produces NADPH and a proton gradient across the thylakoid membrane.
NADPH is used directly in the Calvin cycle(below). The proton gradient is
used to drive adenosine triphosphate (ATP)synthase, the enzyme that makes
ATP.
The second photosystem, PS-I, can work independently of PS-II. This occurs
when electrons are transferred from cytochrome b/fback to P 700 via plasto-
cyanin and re-energized by light. This process, termed cyclic electron flowis
illustrated in Fig. 2. As the electrons never reach NADP reductase, the system
only generates a proton gradient.
ATP is produced by ATP synthase(Fig. 3). This is a large protein complex
located in the thylakoid membrane. As the protons flow down the electrochem-
ical gradient from the thylakoid lumen into the stroma, the energy is used to
synthesize ATP by the phosphorylation of adenosine diphosphate (ADP) by the
ATP synthase complex. ATP synthesis driven by cyclic electron flow (PS-I only)
is termed cyclic photophosphorylation, while ATP synthesis driven by non-
cyclic electron flow (PS-I and PS-II) isnon-cyclic photophosphorylation.
The next stages of photosynthesis do not require light and are termed the
carbon-fixation reactions. They do require the ATP and NADPH generated by
the light reactions and result in the incorporation (fixation) of carbon into carbo-
hydrates. The carbon-fixation reactions in many plants occur in the stroma of
the chloroplast by the Calvin cycle.
The carbon-
fixation
reactions
140 Section J – Metabolism
Redox
potential
(V)
P700
Photosystem
I
P700*
Light
Fd
PC
Cytochrome
bf
complex H+ pump
+
Fig. 2. Cyclic phosphorylation in green plants.