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Essential Cell Biology 5th edition

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CHAPTER FOURTEEN

14

Energy Generation in

Mitochondria and Chloroplasts

The fundamental need to generate energy efficiently has had a profound

influence on the history of life on Earth. Much of the structure, function,

and evolution of cells and organisms can be traced to their quest for

energy. Oxygen did not appear in the atmosphere until more than a billion

years after the first cells appeared on Earth. It is therefore thought

that the earliest cells may have produced ATP by breaking down organic

molecules that had been generated by geochemical processes. Such fermentation

reactions, discussed in Chapter 13, can occur in the cytosol

of present-day cells, when they use the energy derived from the partial

oxidation of energy-rich food molecules to form ATP.

But very early in the history of life, a much more efficient mechanism

for generating energy and synthesizing ATP appeared—one based on the

transport of electrons along membranes. This mechanism is so central

to the survival of life on Earth that we devote this entire chapter to it.

Membrane-based electron transport first appeared in bacteria more than

3 billion years ago, and the progeny of these pioneering cells currently

crowd every crevice of our planet’s land and oceans in a wild menagerie

of living forms. Perhaps most remarkably, remnants of these energygenerating

electron-transport systems can be found in the bacterial

descendants that labor within living eukaryotic cells: chloroplasts and

mitochondria.

In this chapter, we consider the molecular mechanisms that enable electron-transport

systems to generate the energy that cells need to survive.

We begin with a brief overview of the general principles central to the

generation of energy in all living things: the use of a membrane to harness

the energy of moving electrons. We describe how such processes

MITOCHONDRIA AND

OXIDATIVE PHOSPHORYLATION

MOLECULAR MECHANISMS OF

ELECTRON TRANSPORT AND

PROTON PUMPING

CHLOROPLASTS AND

PHOTOSYNTHESIS

THE EVOLUTION OF ENERGY-

GENERATING SYSTEMS

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