Respiration
Respiratory System |
Cellular
Respiration, process in which cells produce the energy they
need to survive. In cellular respiration, cells use oxygen to break down the
sugar glucose and store its energy in molecules of adenosine
triphosphate (ATP). Cellular respiration is critical for the survival of
most organisms because the energy in glucose cannot be used by cells until it is
stored in ATP. Cells use ATP to power virtually all of their activities—to grow,
divide, replace worn out cell parts, and execute many other tasks. Cellular
respiration provides the energy required for an amoeba to glide toward food, the
Venus fly trap to capture its prey, or the ballet dancer to execute stunning
leaps. Cellular respiration occurs within a cell constantly, day and night, and
if it ceases, the cell—and ultimately the organism—dies.
Two critical ingredients required for
cellular respiration are glucose and oxygen. The glucose used in cellular
respiration enters cells in a variety of ways. Plants, algae, and certain
bacteria make their own glucose through photosynthesis, the process
by which plants use light to convert carbon dioxide and water into sugar.
Animals obtain glucose by eating plants, and fungi and bacteria absorb glucose
as they break down the tissues of plants and animals. Regardless of how they
obtain it, cells must have a steady supply of glucose so that ATP production is
continuous.
Adenosine Triphosphate |
Oxygen is present in the air, and also is
found dissolved in water. It either diffuses into cells—as in bacteria, fungi,
plants, and many aquatic animals, such as sponges and fish—or it is inhaled—as
in more complex animals, including humans. Cellular respiration sometimes is
referred to as aerobic respiration, meaning that it occurs in the presence of
oxygen.
Cellular respiration transfers about 40
percent of the energy of glucose to ATP. The rest of the energy from glucose is
released as heat, which warm-blooded organisms use to maintain body temperature,
and cold-blooded organisms release to the atmosphere. Cellular respiration is
strikingly efficient compared to other energy conversion processes, such as the
burning of gasoline, in which only about 25 percent of the energy is used and
about 75 percent is released as heat.
While most organisms carry out cellular
respiration to produce ATP, some cannot produce ATP through this process because
they live in anaerobic environments, or environments that lack oxygen. These
organisms, typically bacteria, rely on anaerobic processes such as
fermentation to generate their ATP.
CHEMICAL REACTIONS AND METABOLIC PATHWAYS
To understand cellular respiration, it is
necessary to understand the nature of chemical reactions. Chemical
reactions can occur outside of living organisms—the rusting of a car, for
example, is a chemical reaction—or they can occur within organisms, where they
are termed biochemical reactions. In a chemical or biochemical reaction, the
bonds between atoms that hold molecules together break apart, and the atoms
rearrange to form new molecules. Water molecules, for example, are composed of
hydrogen and oxygen atoms, and under certain conditions, the bonds between these
atoms can break and reform to yield separate molecules of hydrogen and oxygen
gas. In living organisms, most biochemical reactions occur with the help of
enzymes, specialized proteins designed to carry out specific
reactions. All biochemical reactions release energy in the form of heat as they
occur.
Cells carry out biochemical reactions to
create needed molecules—such as proteins or starch—or to destroy these molecules
once they are no longer needed. If certain molecules are built or destroyed in a
single biochemical reaction, the reaction may release too much heat, which could
incinerate the cell. To control the release of heat, cells build up and break
down most molecules in a linked series of small reactions that release only a
little bit of heat at a time. The series of linked biochemical reactions is
called a metabolic pathway.
Cellular respiration is one of the most
important metabolic pathways found in cells. This enzyme-assisted, step-by-step
process not only protects the cell from lethal temperature increases but also
provides the cell with a mechanism of transferring the energy of glucose to ATP
in a controlled manner.
HOW CELLULAR RESPIRATION WORKS
The process of cellular respiration occurs in four stages: glycolysis; the
transition stage; the Krebs cycle, also known as the citric acid
cycle; and the electron transport chain. Each stage accomplishes
different tasks.
A. Glycolysis
Glucose is the primary fuel used in
glycolysis, the first stage of cellular respiration. This all-important molecule
is found in the cell’s cytoplasm, the gel-like substance that fills the cell.
Glucose consists of 6 carbon, 12 hydrogen, and 6 oxygen atoms bonded together,
along with electrons (negatively charged atomic particles) associated with each
atom. Of these components, only the hydrogen atoms and certain electrons
participate directly in glycolysis.
In glycolysis, glucose is broken down
with the help of enzymes and other molecules found in the cytoplasm. Enzymes
first attach two phosphate groups to glucose to make it more reactive. A
phosphate group is a cluster of one phosphorus and four oxygen atoms. The
addition of the two phosphate groups prepares glucose for the action of another
enzyme. This enzyme splits glucose in half to produce two three-carbon
molecules, each with one phosphate group attached.
In the next step, an enzyme removes one
hydrogen atom and two electrons from each three-carbon molecule. Both hydrogen
atoms are modified to hydrogen ions, positively charged particles. A hydrogen
ion and two electrons from each three-carbon molecule are transferred as a unit
to a large molecule called nicotinamide adenine dinucleotide (NAD+)
to form two molecules of NADH. The hydrogen ions and electrons stored in each
molecule of NADH are used to make ATP in later stages of cellular respiration.
In the final steps of glycolysis, two
hydrogen atoms are removed from each three-carbon compound. These hydrogen atoms
bond to free-floating oxygen atoms in the cytoplasm to form water. This step
prepares the two three-carbon compounds for action by the next enzyme in the
pathway. This enzyme removes the phosphate group from each three-carbon
compound. Each phosphate group then bonds to a single molecule of adenosine
diphosphate (ADP). ADP is composed of three carbon-based rings and a tail of two
phosphate groups. The addition of the third phosphate group to the tail forms
ATP. In this step, two new ATP molecules are produced. When cells require
energy, another enzyme breaks off the third phosphate group, releasing energy
that powers the cell. The removal of the third phosphate from ATP converts ATP
back to ADP, which is used again in cellular respiration to make more ATP.
When the two three-carbon compounds are
separated from the phosphate groups, the three-carbon compounds are converted to
two molecules of pyruvate, each composed of three carbon, three oxygen, and
three hydrogen atoms. When glycolysis is complete, important products used in
the next steps of cellular respiration have been produced: two molecules of NADH
and two molecules of pyruvate. The two ATP molecules gained in glycolysis are
used for reactions in the cell that require energy.
Mitochondria |
B. Transition Stage
The pyruvate molecules move from the
cytoplasm to special structures in the cell called mitochondria,
where the remaining steps of cellular respiration are carried out. Each
mitochondrion contains a membrane that is folded back and forth many times. This
extensive membrane is studded with hundreds of thousands of enzymes that direct
cellular respiration. The numerous enzymes enable great quantities of ATP to be
produced simultaneously in one mitochondrion. Without mitochondria or a similar
structure, most cells could not generate enough ATP to survive.
The transition stage is a short
biochemical pathway that links glycolysis with the Krebs cycle. In this brief
stage, enzymes transfer hydrogens and electrons from the two pyruvate molecules
to two molecules of NAD+ to form two more molecules of NADH. Another
enzyme breaks off one carbon and two oxygen atoms from each pyruvate molecule.
These atoms combine to form carbon dioxide, the primary waste product of
cellular respiration, which diffuses out of the cell. As a result of these
reactions, each pyruvate molecule is transformed into a two-carbon compound
called an acetyl group. The two acetyl groups unite with two molecules of
coenzyme A to form two acetyl coenzyme A molecules. The acetyl coenzyme A
molecules are the molecules that enter the Krebs cycle.
C. Krebs Cycle
Krebs Cycle |
During the Krebs cycle, the acetyl coenzyme A molecules are processed. As this complex pathway progresses, six molecules of NADH are formed. Additional carbon dioxide is created, and this process releases energy that is used to build two molecules of ATP from a pool of ADP and phosphate groups in the mitochondria. Hydrogens and electrons then are transferred to a molecule of flavin adenine dinucleotide (FAD++)to form FADH2, a molecule like NADH that temporarily stores hydrogen and electrons for later use. By the end of the Krebs cycle, most of the usable energy from the original glucose molecule has been transferred to ten molecules of NADH (two from glycolysis, two from the transition stage, and six from the Krebs cycle); two molecules of FADH2; and four molecules of ATP, two of which were formed in glycolysis.
D. Electron Transport Chain
The reactions of the electron transport
chain occur in several closely spaced molecules embedded in the mitochondrial
membrane. Acting like specialized delivery trucks, the NADH and FADH2
molecules dump off their load of electrons and hydrogen ions near these electron
transport chain molecules. The first molecule in the chain has an attraction for
electrons and grabs them, but the molecule next to it in the chain has an even
stronger attraction and grabs the electrons away from the first molecule. The
electrons are passed down the chain in this manner, until they reach oxygen, the
final molecule in the chain. Oxygen has a stronger appetite for electrons than
any molecule in the chain, and the electrons therefore are held by oxygen. They
are joined by the hydrogen ions that were dropped off by NADH and FADH2
at the beginning of the electron transport chain. The combination of the
electrons, hydrogen ions, and oxygen forms water, used by the cell in other
biochemical reactions. As NADH and FADH2 release hydrogen and
electrons in the electron transport chain, they are converted back to
NAD+ and FAD++, respectively, providing the cell with a
steady supply of these molecules so that cellular respiration can be carried out
over and over again.
As the electrons flow down the electron
transport chain, they release a veritable windfall of energy that is used by an
enzyme to make more ATP, again from the pool of ADP and phosphate groups in the
mitochondria. In most cells, the electron transport chain produces 32 molecules
of ATP. Together with the two ATP molecules gained in glycolysis and the four
generated in the Krebs cycle, cellular respiration produces a grand total of 38
molecules of ATP for every molecule of glucose processed.
Glucose molecules enter the cell by the
hundreds of thousands. They are processed simultaneously to generate millions of
ATP molecules every second. Some of the ATP molecules remain in the mitochondria
to supply it with needed energy, but the majority stream from the mitochondria
to the cytoplasm, where they fuel the cell’s activities. It is estimated that a
single human brain cell uses a staggering 10 million ATP molecules per second to
carry out its tasks.
Although glucose is the primary fuel for
cellular respiration, cells can rely on other molecules to produce ATP. The
cellular respiration pathway is connected to other metabolic pathways that can
donate molecules to cellular respiration at different steps along the way. For
example, glycerol, a breakdown product of fat, can enter the cellular
respiration pathway in the middle of glycolysis. Another product of fat
digestion, fatty acids, can enter at the transition stage. Glycerol is modified
in glycolysis to pyruvate, and fatty acids are modified to acetyl coenzyme A in
the transition stage. The pyruvate and acetyl coenzyme A are processed through
the remaining steps of cellular respiration to yield ATP. The ability to use
alternate molecules enables cells to keep ATP production going even if they run
out of glucose. Marathon runners, for example, first use up their glucose
reserves to make ATP, and then draw on their fat reserves to generate the ATP
needed to get to the finish line.
IV. ANAEROBIC PATHWAYS TO ATP
Although most organisms on Earth carry out
cellular respiration to generate ATP, a few rely on alternative pathways to make
this vital molecule. These pathways are anaerobic—that is, they do not require
oxygen. Fermentation is a type of anaerobic pathway used by certain species of
bacteria that live in anaerobic environments, such as stagnant ponds or decaying
vegetation. Some cells produce ATP using both anaerobic and aerobic pathways.
For example, muscle cells typically carry out cellular respiration, but if they
do not receive enough oxygen, as can occur during strenuous exercise, muscles
switch to fermentation. Yeast cells also carry out both pathways, depending on
whether they are in an aerobic environment, such as soil, or an anaerobic one,
such as inside a wet lump of dough. Fermentation is much less efficient than
cellular respiration, however, producing only two molecules of ATP for every
glucose molecule processed.
Fermentation occurs in two stages:
glycolysis and the recycling stage. The glycolysis pathway in fermentation is
virtually the same as the glycolysis pathway of cellular respiration, relying on
enzymes, NAD+, and other molecules to transfer the energy of glucose
to two molecules each of pyruvate, NADH, and ATP. This is the only stage of
fermentation that yields ATP, and while it produces a relatively small amount,
it is ample for simple cells.
Molecules of NAD+, like other
molecules required in glycolysis, must be in constant supply for glycolysis to
continue without interruption. In the recycling stage of fermentation, the NADH
produced in glycolysis is recycled back into NAD+ to be used again in
glycolysis.
While all cells that carry out fermentation
use the recycling stage to make more NAD+, slight variations exist,
depending on the organism involved. Many variations produce a number of
interesting byproducts that are waste as far as the cell is concerned but are
useful to humans. In yeast, for example, the recycling stage produces the carbon
dioxide gas that makes dough rise in bread making and the alcohol
that makes beer and wine intoxicating. In certain
kinds of bacteria, the recycling stage produces the lactic acid that turns
milk into yogurt or cheese.
Other types of anaerobic pathways are
carried out by certain species of archaebacteria that live in extreme
environments, such as hydrothermal vents, springs of hot water in
the deep ocean. These anaerobic pathways begin with glycolysis and are followed
by two subsequent stages that resemble the Krebs cycle and electron transport
chain of cellular respiration. However, these anaerobic pathways yield about
half the amount of ATP that cellular respiration yields