In addition to being involved in protein sorting and transport, cytoplasmic organelles provide specialized compartments in which a variety of metabolic activities take place. The generation of metabolic energy is a major activity of all cells, and two cytoplasmic organelles are specifically devoted to energy metabolism and the production of ATP. Mitochondria are responsible for generating most of the useful energy derived from the breakdown of lipids and carbohydrates, and chloroplasts use energy captured from sunlight to generate both ATP and the reducing power needed to synthesize carbohydrates from CO2 and H2O. The third organelle discussed in this chapter, the peroxisome, contains enzymes involved in a variety of different metabolic pathways, including the breakdown of fatty acids.

Mitochondria, chloroplasts, and peroxisomes differ from the organelles discussed in the preceding chapter not only in their functions but also in their mechanism of assembly. Rather than being synthesized on membrane-bound ribosomes and translocated into the endoplasmic reticulum, most proteins destined for mitochondria, chloroplasts, and peroxisomes are synthesized on free ribosomes in the cytosol and imported into their target organelles as completed polypeptide chains. Mitochondria and chloroplasts also contain their own genomes, which include some genes that are transcribed and translated within the organelle. Protein sorting to the cytoplasmic organelles discussed in this chapter is thus distinct from the pathways of vesicular transport that connect the endoplasmic reticulum, Golgi apparatus, lysosomes, and plasma membrane.