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Over a billion years ago, aerobic bacteria were engulfed by eukaryotic cells and evolved into
mitochondria, providing the eukaryotic cells with the ability to form the energy-rich compound
ATP by oxidative phosphorylation. Mitochondria perform other functions, including a role in
the regulation of apoptosis (programmed cell death), but oxidative phosphorylation is the most
crucial. Each eukaryotic cell can have hundreds to thousands of mitochondria. In mammals, they
are generally depicted as sausage-shaped organelles (Figure 21), but their shape can be quite
dynamic. Each has an outer membrane, an intermembrane space, an inner membrane, which is
folded to form shelves (cristae), and a central matrix space. The enzyme complexes responsible
for oxidative phosphorylation are lined up on the cristae (Figure 24).
Figure 24
As zygote mitochondria are derived from the ovum, their inheritance is maternal. This maternal
inheritance has been used as a tool to track evolutionary descent. Mitochondria have an
ineffective DNA repair system, and the mutation rate for mitochondrial DNA is over 10 times
the rate for nuclear DNA. A large number of relatively rare diseases have now been traced to
mutations in mitochondrial DNA. These include for the most part disorders of tissues with high
metabolic rates in which energy production is defective as a result of abnormalities in the
production of ATP.
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Mitochondria
Mitochondria (mito-kondre-ah) are threadlike (mitos = thread) or sausage-shaped membranous
organelles (Figure 3.17). In living cells they squirm, elongate, and change shape almost continuously.
They are the power plants of a cell, providing most of its ATP supply. The density of mitochondria in a
particular cell reflects that cells energy requirements, and mitochondria are generally clustered where
the action is. Busy cells like kidney and liver cells have hundreds of mitochondria, whereas relatively
inactive cells (such as unchallenged lymphocytes) have just a few. Mitochondria are enclosed by two
membranes, each with the general structure of the plasma membrane. The outer membrane is smooth
and featureless, but the inner membrane folds inward, forming shelflike cristae (krste; crests) that
protrude into the matrix, the gel-like substance within the mitochondrion. Intermediate products of
food fuels (glucose and others) are broken down to water and carbon dioxide by teams of enzymes,
some dissolved in the mitochondrial matrix and others forming part of the crista membrane.
As the metabolites are broken down and oxidized, some of the energy released is captured and used
to attach phosphate groups to ADP molecules to form ATP. This multistep mitochondrial process
(described in Chapter 24) is generally referred to as aerobic cellular respiration (a-er-obik) because it
requires oxygen.
Mitochondria are complex organelles: They contain their own DNA and RNA and are able to reproduce
themselves. Mitochondrial genes (some 37 of them) direct the synthesis of some 5% of the proteins
required for mitochondrial function, and the DNA of the cells nucleus encodes the remaining proteins
needed to carry out cellular respiration. When cellular requirements for ATP increase, the mitochondria
synthesize more cristae or simply pinch in half (a process called fission) to increase their number, then
grow to their former size.
Intriguingly, mitochondria are similar to a specific group of bacteria (the purple bacteria phylum), and
mitochondrial DNA is similar to that found in bacterial cells. It is now widely believed that mitochondria
arose from bacteria that invaded the ancient ancestors of plant and animal cells.
The Cytoskeleton
The cytoskeleton is a collection of protein filaments and
cylinders that determine the shape of a cell, lend it structural
support, organize its contents, direct the movement of
substances through the cell, and contribute to movements
of the cell as a whole. It can form a very dense supportive
scaffold in the cytoplasm (fig. 3.31). It is connected to
transmembrane proteins of the plasma membrane, and
they in turn are connected to protein fibers external to the
cell, so there is a strong structural continuity from extracellular
material to the cytoplasm. Cytoskeletal elements may