energy needed for it to undergo splitting/lysis

Explain why this phosphorylated glucose does not diffuse out of the cell?
• Phosphorylated glucose cannot pass through the phospholipid bilayer;
• It’s too large to be transported via glucose protein channels;
• At times, it rapidly gets used up by the cell.
2. The phosphorylated 6C sugar undergoes lysis to produce two 3C sugar phosphate molecules.
3. Each of these 3C sugar phosphates undergoes a dehydrogenation reaction to produce NADH
from NAD+. The reaction also produces 2 ATP molecules for each 3C sugar phosphate
undergoing the reaction.Four ATP molecules are thus produced by substrate-level
phosphorylation.

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The overall Glycolysis step can be written as a net equation:
Glucose + 2ADP + 2NAD+---------> 2Pyruvate + 2ATP + 2NADH + 2H+
Yield of Glycolysis/ molecule of glucose:

• 2 molecules of pyruvate
• 2 molecules of ATP (net)
• 2 molecules of NADH

Glycolysis is the common pathway for aerobic and anaerobic respiration. In presence of oxygen and
mitochondrion, aerobic respiration occurs. The following reactions will ONLY occur if respiration is
AEROBIC:
• Link Reaction
• Krebs Cycle
• Electron Transport Chain & Oxidative Phosphorylation

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LINK REACTION

The pyruvate produced at the end of glycolysis enters the matrix of mitochondrionby active transport to
undergo a dehydrogenation and a decarboxylation step to form Acetyl- CoA.The enzyme complex
required for this process is known as the pyruvate dehydrogenase complex. It’s present in the matrix of
mitochondrion.
• The carbon lost is in the form of Carbon dioxide (CO2);

• The dehydrogenation step produces reduced NAD (NADH).
Please note that the link reaction occurs twice for one molecule of glucose because each glucose
molecule produces two pyruvate molecules.
Yield of Link Reaction/ molecule of glucose:

• 2 molecules of Acetyl CoA
• 2 molecules of CO2

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KREBS CYCLE (Tri Carboxylic Acid Cycle; Citric Acid cycle)

The Acetyl CoA molecules produced in the link reaction enter what we call the Krebs Cycle. It occurs in
the matrix of the mitochondrion mainly because the enzymes needed for the process are present
there.The process involves a series of decarboxylation and dehydrogenation reactions with little
production of ATP.
A brief simplified version of the cycle is highlighted and shown below:
• Initially the Acetyl CoA (2C) combines with oxaloacetate (4C) to produce citric acid (6C)
• The citric acid undergoes a decarboxylation and a dehydrogenation reaction to produce alpha-
ketoglutarate (5C).
• Alpha- Ketoglutarate then undergoes further decarboxylation and dehydrogenation steps to
reproduce oxaloacetate to complete the cycle.
• The energy released from these dehydrogenation reactions is used to reduce the electron
carriers NAD+ and FAD+ to produce NADH + H+ and FADH2, and to produce ATP by substrate-
level phosphorylation.
The cycle produces/ molecule of glucose (2 x Acetyl CoA):

• 4 molecules of CO2
• 2 molecules of ATP

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• 2 molecules of FADH2

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ELECTRON TRANSPORT CHAIN & OXIDATIVE PHOPHORYLATION

The process extracts energy from NADH and FADH2 in order to add a phosphate group to ADP to make
ATP. It occurs in mitochondrialcristae. Oxidative Phosphorylation requires NADH or FADH2, ADP, O2 and
the Electron Transport Chain. The Electron Transport Chain is a collection of proteins, embedded in the
inner membrane. It is used to transport the electrons from NADH and FADH2.
The final stage of aerobic cellular respiration is oxidative phosphorylation. The key event of this stage is
the formation of a proton gradient that drives the synthesis of ATP. Here’s a summary of how it happens
followed by more details below:
Summary
• In the first step, an electron carrier (such as NADH + H+) donates high energy electrons to the
first protein in the chain (complex I), resulting in the protein becoming reduced.
• This is followed by the sequential oxidation and reduction of a “chain” of proteins and other
molecules that comprise the electron transport chain (ETC).
• The high energy electrons transported via ETC lose energy in these sequential redox reactions.
• This energy is tapped to transfer protons from the matrix across the inner membrane to inter-
membranous space (IMS) creating an electrochemical gradient that generates a proton motive
force.
• The protons within the IMS travel down their electrochemical gradient via ‘stalked particles’
which contains the enzyme ATPase. As these protons travel via these stalked particles, ADP and
Pi combine to form ATP.
• Oxygen acts as the final electron acceptor, and hence the formation of ATP via this process is
referred to as oxidative phosphorylation.

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Detailed Account:
The electron transport chain consists of several proteins and other compounds that are associated with
the inner membrane of mitochondria. As mentioned in the first step, the electron carrier NADH donates
high energy electrons to the first protein in the chain – the NADH dehydrogenase complex (complex I).
Once NADH dehydrogenase complex is reduced (by gaining electrons), it can in turn become oxidized by
reducing the next component of the chain which is ubiquinone (Coenzyme Q).Ubiquinone then becomes
oxidized by reducing the next component of the chain, and so on. The high energy electrons lose energy
as they move from one electron carrier to the next. Note that each component of the chain undergoes a
cycle of being reduced and then oxidized as it gains and then donates electrons. At the end of the
electron transport chain, the electrons are finally used by cytochrome oxidase (complex IV) to reduce
the “terminal electron acceptor”, which in aerobic metabolism is oxygen. The reduction of oxygen yields
water - the final product of this process.
At this point we have finally found where oxygen takes part in aerobic cellular respiration, or, in other
words, why aerobic cellular respiration is ‘aerobic.’ In fact, this is the only step in aerobic cellular
respiration that directly requires molecular oxygen. If oxygen is not present, however, oxidative
phosphorylation, the citric acid cycle, and link reaction are not able to occur.
The sequence of reduction and oxidation reactions allows energy to be gradually released, and it is
productively used to transport protons (H+) across the membrane. The transport of protons requires
energy because it is accompanied by the formation of anelectrochemical gradient. An electrochemical
gradient is a form of potential energy – remember this energy, we’ll see how it is used to produce ATP in
just a moment.
As you recall, in the citric acid cycle FAD+ is reduced to form FADH2. The electrons carried by FADH2
enter the electron transport chain at ubiquinone via the succinate dehydrogenase complex (complex II).
From this point on, the electrons follow the same path as with NADH and are ultimately used to reduce
oxygen. Two other important features of electron transport are notable.First, notice that each time a
redox reaction occurs, the amount of free energy present decreases. In other words, each time an
electron is transferred between carriers, a small amount of energy is released. This, again, is the energy
that is used to build a proton gradient across the membrane. Second, it is important to notice that all

components of electron transport chains share a common feature - they are all capable of undergoing
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oxidation and reduction.


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As a result, they cycle from being oxidized, to being reduced, to being oxidized, and so on. This has
several important implications, one of which is that the NADH produced during metabolism of glucose
is continually being re-oxidized to form NAD+ that can cycle back to participate in glycolysis, pyruvate
oxidation, and the citric acid cycle.
Like many other membranes in cells, the inner mitochondrial membrane is highly selective and H+
cannot readily cross it (as mentioned earlier). This is a very important feature since it provides a way to
form a H+ gradient. During electron transport, protons are pumped across the membrane via the
electron transport chain, forming just such a gradient.
Given a chance, protons will diffuse back across the membrane toward equilibrium. In order for H+ to
move back across the membrane, however, they must pass through special channels within the ATP
synthase complex. The passage of H+ through the channels is “driven” by the electrochemical gradient
that exists due to the difference in the H+ concentrations on either side of the membrane.
This electrochemical gradient, which is also called the proton motive force, provides the energy needed
by ATP synthase to synthesize ATP from ADP and inorganic phosphate. As protons flow down their
electrochemical gradient and through ATP synthase, they cause part of the synthase complex to rotate.
The energy thus provided is used to form ATP from ADP and inorganic phosphate.This phenomenon is
also referred to as the chemiosmotic hypothesis of energy production.
One molecule of NADH on complete oxidation provides sufficient energy to produce 3 ATP molecules via
ETC. Additionally, one molecule of FADH2 on oxidation provides enough energy to form 2 ATP molecules.
Outlining Oxidative Phosphorylation_Highlight

The processes that entail the Electron Transport Chain includes:
• Regeneration of oxidized NAD from reduced NADH, releasing H
• H from NAD is split to produce protons and electrons
• The electrons travel down a chain of carriers to attain a lower energy state
• Oxygen acts as the final electron acceptor
• Protons + Oxygen + electrons produce water as the waste product
Chemiosmosis involves:
• Transfer of electrons from one carrier to the next releasing energy
• The energy released is used to pump the hydrogen ions from the matrix into the intermembranous space
creating a proton motive force
• These hydrogen ions move back into the matrix down their electrochemical gradient via ATP synthases
• Synthesizing ATP from ADP and Pi

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Yield of Electron Transport Chain & Oxidative Phosphorylation

• 10 NADH---------(10 x 3)=30 ATP
• 2 FADH-----------(2 x 2)= 4 ATP

+
What is the role of NAD in the entire process?
• Coenzyme
• Hydrogen and electron carrier
• Carries protons and electrons produced in glycolysis, link reaction and Krebs cycle;
• To Electron Transport Chain (ETC); and
• Gets oxidized in the electron transport chain (ETC) indirectly providing energy for ATP synthesis
• Oxidizes triose phosphate to pyruvate in glycolysis
What will happen if O2 is absent?

• No Oxygen present to accept protons and electron
• Proton motive force does not develop
• Oxidative phosphorylation does not occur
• NADH cannot be oxidized and reused
• Krebs cycle and Link reaction stops

TEMMEEEE WHY?
We discussed previously that we produce 3 ATP molecules per NADH molecule and 2 ATP molecules per
FADH2 molecule.
However, in many physiological systems like ours, that is NOT exactly the case. Effective production of
number of ATP molecules in our body is as follows:
Yield of Electron Transport Chain & Oxidative Phosphorylation

• 1 NADH---------2.5ATP , therefore 10NADH will produce 25 ATP
• 2 FADH----------1.5 ATP, therefore 2FADH2 will produce 03 ATP

Can you figure out why this happens?
Respiratory Inhibitors
Many inhibitors block the ETC and hence inhibit the process of oxidative phosphorylation and Krebs
Cycle. Consequently, the link reaction stops. Oxygen is no longer consumed because of lack of electrons
needed to reduce oxygen to produce water in the final stages of respiration. Common inhibitors include:
• Cyanide
• Sodium azide
Read up more on inhibitors and specifically uncouplers J

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SUMMARY OF CELLULAR AEROBIC RESPIRATION

Enlist three differences between substrate level and oxidative phosphorylation.

Substrate Level Phosphorylation Oxidative Phosphorylation

Stage Site NADH/molecule FADH2/molecule ATP/ CO2/

of glucose of glucose molecule of molecule of
glucose glucose
Glycolysis Cytoplasm 2 - 2 -
Link Reaction Matrix of 2 - - 2
Mitochondrion
Krebs Cycle Matrix of 6 2 2 4

Mitochondrion
ETC & OP Cristae of - - 34 -

Mitochondrion NADH: 30
FADH2: 4

Total - 10 2 38 6
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CELLULAR RESPIRATION- ANAEROBIC
ANAEROBIC RESPIRATION

The term anaerobic means without air and hence anaerobic respiration refers to the special type of
respiration, which takes place without oxygen. Anaerobic respiration is the process of oxidation of
molecules in the absence of oxygen, which results in production of energy in the form of ATP. The
process occurs differently inyeast and mammals. Glycolysis, however,is the common pathway in the two
leading to production of pyruvate. It’s also the ONLY energy producing stage in anaerobic respiration
leading to net production of 2 ATP molecules per molecule of glucose.
Anaerobic respiration occurs in the cytoplasm. There are two common forms of anaerobic respiration:
• Lactate fermentation occurs in mammals. Each pyruvate is converted to lactate and one NADH
is used.
• Alcoholic fermentation occurs in yeast, and certain bacteria. Each pyruvate is converted to a
molecule of ethanol and one NADH is used in the reaction.
The purpose of both fermentation processes is to free NADH for use in glycolysis.
Lactate fermentation
In periods of strenuous activity, anaerobic respiration operates in additionto aerobic respiration. It is

NOT an alternative to aerobic respiration, but an adjunct to meet the high energy demand when the
oxygen supply isn’t enough to support aerobic respiration. In mammals, this is coupled with
productionof lactic acid in muscles. Lactic Acid is a toxin and causes fatigue, soreness and stiffness in
muscles.No ATP is produced in converting pyruvate to lactate. Instead, the NADH produced
duringglycolysis is used to reduce pyruvate to lactic acid. Most of the energy is locked up in lactate.

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NAD+ is regenerated in the process. The enzyme used in the process is lactate dehydrogenase.

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MAMMALS: Glucose (Broken down to) → 2 Lactic acid + 2 ATP + 2 NAD+
With provision of oxygen, this lactate can be reconverted to pyruvate in the liver cells (heart muscle may
also do it). Pyruvate can then enter mitochondrion to participate in the link reaction.
Alcoholic Fermentation
Alcoholic fermentation occurs in fungi, bacteria and a few plants (yes!). Again, glycolysis converts
glucose to pyruvate producing 2 molecules of ATP and 2 molecules of NADH. Pyruvategets reduced and
decarboxylated to form ethanal (acetaldehyde) and carbon dioxide. Ethanal is reduced using NADH to
form ethanol. NAD+ is regenerated in the process. Most of the energy is therefore locked up in ethanol
molecules. The enzymes required for the process are pyruvate decarboxylase (Step 1: Pyruvate to
Acetaldehyde) and alcohol dehydrogenase (Step 2: Acetaldehyde to Ethanol).
YEAST: Glucose (Broken down to) →2 ATP + 2 Ethanol + 2 CO2 + 2 NAD+
Unlike lactic acid fermentation, alcoholic fermentation is an irreversible process. Ethanol cannot be

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reconverted back to pyruvate.


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Few important factors affecting the rate of anaerobic respiration in yeast:
• Temperature…….find out how?

• Glucose concentration…….find out how?
• Yeast concentration…….find out how?
In everyday life, alcohol industry and baking industry use this process for commercial purposes.
How is lactate fermentation (LF) different from alcoholic fermentation (AF)?

• No decarboxylation in LF
• Reversible when oxygen is available
• Single step reaction
• Enzyme involved is lactate dehydrogenase as compared to alcohol dehydrogenase in AF

What is the role of NADH in Anaerobic Respiration?
• Reduction of pyruvate to lactate/ pyruvate to ethanal and consequently ethanol
• NAD+ is regenerated
• Its regeneration enables glycolysis to continue

Summary of Reactions of Lactate Fermentation and Alcoholic Fermentation

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AEROBIC VS ANAEROBIC RESPIRATION

• Anaerobic respiration produces considerably less amount of energy when compared with
aerobic respiration (2 vs 38 ATP). The reason is that most of the energy is locked in molecules
like lactic acid or ethanol. Thus, more glucose needs to be broken down to produce the same
amount of energy----depletes glycogen reserves.
• Anaerobic respiration is 2.5 times faster than aerobic respiration.
• Both ethanol and lactate are toxic products produced from anaerobic respiration. Ethanol kills
yeast at higher concentration (reason why wines and beers do not have alcohol concentration
beyond 13-15%). Lactate causes muscle stiffness, soreness and cramps. Aerobic respiration
produces carbon dioxide and water.

EXERCISE AND OXYGEN DEBT

During muscular exercise, blood vessels in muscles dilate and blood flow is increased in order to
increase the available oxygen supply. Up to a point, the available oxygen is sufficient to meet the energy
needs of the body. However, when muscular exertion is intense, oxygen cannot be supplied to muscle
fibers fast enough, and the aerobic breakdown of pyruvic acid cannot produce all the ATP required for
further muscle contraction. The body therefore develops an oxygen deficit due to real oxygen uptake
being less than the ideal required one. ATP production therefore needs to be supported by other
mechanisms to meet the energy needs. These include:
• Anaerobic respiration forming lactic acid (muscle fatigue and depletion of glycogen reserves)
• Oxygen released from oxymyoglobin in muscles
• Creatine phosphate in muscles (serves as a source of phosphate to convert ADP into ATP)

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After a strenuous exercise (like the one I do every day to keep myself fit….lols!), there are four tasks that
need to be completed:
• Removal of lactic acid (link to Cori Cycle)

• Replenishment of myoglobin with oxygen
• Replenishment of glycogen and ATP production
• Replenishment of phosphocreatine (PC) inside muscles
The need for oxygen to replenish ATP and remove lactic acid is referred to as the "Oxygen Debt" or
"Excess Post-exercise Oxygen Consumption" (EPOC) - the total oxygen consumed after exercise in excess
of a pre-exercise baseline level. This is required to support increased metabolic rate (including increased
heart rate, breathing rate and body temperature) following exercise. The following chart summarises
the need to have EPOC.
How to measure Oxygen Debt (EPOC)?

• Oxygen consumption to be measured as: Oxygen inhaled – Oxygen exhaled
• Oxygen debt is the excess post exercise oxygen consumption
• Measure oxygen consumption at rest (a) and after exercise (b)

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• Oxygen debt will therefore be = a - b


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RESPIRATORY SUBSTRATES- Relative Energy Values

Carbohydrates, fats and proteins all serve as respiratory substrates. All of them may be used to produce
ATP from respiration. The amount of ATP they produce however varies for a fixed mass of the substrate.
This difference in energy production results due to differences in their organic structures. Carbohydrates
and proteins release an almost similar amount of energy on oxidation of a fixed mass. Lipids however,
have greater number of C-H bonds and therefore produce more energy.
• The more hydrogens (and C-H bonds) there are in an organic molecule, the more NADH and
FADH2 molecules formed in glycolysis, link reaction and Krebs cycle.
• More ATP is therefore produced in the electron transport chain via oxidative phosphorylation
• Some molecules have more hydrogens than others
• If there are more hydrogen atoms per mole (fixed amount) of substrate, more oxygen is needed
as the final acceptor
• Proteins and carbohydrates contain similar ratios of C, H and O, but lipids containless O than C
and H, so lipid yields more energy.
• Carbohydrates and proteins, therefore, have lower energy density(less energy per mole)when
compared with lipids.
Respiratory Substrate Mean Energy Value (kJ/g)

Carbohydrate 15.8
Protein 17.0
Lipid 39.4

RESPIRATORY QUOTIENT (RQ)

The volume of carbon dioxide produced divided by the volume of oxygen used during respiration is
referred to as the RQ.
RQ = CO2 produced/ O2 used
It can be found by experimentation, using a respirometer with and without soda-lime to absorb the
carbon dioxide.
e.g. for a carbohydrate: C6H12O6 + 6O2 = 6CO2 + 6H2O RQ = 6/6 = 1
e.g. for a lipid: 2C57H110O6 + 163O2 = 114CO2 + 110H2O RQ = 114/163 = 0.7
General Equation for balancing oxidation of organic compounds with C, H & O:

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CxHyOz + (x + y/4 - z/2) O2 ---> x CO2 + (y/2) H2O


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RQ is a good way of predicting what respiratory substrates are being used for respiration. Substrates
with more H atoms in their structure will need more Oxygen as the final proton and electron acceptor
and hence will have lower RQs. Proteins and fats if used exclusively for respiration will therefore have
lower RQ values compared to RQ obtained from oxidation of carbohydrates exclusively.
RQ values are greater than 1.0 when:
• Anaerobic respiration is occuring in addition to aerobic; or

• Organic acids are being oxidised
RQ values of germinating seeds

Time RQ Explanation
Seeds soaked in 5.5-7.5 With little dissolved oxygen in water, anaerobic respiration
water occurs.
After 14 hours in soil 0.8 As oxygen becomes available, the amount of aerobic
respiration increases. A mixture of lipds and carbohydrates
from the stores in the seed is being respired. The conversion of
stored lipids to carbohydrate is also taking place.
After 14 days 1.0 Aerobic respiration occurs. The leaves have emerged and
photosynthesis is producing carbohydrates, which are being
respired.

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Adaptations of Rice for wet environment
UNIT 12: PRACTICAL TASKS

• Use of respirometers to find out RQ and the effect of temperature on respiratory
rate(reference below).
• Carrying out an investigation into the activity of dehydrogenase enzymes during respiration,
using DCPIP
• Practical work using tetrazolium chloride (TTC) as an artificial hydrogen acceptor, to illustrate
the activity of dehydrogenase enzymes during respiration.
• Investigate the effect of temperature, glucose concentration and ethanol concentration on the

rate of anaerobic respiration in yeast

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The Respirometer, Rates of Respiration and Respiratory Quotient (RQ)

Respirometers are generally used to measure oxygen consumption rates in small invertebrates,
germinating seeds and respiring yeast. It works on the principle that CO2 produced by respiration is
absorbed by a strong alkali such as KOH. This results in a decrease in volume in the vessel containing the
respiring organism and thus a decrease in pressure. This decrease in pressure causes the manometer
fluid to move. It is important that there are no fluctuations in temperature and pressure as these will
affect the level of the manometer fluid. The control tube has an equal volume and also contains an inert
material such as glass beads equal to the volume of the respiring organisms and is used to help
compensate for changes in atmospheric pressure. The respirometer can be used to measure rates of
respiration and respiratory quotients.

Fig. 1.0 A Respirometer.

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Measuring the effects of temperature on the rate of reaction
As outlined above, the respirometer can be used to measure rates of respiration. Look at the data below
and then answer the questions that follow.
The experiment involved investigating the effects of temperature on the rate of respiration in woodlice.
The change in O2 volume was measured over 3 minutes.
1. Which tube is the control (compensation) tube: A or B?

2. Which gas does the potassium hydroxide absorb and why is the filter paper added to each
tube?
3. What is missing from tube B?
4. Indicate using an arrow which way you would expect the manometer fluid to move during
the experiment.
5. At the end of each repeat of each experiment what is the syringe connected to tube A used
for?
6. Complete the table of data below and plot temperature vs. average rate of oxygen
consumption.
7. What part of the respiratory pathway is this measuring the rate of?
8. Using the results of this experiment, explain HOW does temperature affect the rate of
respiration? (link it to AS Bio)

Temperature Start point of Finish point Change in Rate of oxygen Average rate of oxygen
(°C) manometer of volume comsumption comsumption
fluid (ml) manometer (ml) -1 -1
fluid (ml) (ml O2 min ) (ml O2 min )
5 15.00 16.70
5 17.00 19.00
5 14.00 16.30
10 12.00 15.00
10 16.00 19.20
10 17.00 20.00
20 18.00 24.30
20 13.00 19.20
20 15.00 21.00
40 16.00 27.10

40 12.00 22.30
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40 19.00 30.00
60 16.00 16.80
60 14.00 14.10
60 15.00 15.30

Using respirometers to determine Respiratory Quotient (RQ)

This apparatus can also be used to determine respiratory quotients. First oxygen consumption at a
particular temperature is found (x ml O2 min-1). Then the respirometer is set up with the same organism
at the same temperature but with no CO2 absorbing chemical. The manometer will show whether the
volumes of oxygen and carbon dioxide are the same.
ü When the volumes are the same the level of the manometer fluid will stay the same.
ü Or the level of the manometer fluid may show an increase in the volume of gas by y ml O2 min-1.
The RQ can be calculated by the equation:
RQ = CO2/O2 = (x + y)/x
ü Or the manometer fluid may show a decrease in the volume of gas by z ml O2 min-1.Then the RQ
would be:

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RQ = CO2/O2 = (x - z)/x

Page59

Questions.
1. Label on the respirometer which way you would expect the manometer fluid to move if the
organisms were respiring anaerobically when there was and was no CO2 absorbing substance in
both tubes A and B.
2. Calculate these Respiratory quotients from the data in the table below.

Temperature Rate of oxygen Rate of carbon Respiratory Substrate being

(°) consumption dioxide quotient respired/Type
consumption of respiration
(ml O2 min-1)
(ml CO2 min-1)
W 30 2.55 3.21
X 30 2.63 2.65
Y 30 2.23 1.55
Z 30 2.68 2.47
Sketch a Respirometer here and assign direction for organisms W, X, Y & Z:
ACTIVITY
The type of respirometer mentioned above is a manometer based respirometer. Another type
commonly used in school labs is a syringe based respirometer. Look it up for class discussion!!!

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APPENDIX (just for personal reference- you won’t be asked about these structures in Exams)

Structure of NAD+
Nicotinamide Adenine Dinucleotide is composed of two nucleotide molecules: Adenosine monophosphate

(adenine plus ribose-phosphate) and nicotinamide ribotide (nicotinamide plus ribose-phosphate).
Nicotinamide (derivative of vitamin B3) is the active part of the molecule where the reversible oxidation
and reduction takes place. The oxidized form of NAD has one hydrogen atom less than the reduced form
and, in addition, has a positive charge on the nitrogen atom which allows it to accept a second electron
upon reduction. The correct way to symbolize the reaction is: NAD+ + 2H----->NADH + H+.
Structure of FAD+
The structure shown is for FAD and is similar to NAD+ in that it contains a vitamin-riboflavin (vitamin B2),
adenine, ribose, and phosphates. As shown it is the diphosphate, but is also used as the monophosphate
(FMN).

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Coenzyme Q- Co Q (Ubiquinone)
This oil-soluble, vitamin-like substance (a form of lipid) is present in most eukaryotic cells, primarily in the
mitochondria. All the natural forms of CoQ are insoluble in water, but soluble in membrane lipids where
they function as a mobile electron carrier in the electron transport chain. The long hydrocarbon chain
gives the non-polar property to the molecule. CoQ acts as a bridge between enzyme complex 1 and 3 or
between complex 2 and 3.
Cytochromes
• Are proteins
• One class of such proteins is present in the cristae of mitochondrion forming an ETC
• All contain haem or haem-like prosthetic group
• Carries electrons
• cytochromes b, c1, c, a, a3 relay electrons,one at a time in this order
• Complex III = cytochrome b + cytochrome c1
• Cytochrome c = mobile electron carrier to transfer electrons from complex III to complex IV
• Complex IV = cytochrome a + cytochrome a3 = cytochrome oxidase

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Energy &
Respiration

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With

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ADVANCED LEVEL
BIOLOGY 9700

CIE PAST PAPERS: S02-N18/P04

Pre U PAST PAPERS: S11-S13

UNIT 12:
ENERGY AND RESPIRATION

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Page70
Q1.

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Q2.

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Q3.

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Q4.

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Q5.

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Q6.

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Q7.

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Q8.

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Q9.

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Q10.

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Page88

Q11.

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Q12.

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Page92

Q13.

Page93

Q14.

Q15.

Page94
Q16 (J10/41/q7)

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Page96

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Q17 (J10/43/q7)

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Page99
Q18 (N10/41/q10)

Q19 (N10/43/q10)

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Q20 (J11/41/q7)

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Page103

Page104

Q21 (J11/43/q6)

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Page106

Q22 (N11/41/q6)

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Page108

Q23 (N11/43/q6)

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Q24 (J12/41/q9)
Q25 (J12/42/q9)

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Q26 (N12/41/q8)

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Page112

Page113

Q27 (N12/43/q8)

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Page115

Q28 (J13/42/q4:a)

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Q29 (J13/43/q4)

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Q30 (N13/41/q3)

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Q31 (N13/43/q3)

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Q32 (J14/P41/Q8)

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Q33 (J14/P42/Q8b)

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Q34 (N14/P41/Q5)

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Q35 (N14/P43/Q4)

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Q36 (J02/P4/Q2)

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Q37 (J15/41/q7)

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Q38 (J15/42/q7)

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Q39 (N15/41/q1)

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Q40 (N15/43/q1)

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Q41 (J16/41/q1)

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Q42 (J16/42/q1)

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Q43 (N16/42/q8)

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Q44 (N16/43/q8)

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Q45 (J17/41/q4)

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Q46 (J17/42/q1)

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Page157

Q47 (N17/42/q7)

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Page159

Q48 (N17/42/q9)

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Q49 (J18/41/q6)

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Q50 (J18/42/q6)

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Page164

Q51 (N18/41/q6)

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Q52 (N18/42/q7)

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P1 (J11/P3/Q5)

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P2 (J11/P4/Q3)

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P3 (J12/P2/Q4)

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P4 (y16/SP_1/Q21a)
P5 (y16/SP_3/Q5) {Attempt after Unit M: Photosyntheis}

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ADVANCED LEVEL
BIOLOGY 9700

MARKING SCHEME

UNIT 12

ENERGY AND RESPIRATION

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Q1.

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Q2.

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Q3.

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Q4.
Q5.

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Q6.
Q7.

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Q8.

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Q9.

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Q10.
Q11.

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Q12.
Q13.

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Q14.

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Q15.

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Q16.

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Q17.

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Q18.

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Q19.

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Q20.

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Q21.
Q22.

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Q23.

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Q24.

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Q25.

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Q26.

Page200

Q27.

Q28.

Page201

Q29.

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Q30 (N13/41/q3)

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Q31 (N13/43/q3)

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Q32

Q33

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Q34

Q35

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Q36 (J02/P4/Q2)

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Q37

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Q38

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Q39

Q40

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Q41

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Q42

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Q43

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Q44

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Q45

Q46

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Q47

Q48

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Q49

Q50

Q51

Q52

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P1

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P2

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P3

P4

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P5

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A2 Biology Notes - PP & MS - Energy and Respiration - 2019

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Code: A2B-Unit 12: Energy & Respiration

Mohammad Hussham Arshad, MD

Mohammad Hussham Arshad, MD

Glycolysis -is defined as the oxidation of glucose to pyruvate.

Oxidative Phosphorylation is the enzymatic phosphorylation of ADP to ATP coupled to electron

RQ = CO2 produced/ O2 used

WHY WE NEED ENERGY?

• Active transport (ref. AS syllabus: against the [gradient]/ATP/Carrier proteins/shape change)

ADENOSINE TRIPHOSPHATE (ATP)

ATP- Energy Currency

b)Condensation: ADP + Pi+ energy --> ATP + H2O

ATP + H2O --> AMP + PP + energy

The Outer Membrane

The Intermembrane Space

The Inner Membrane

• Production of ATP via cellular respiration (see below)

Structural adaptations of mitochondrion for energy production

Structure Function Structural Adaptation

Aerobic respiration involves the followingfour stages:

Anaerobic respration is of two types:

a) Lactate fermentation (glycolysis followed by lactate formation)

Glycolysis is the common pathway in aerobic and anaerobic respiration.

Dehydrogenation & Decarboxylation

CELLULAR RESPIRATION- AEROBIC

• Activates glucose to participate actively in the process (gains activation energy)

The overall Glycolysis step can be written as a net equation:

Glucose + 2ADP + 2NAD+---------> 2Pyruvate + 2ATP + 2NADH + 2H+

Yield of Glycolysis/ molecule of glucose:

• The carbon lost is in the form of Carbon dioxide (CO2);

Yield of Link Reaction/ molecule of glucose:

KREBS CYCLE (Tri Carboxylic Acid Cycle; Citric Acid cycle)

A brief simplified version of the cycle is highlighted and shown below:

The cycle produces/ molecule of glucose (2 x Acetyl CoA):

ELECTRON TRANSPORT CHAIN & OXIDATIVE PHOPHORYLATION

oxidation and reduction.

Outlining Oxidative Phosphorylation_Highlight

Yield of Electron Transport Chain & Oxidative Phosphorylation

What will happen if O2 is absent?

Yield of Electron Transport Chain & Oxidative Phosphorylation

Read up more on inhibitors and specifically uncouplers J

SUMMARY OF CELLULAR AEROBIC RESPIRATION

Enlist three differences between substrate level and oxidative phosphorylation.

Stage Site NADH/molecule FADH2/molecule ATP/ CO2/

Krebs Cycle Matrix of 6 2 2 4

ETC & OP Cristae of - - 34 -

CELLULAR RESPIRATION- ANAEROBIC

In periods of strenuous activity, anaerobic respiration operates in additionto aerobic respiration. It is

MAMMALS: Glucose (Broken down to) → 2 Lactic acid + 2 ATP + 2 NAD+

YEAST: Glucose (Broken down to) →2 ATP + 2 Ethanol + 2 CO2 + 2 NAD+

reconverted back to pyruvate.

Few important factors affecting the rate of anaerobic respiration in yeast:

• Temperature…….find out how?

How is lactate fermentation (LF) different from alcoholic fermentation (AF)?

AEROBIC VS ANAEROBIC RESPIRATION

EXERCISE AND OXYGEN DEBT

• Removal of lactic acid (link to Cori Cycle)

How to measure Oxygen Debt (EPOC)?

• Oxygen debt will therefore be = a - b

RESPIRATORY SUBSTRATES- Relative Energy Values

• Some molecules have more hydrogens than others

Respiratory Substrate Mean Energy Value (kJ/g)

RESPIRATORY QUOTIENT (RQ)

RQ = CO2 produced/ O2 used