Ray F. Evert • Susan E.

Eichhorn

Raven Biology of Plants

Eighth Edition

CHAPTER 6
Respiration
Lecture presentation by Dr. Mohammad Brake
Jerash University

© 2013 W. H. Freeman and Company

CHAPTER OUTLINE
• An Overview of Glucose Oxidation
• Glycolysis
• The Aerobic Pathway
• Other Substrates for Respiration
• Anaerobic Pathways
• The Strategy of Energy Metabolism
ATP takes part in a great variety of cellular event like:
1. Biosynthesis of organic molecules.
2. Streaming of cytoplasm.
3. The active transport of molecules a cross the plasma membrane.
An overview of glucose oxidation:
Respiration: the complete oxidation of sugar or other molecules to carbon
dioxide and water.
CH2O → CO2 + H2O + energy
The energy storage molecules in plants are starch or sucrose.
Necessary steps for respiration:
•Hydrolysis of starch and sucrose molecules to glucose and fructose.
•Oxidation of glucose or fructose to carbon dioxide and water.
During the oxidation process:
1. Carbon atom in glucose loss electrons in the
form of H atom.
1. The Oxygen atom gain electrons in the form of H atom.
2. Free energy is released.
The oxidation of glucose could be:
a) In the presence of Oxygen (aerobic) and the overall reaction for the
complete oxidation is:
C6H12O6 (oxidized) + 6O2 → 6CO2 + 6H2O (reduced) + energy (686
kcal/mole)
b) In the absence of Oxygen (anaerobic):
The process called fermentation.
Yields less energy.
The main steps in respiration are:
Glycolysis:
Occurs in cytosol
The six carbon glucose is break down to a pair of three-carbon molecule
(pyruvate).
The pyruvate molecules are then oxidized to two molecules of acetyl CoA.
Small amount of ATP is synthesized from ADP + P.
Few electrons are transferred to coenzymes that function as electron carriers.
Citric acid cycle:
Occurs in mitochondria.
The acetyl CoA molecules are completely oxidized to carbon dioxide.
ATP is synthesized.
Electrons and proton are transferred to coenzyme.
Electron transport chain:
The coenzyme transfers the electrons to an electron transport chain, in which
the electrons drop (step by step) to lower level generating a proton gradient.
Consider amount of ATP is synthesized (oxidative phosphorlyation).
At the end of this step, electrons reunite with proton and combine with Oxygen
to form water.
Most of the energy produced from the oxidation of glucose is lost as heat
energy.
Glycolysis:
Is an anaerobic process that occurs in the cytosol.
Happened in all living cells from bacteria to plant and animal cells.
In glycolysis, the glucose split into two molecules of pyruvate and four H
atoms in a series of 10 reactions each is catalyzed by a specific enzyme.
In glycolysis, ATP, NADH will formed.
In glycolysis, 1 molecule of glucose is used.
The series of reactions in glycolysis are subdivided into two phases:
1) Preparatory phase: energy investment
From steps 1 to step 5.
2 ATP molecules are consumed, in step 1 and step 3.
The end product of this phase is 2 molecules of glyceraldehydes-3-
phosphate (3 carbon molecule).
2) Payoff phase:
From step 6 to step 10.
Two molecules of NADH formed in step 6.
In step 7 and 10, 4 molecules of ATP are formed.
The net result of ATP formed from glycolysis of one (1) molecule of glucose
equal 4-2= 2 ATP and 2 NADH.
Glycolysis ends with most of the energy of the original glucose
molecule still present in the two pyruvate molecules.
Glycolysis (from glucose to pyruvate) can be summarized by the overall
equation:
Glucose + 2 NAD+ + 2ADP + 2P → 2 Pyruvate + 2 NADH + 2 H+ + 2ATP +
2H2O
2 pyruvate molecules are still having 546 kcal/mole (80%) of the glucose
energy.
NADH formed during glycolysis can yield additional ATP molecules in the
mitochondria when used as electron donors to the electron transport chain
of the aerobic pathway.

The aerobic pathway:

Respiration composed of two main phases:
1. Glycolysis: anaerobic process happened in cytosol
2. Aerobic pathway: happened in mitochondria
•Citric acid cycle
•Electron transport chain
The mitochondria is an organelle surrounded by two membrane
1. Outer membrane
2. Inner membrane: inward folds to form cristae
3. The inner part of the mitochondria called matrix
The matrix is a dense solution containing enzymes, coenzymes, water,
phosphates, and other molecules involved in respiration.
Mitochondria resemble a self-contained chemical factory in which the outer
membrane allows the passage of small molecules freely, while the inner
membrane allows the passage of certain molecules such as pyruvate and ATP
and prevent the passage of others.
The enzymes of the citric acid cycle are found in:
•Most of them on the matrix
•Few are embedded in the inner membrane
The component for the electron transport chain are embedded in the inner
membrane
A preliminary step: pyruvate enters the mitochondria and is both
oxidized and decarboxylated.
The pyruvate produced by glycolysis passes from the cytosol to the matrix of
mitochondria
Before the starting of the citric acid cycle, the pyruvate molecules must be
oxidized and decarboxylated (electrons are removed and CO2 is split out of
the molecules). In this reaction 2 NADH (5 ATP) will form and 2 CO2 and 2
acetyl groups.
Acetyl group will attached to conenzyme A to from acetylcoenzyme A, which
will enter the citric cycle.
The citric acid cycle oxidized the acetyl groups of the acetyl CoA
molecules:
Citric acid cycle is known also as krebs cycle.
Citric acid cycle starts with the formation of citrate, which has three carboxylic
acid (-coo-) groups.
Citric acid cycle begins when acetyl group CoA combined with oxaloacetate
(4 C) to produce citrate (6 C).
The CoA is released to combine with a new acetyl group when another
molecule of pyruvate is oxidized.
The citric acide cycle use one acetyl group and regenerates one molecule of
oxaloacetate in 8 reactions.
The overall equation of citric acid cycle is:
Oxaloacetate + Acetyl Co A + 3H2O + ADP + P + 3NAD+ + FAD→
Oxaloacetate + 2CO2 + CoA + ATP + 3NADH + 3H+ + FADH2
The net energy released:
1. 1 ATP → 2X ATP = 2
2. 3 NADH → 2 X 3X2.5= 15
3. 1 FADH2 → 2X 1.5 = 3
In the electron transport chain, electrons removed from the glucose
molecule are transferred to oxygen
After the citric acid cycle, the glucose molecule is completely oxidized to
CO2.
Some of the energy was released during glycolysis and citric acid cycle and
most of the energy is still remains in NADH, and FADH2.
In the electron transport chain, the high-energy electrons of NADH and
FADH2 are passed from a high energy level to low energy level (O2).
The electron transport chain composed of a series of electrons carriers.
Most of the electron carriers in the transport electron chain are embedded in
the inner membrane of mitochondria.
Electron carriers are:
1) Cytochromes:
Protein molecules
Have iron-containing porphyrin, or heme group
Pick up electrons in the iron atoms
Each cytochrome differs in its protein structure
Carry single electron without protons
2) Iron-sulfur proteins:
Protein molecules
Have iron atoms attached to sulfur atoms
Pick up single electrons without protons on the iron atoms
3) Coenzyme Q (quinone):
Not associated with proteins
Small and hydrophobic molecules
Carry one or two electrons with protons (H+)
Co Q can shuttle protons across the inter mitochondrial membrane?
Co Q is embedded in the inner membrane
When Co Q pick up e-, it is also pick up proton (H+)
The proton (H+) will be released in the inter membrane space
Proton gradient will be generated a cross the inner mitochondrial membrane
This gradient is important for ATP production
At the beginning of the electron transports chain, the electrons held by NADH,
FADH2.
The electrons held by NADH are transferred to FMN carrier –and the
electrons held by FADH2 are transferred to Co Q.
When the electrons flow form high energy level to low energy level a big
amount of energy will be released and will be used to create proton gradient
across the inner membrane of mitochondria.
The proton gradient will drive the formation of ATP from ADP + P by oxidative
phosphorylation.
At the end of the chain, electrons are accepted by O2 and combine with proton
to form H2O.
Pair of electrons from NADH will generate 2.5 ATP
Pair of electrons from FADH2 will generate 1.5 ATP
Oxidative phosphorylation is achieved by chemiosmotic coupling
mechanism
Most of the components of the electron transport chain (complexes I-IV) are
embedded in the inner membrane of the mitochondria.
The protein complexes (I-IV) are proton pumps.
When the electrons pass from high energy level to lower one, the energy will be
released and used to pump protons (H+) from the matrix to the intermembrane
space through the protein complexes.
Each pair of electrons from NADH pumps 10 H+ protons.
The inner membrane of mitochondria is impermeable (protons that are pumped
into the inner membrane space cannot move back to the matrix →the result:
concentration gradient of protons)
Concentration gradient of protons means:
1. More H+ protons in inner membrane space than matrix
2. More + positive charge in the inter membrane space than matrix
Results: electrochemical gradient
When the protons back into the matrix through ATP synthase, the energy
released powers the synthesis of ATP from ADP+ P
ATP synthase:
•Large enzyme complex
•Embedded in the inner membrane of mitochondria
•Has binding sites for ADP+ P
The mechanism of ATP synthesis is known as chemiosmotic coupling.
Two distinct events take place in chemiosmotic coupling:
•Proton gradient is established a cross the inner membrane of the
mitochondria
•Potential energy stored in the gradient is used to generate ATP from ADP+ P
The overall energy harvest involves NADH and FADH2 as well as ATP
All but 2 of the 32molecules of ATP come from reactions in the mitochondria
All but 4 involve the oxidation of NADH and FADH2.
The total differences in free energy (∆G) between glucose and CO2 + H2O =
686 kcal/mole
The total energy stored in 32 ATP molecules = 233.6 kcal/mole
453kcal/mole (66%) is lost as a heat.
Other substrate for respiration: fats and proteins
Fats: the carboxyl end of the fatty acids will be removed as a cetyl CoA →
enter acid cycle
Protein: some residual carbon skeleton enters the citric acid cycle.
Anaerobic pathways:
In the absence of Oxygen, purvate is not the end product of glycolysis.
In many bacteria, fungi, protests , and animal cells, the end product of
glycolysis is lactate (3C, similar to pyurvate) and the process called lactate
fermentation.
In yeast and most plant cells, the end product of glycolysis is ethanol and the
process called the fermentation.
The complete equation of the fermentation of glucose is:
•Glucose + 2 ADP + 2 Pi → 2 ethanol + 2 CO2 + 2 ATP + 2 H2O
•Glucose + 2 ADP + 2 Pi → 2 lactate + 2 CO2 + 2 ATP + 2 H2O
The strategy of energy metabolism:
Metabolism: the totality of an organism’s chemical reactions consisting of
catabolism and anabolism.
Catabolic pathway: metabolic pathway that releases energy by breaking
down complex molecules to simpler compounds
Anabolic pathway: metabolic pathway that synthesizes a complex
molecule from simpler compounds
The large macromolecules like protein can be broken down to release
energy or serve as building units in biosynthesis.
Without building units (from the breakdown of macromolecule) the function
of the cell will stop.
eg: heterotrophic (root cells) depends on autotrophic cell (leaves cells) for
energy and building units molecules.
The end