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All cells need energy to stay alive and maintain an ordered cellular environment.
Cell growth, development and reproduction all require energy. Movement of
materials through membranes often requires energy, as do intracellular
movements.
Cells obtain the energy to do work by oxidizing organic molecules, a process called
cellular respiration*. Although many organic molecules can be oxidized,
glucose, the main product of photosynthesis, is the primary f uel molecule for the
cells of living organisms. Cell respiration pathways are catabolic – the end
products have less energy than the reactants. Some of the energy released during
cell respiration is heat energy; the rest is used to make molecules of A TP.
All organisms, autotrophs and heterotrophs, must do cell respiration. Recall that
organisms that do photosynthesis (or properly, manufacture their own fuel
molecules) are called a utotrophs. H eterotrophs obtain their fuel molecules
"pre-formed" by other organisms. Animals, fungi, many protists and many
bacteria are heterotrophs. Plants and some protists are autotrophs, as are some
bacteria, in particular, the cyanobacteria.
The metabolic pathways of cell respiration vary depending on the type of organism,
the enzymes the organism has, oxygen use, and what the final product molecule in
the cell respiration process is. We will focus on the metabolism of glucose in cell
respiration, but we shall also discuss how alternative fuel molecules fit into the cell
respiration pathways.
In complete aerobic respiration, glucose is broken down into water and carbon
dioxide. This process requires oxygen.
Not all cell respiration is aerobic. Organisms that do cell respiration without oxygen
are said to be anaerobic. Fuel molecules can be oxidized without oxygen to yield
smaller amounts of ATP. The f ermentations involve the partial breakdown of
glucose without using oxygen. Many prokaryotes have a variety of fermentation
pathways, using a number of different fuel molecules. The final electron acceptor
for the fermentations is an o rganic molecule.
Glycolysis
The initial stage of glucose metabolism, or cell respiration, is a process called
glycolysis, which splits a glucose molecule into two molecules of pyruvate, a
3-carbon compound. Glycolysis occurs in the cytosol of the cell.
The final stage of aerobic respiration is the e lectron transport chain and the
chemiosmotic synthesis of ATP. Since the energy to synthesize ATP is from the
oxidation-reduction reactions, such synthesis is called o xidative
phosphorylation.
• Oxygen is the final electron acceptor for the oxidation-reductions that start
with NADH in the electron transport system.
• The electron transport system takes place in the inner membrane of the
mitochondria.
• When oxygen is available, as much as 38 ATP can be generated from one
glucose molecule
Glycolysis
Cell Respiration - 6
Summary of Glycolysis
Inputs
Glucose
2 ATP*
2 NA D+
2 ADP + 2Pi
Outputs
2 Pyruvate
2 NADH
4 ATP*
* Therefore the
net energy yield
is 2 ATP
Notes:
• The ATP generated is by s ubstrate-level phosphorylation
After Glycolysis
Pyruvate is a critical intermediate in the cellular respiratory pathways. As stated,
virtually all organisms do glycolysis. The respiratory pathways diverge at
pyruvate. If oxygen is available, and the organism has the appropriate enzymes,
pyruvate is oxidized and follows the aerobic respiratory pathway. In the absence
of oxygen, pyruvate will be reduced in a fermentation pathway or the anaerobic
respiratory pathway. Pyruvate is also an important intermediate in the use of fuel
molecules other than glucose.
Cell Respiration - 7
The enzymes needed to do the Krebs cycle are located in the mitochondrial matrix
or embedded in the inner membrane as integral proteins. The enzymes and
electron carrier complex for electron transport in the respiratory chain are
located in the inner membrane.
Cell Respiration - 8
The two Pyruvate molecules are transported into the inner matrix of the
mitochondria via facilitated diffusion where is undergoes o xidative
decarboxylation within the pyruvate dehydrogenase complex.
• CO2 is removed from pyruvate, producing a 2-carbon compound.
• The 2-carbon fragment is oxidized releasing H+ to reduce NAD+ to NADH,
leaving A cetyl
• Acetyl combines with Co-enzyme A (formed from the B vitamin, pantothenic
acid and added sulfur groups) to form A cetyl-CoA, which can enter the
Krebs cycle.
Pyruvate dehydrogenase
Note: When the level of ATP is high in a cell, the cell can convert acetyl-CoA into
lipid molecules that can be stored for later energy use. This is one way that
excess calories, no matter the nutrient source, are converted to fat.
Cell Respiration - 9
The acids of the Krebs cycle under the right conditions (i.e., The Krebs Cycle) can
be oxidized. They donate H+ and its electron to the appropriate energy transfer
molecule. Once the hydrogen is removed, carbon can also be removed as the waste
product, CO2.
In addition, for each original glucose molecule, two ATP are produced in the Krebs
cycle by s ubstrate-level phosphorylation, one for each acetyl Co-A molecule
that enters the Krebs cycle. (Remember that the glucose molecule has already
gone through glycolysis and been converted to two molecules of pyruvate in the
cytoplasm prior to starting the Krebs cycle.)
In biology, a cycle is a metabolic pathway that starts and ends with the same
molecule. The Krebs cycle starts with Oxaloacetic acid (A 4-carbon acid), which
is regenerated at the end of the cycle. O xaloacetic acid combines with A cetyl-
CoA to begin the Krebs cycle. Most of the enzymes needed to do the Krebs cycle
are located in the mitochondrial matrix. Two, succinate dehydrogenase and -
ketodehydrogenase, are integral membrane proteins of the inner mitochondrial
membrane.
Note: The acids in this process are ionized, and the naming convention is to use
the suffix -ate. For example, oxaloacetic acid may be called oxaloacetate in the
cycle.
The Krebs cycle must turn two times to oxidize the two molecules of Acetyl-CoA
that are what's left of the original glucose molecule.
It should be noted, for preciseness, that a molecule of H2O is also consumed in the
Krebs cycle. Don't worry about it.
Cell Respiration - 10
The Krebs cycle will turn t wo times for each glucose molecule, since glycolysis
produces two pyruvate molecules. Therefore, for each glucose molecule that we
start with, at the completion of the Krebs cycle's two turns, including the
preparation step of pyruvate acetyl we have:
• 6 CO2
• 2 ATP
• 2 FADH2
• 8 NADH
To provide an idea of the potential energy gained from the Krebs cycle, we can look
at the change in free energy as the reactions of the Krebs cycle take place.
Peter Mitchell won the 1978 Nobel prize in chemistry "for his contribution to
the understanding of biological energy transfer through the formulation of
the chemiosmotic theory". ATP is synthesized in the thylakoid membranes
of the chloroplast by a similar mechanism.
• Each NADH that enters the ETS provides sufficient energy to synthesize 3
ATP molecules by chemiosmosis. FADH2 provides energy for 2 ATPs.
ATP Synthase
Total ATP = 36
The maximum ATP may not be realized, since the inner mitochondrial membrane is
leaky to protons and some energy is used to move pyruvate from the cytoplasm
into the mitochondrial matrix. Cells may get about .5 ATP less per reduced carrier
that enters electron transport than the maximum.
Cell Respiration - 15
Plants also have uncoupling proteins for heat generation. Arum lilies attract
carrion beetles and flies for pollination by exuding odors that smell like carrion.
The voodoo lily increases the temperature of the flower so that the odors
volatilize and disperse better. Skunk cabbage elevates its body temperature as
much as 10 - 12° C above ambient air temperature for flowering.
Arum thermogenesis
Cell Respiration - 16
For some prokaryotes and eukaryotes, fermentation is a way of life. Some lack
the enzymes to do the Krebs cycle or oxidative phosphorylation; for others,
oxygen is toxic. These are the strict (or obligate) anaerobes. Others, such as
yeasts and E. coli are f acultative organisms. When oxygen is available, they do
aerobic respiration. When oxygen is not, they perform a fermentation.
In the fermentations the NADH electrons produced in glycolysis are used to reduce
pyruvate to some other organic molecule, which becomes the final electron
acceptor. No more ATP is obtained in the fermentation processes beyond the
two ATP produced during the glycolysis pathway. However, in cells that normally
do aerobic respiration, the rate of glycolysis increases when oxygen is unavailable,
to produce as much ATP as possible.
In the Fermentations:
• Organic molecules serve as the electron acceptors for NADH.
• Among the Prokaryotes there are several different fermentation pathways.
• However only two pathways are found in Eukaryotic organisms:
Alcoholic Fermentation
Lactic Acid Fermentation
Cell Respiration - 17
Any c arbohydrate that can be digested will be converted to glucose. It's just a
matter of time needed to digest and rate of absorption. Polysaccharides and
disaccharides are digested to monosaccharides in the digestive tract. All
monosaccharides absorbed are converted in the liver to glucose for use in
glycolysis.
Fats are energy rich. A gram of fat potentially can produce two times as much
ATP as a gram of carbohydrate or protein. oxidation, which occurs in the
mitochondrial matrix, converts fatty acids to the 2-carbon acetyl. oxidation
uses one ATP, and produces one FADH2 and one NADH along with each acetyl Co-A
formed. A 16-carbon fatty acid can produce 8 acetyl Co-A.
Most cells routinely use a mix of fats and carbohydrates for fuel. The brain, nerve
cells and red blood cells, however, have an absolute glucose requirement; fatty acid
fragments cannot normally cross the brain membrane barriers so that the brain
does not use fats for fuel. The use of fatty acids for fuel is also a strictly aerobic
process. All anaerobic respiration requires glucose.
Two important places for alternative fuel molecules to enter our respiration
pathway are pyruvate and acetyl-CoA. When we have insufficient glucose for our
brain and nerve cells, any molecule that can be converted to pyruvate can
ultimately be used to form glucose, although it is an energy consuming process to
"reverse" glycolysis. However, the step from pyruvate to acetyl is not reversible.
Fuel molecules that are converted to acetyl, or to acids that are a part of the
Krebs cycle are not only unavailable for conversion to glucose, but are useful only
in aerobic respiration.
Cell Respiration - 20
Acetyl is also a major point for the conversion of all excess fuel molecules to fats
as well. If we have sufficient energy, acetyl need not enter the Krebs cycle and is
diverted to the formation of fatty acids for adipose storage. This acetyl can
come from any fuel molecule: glucose, fatty acids or amino acids. All excess
calories consumed, no matter the source, will be converted to adipose.
In general:
• When levels of carbohydrate are high, glucose is metabolized more than fats.
o As levels of glucose fall, stored glycogen (in animals) will be converted
to glucose.
o When glucose supplies diminish, more fat is mobilized to supplement
metabolic needs.
• Protein will be removed from body tissues when carbohydrate is unavailable
to provide glucose for brain and nerve cells.
• When specific nutrients are high, biosynthesis pathways related to nutrient
inter-conversion that would produce those nutrients are stopped.
All excess calories are converted to fat. Fat to fat conversion is efficient, and
any fat consumed not needed for structural or fuel purposes is readily converted
to adipose for storage. Excess carbohydrate beyond the maximum glycogen
stores is also converted to fat in an endergonic process. Some of the caloric
value of the carbohydrate is lost in the conversion. Excess amino acids will be
converted either to glucose, if carbohydrate reserves are low, or to adipose. Much
of this excess nutrient conversion occurs in the respiratory pathway at acetyl.
Just as we convert fatty acids to acetyl to "feed" the Krebs cycle, acetyl not
needed for the Krebs cycle is readily converted to fat.