23.
Energy Transformations: Respiration and Photosynthesis
Respiration: Reoxidizing The Carriers
Ubiquinone is a small carrier molecule with a long
isoprene-derived tail related to the phytol tail of chlorophyll,
to b-carotene, and to the other terpene
derivatives discussed in Chapter
20. The working head of ubiquinone is a quinone ring that can
accept two hydrogen atoms at the para positions and give them up
again. From NADH to ubiquinone, reduction involves the transfer
of hydrogen atoms. Beyond ubiquinone, the reducing hydrogens are
split into protons, which are released into the solution, and electrons,
which travel through the cytochromes from one heme iron atom to
the next. Each cytochrome molecule is reduced to the Fe (II) state
by the one before it, and then reoxidized to Fe(III).
All of the foregoing respiratory reactions take place
inside mitochondria within the cell. The components of the respiratory
chain are embedded in the inner mitochondrial membrane, and are
organized into four complexes. Complex 1 contains a flavoprotein
(a nonheme iron protein of uncertain function) and phospholipid,
and has a molecular weight of around 600,000. Complex III (270,000
molecular weight) contains cytochromes c and c1,
more nonheme iron protein, and phospholipids. Complex IV (eytochrome
oxidase) has a weight of 200,000 and contains cytochromes a
and a3, copper atoms, and phospholipids. Each
pair of complexes is connected by a mobile shuttle, ubiquinone between
complexes I and III, and the small cytochrome c molecule between
III and IV.