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Through a set of reactions that occur in the cytosol, energy derived from the partial oxidation of
energy-richcarbohydrate molecules is used to form ATP, the chemical energy currency of cells
(discussed in Chapter 2). But a much more efficient method of energy generation appeared very
early in the history of life. This process is based on membranes, and it enables cells to acquire
energy from a wide variety of sources. For example, it is central to the conversion of light energy
into chemical bond energy in photosynthesis, as well as to the aerobic respiration that enables us
to use oxygen to produce large amounts of ATP from food molecules.
The membrane that is used to produce ATP in procaryotes is the plasma membrane. But in
eucaryotic cells, the plasma membrane is reserved for the transport processes described in
Chapter 11. Instead, the specialized membranes insideenergy-converting organelles are
employed for the production of ATP. The membrane-enclosed organelles aremitochondria,
which are present in the cells of virtually all eucaryotic organisms (including fungi, animals, and
plants), and plastidsmost notably chloroplastswhich occur only in plants.
In electron micrographs the most striking morphological feature of mitochondria and
chloroplasts is the large amount of internal membrane they contain. This internal membrane
provides the framework for an elaborate set of electron-transport processes that produce most of
the cell's ATP.
The common pathway used by mitochondria, chloroplasts, and procaryotes to harness energy for
biological purposes operates by a process known as chemiosmotic couplingreflecting a link
between the chemical bond-forming reactions that generate ATP (chemi) and membranetransport processes (osmotic). The coupling process occurs in two linked stages, both of which
are performed by protein complexes embedded in a membrane:
Stage 1. High-energy electrons (derived from the oxidation of food molecules, from the
action of sunlight, or from other sources discussed later) are transferred along a series
of electron carriers embedded in the membrane. These electron transfers release energy
that is used to pump protons (H+, derived from the water that is ubiquitous in cells) across
the membrane and thus generate an electrochemical proton gradient. As discussed in
Chapter 11, anion gradient across a membrane is a form of stored energy, which can be
harnessed to do useful work when the ions are allowed to flow back across the membrane
down their electrochemical gradient.
Stage 2. H+ flows back down its electrochemical gradient through a protein machine
called ATP synthase, which catalyzes the energy-requiring synthesis of ATP from ADP
and inorganic phosphate (Pi). This ubiquitous enzymeplays the role of a turbine,
permitting the proton gradient to drive the production of ATP (Figure 14-1).
Figure 14-1
Harnessing energy for life. (A) The essential requirements for chemiosmosis are a membrane
in which are embedded a pump protein and an ATP synthase, plus a source of high-energy
electrons (e -). The protons (H+) shown are freely available from (more...)
The electrochemical proton gradient is also used to drive other membraneembedded protein machines (Figure 14-2). In eucaryotes, special proteins couple the
downhill H+ flow to the transport of specific metabolites into and out of the organelles. In
bacteria, the electrochemical proton gradient drives more than ATP synthesis and transport
processes; as a store of directly usable energy, it also drives the rapid rotation of the bacterial
flagellum, which enables the bacterium to swim.
Figure 14-2
Chemiosmotic coupling. Energy from sunlight or the oxidation of foodstuffs is first used to
create an electrochemical proton gradient across a membrane. This gradient serves as a versatile
energy store and is used to drive a variety of energy-requiring (more...)
It is useful to compare the electron-transport processes in mitochondria, which convert energy
from chemical fuels, with those in chloroplasts, which convert energy from sunlight (Figure 143). In the mitochondrion, electronswhich have been released from
a carbohydrate food molecule in the course of its degradation to CO2are transferred through
themembrane by a chain of electron carriers, finally reducing oxygen gas (O2) to form water. The
free energy released as the electrons flow down this path from a high-energy state to a lowenergy state is used to drive a series of three H+pumps in the inner mitochondrial membrane, and
it is the third H+ pump in the series that catalyzes the transfer of the electrons to O2 (see Figure
14-3A).
Figure 14-3
Electron transport processes. (A) The mitochondrion converts energy from chemical fuels. (B)
The chloroplast converts energy from sunlight. Inputs arelight green, products are blue, and the
path of electron flow is indicated by red arrows. Each of the (more...)
The mechanism of electron transport can be compared to an electric cell driving a current
through a set of electric motors. However, in biological systems, electrons are carried between
one site and another not by conducting wires, but by diffusible molecules that can pick up
electrons at one location and deliver them to another. For mitochondria, the first of these electron
carriers is NAD+, which takes up two electrons (plus an H+) to become NADH, a water-soluble
smallmolecule that ferries electrons from the sites where food molecules are degraded to the
inner mitochondrial membrane. The entire set of proteins in the membrane, together with the
small molecules involved in the orderly sequence of electron transfers, is called an electrontransport chain.
Although the chloroplast can be described in similar terms, and several of its main components
are similar to those of the mitochondrion, the chloroplast membrane contains some crucial
components not found in the mitochondrial membrane. Foremost among these are
the photosystems, where light energy is captured by the green pigment chlorophyll and harnessed
to drive the transfer of electrons, much as man-made photocells in solar panels absorb light
energy and use it to drive an electric current. The electron-motive force generated by the
chloroplast photosystems drives electron transfer in the direction opposite to that in
mitochondria: electrons are taken from water to produce O2, and they are donated (via NADPH, a
compound closely related to NADH) to CO2 to synthesize carbohydrate. Thus, the chloroplast
generates O2and carbohydrate, whereas the mitochondrion consumes them (see Figure 14-3B).
It is generally believed that the energy-converting organelles of eucaryotes evolved from
procaryotes that were engulfed by primitive eucaryotic cells and developed a symbiotic
relationship with them. This would explain why mitochondria and chloroplasts contain their
own DNA, which codes for some of their proteins. Since their initial uptake by a host cell, these
organelles have lost much of their own genomes and have become heavily dependent on proteins
that are encoded by genes in the nucleus, synthesized in the cytosol, and then imported into
the organelle. Conversely, the host cells have become dependent on these organelles for much of
the ATP they need for biosyntheses, ion pumping, and movement; they have also become
dependent on selected biosynthetic reactions that occur inside these organelles.
Mitochondria
vs
Chloroplast
are
larger
and
have
greater
complexity
than
are
the
building
blocks
of
life
for plants
and
found
in
chloroplast
membranes.
both
plants
and
animals.