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Aerobic respiration involves the controlled oxidation
of reduced organic compounds to CO2 and H2O.
Much of the free energy released during respiration
is conserved in the form of ATP. In glycolysis, the
first stage of respiration, carbohydrates are oxidized
to organic acids in the cytosol. The organic acids produced
during glycolysis are completely oxidized to CO2
in the mitochondrial matrix via the citric acid cycle.
The electrons released during the operation of the citric
acid cycle are transferred through a series of multiprotein
complexes located in the inner mitochondrial membrane,
ultimately reducing O2 to H2O.
The free energy released during mitochondrial electron
transfer is used to generate a proton electrochemical
gradient across the inner membrane. The energy available
in the proton gradient is subsequently used by another
protein complex, ATP synthase, to synthesize ATP from
ADP and Pi. Mitochondrial respiration is
regulated by the availability of ADP and Pi
and by the presence of additional electron transfer
complexes that allow respiration to proceed without
forming a proton gradient. Plant mitochondria participate
in several metabolic processes besides respiration,
including providing reducing equivalents to other cellular
compartments and carbon skeletons for amino acid biosynthesis.
Plant mitochondria also
participate in the biosynthesis of sugars from lipids
in some germinating seeds and in the decarboxylation
reactions associated with photosynthesis in some plants
having C4 and CAM pathways. The movement
of metabolites into and out of mitochondria requires
specific transporters in the inner mitochondrial membrane,
some of which are regulated by the proton gradient.
Photorespiration involves
the light-dependent uptake of O2 and evolution
of CO2 during photosynthesis in green plant
tissues. The first step in photorespiration is associated
with the oxygenase activity of the photosynthetic enzyme
Rubisco. Phospho-glycolate formed during the oxygenase
reaction is metabolized through the photorespiratory
carbon cycle to save 75% of the carbon in the form of
phosphoglycerate; the remaining 25% is lost as CO2.
The reactions of the photorespiratory carbon cycle occur
in three organelles: chloroplasts, peroxisomes, and
mitochondria. The loss of CO2 during photorespiration
can represent an appreciable percentage of the carbon
fixed during photosynthesis, decreasing the overall
efficiency of photosynthesis. Photorespiration reflects
the evolutionary origin of Rubisco in an anaerobic environment
and may influence the competitiveness of some plants
in response to future changes in atmospheric CO2
concentrations.
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