(N/A) The chemiosmotic hypothesis explains the mechanism of $ATP$ synthesis in chloroplasts.
- Similar to respiration,in photosynthesis,$ATP$ synthesis is linked to the development of a proton gradient across a membrane,specifically the thylakoid membrane.
- Main difference: In respiration,protons $(H^{+})$ accumulate in the intermembrane space of the mitochondria during electron transport system $(ETS)$ activity.
Formation of proton gradient:
$(a)$ Since water splitting occurs on the inner side of the membrane,the protons $(H^{+})$ produced accumulate within the lumen of the thylakoids.
$(b)$ As electrons move through the photosystems,protons are transported across the membrane. The primary electron acceptor,located on the outer side of the membrane,transfers its electron to an $H$ carrier rather than an electron carrier. This molecule removes a proton from the stroma while transporting an electron. When it passes the electron to the carrier on the inner side,the proton is released into the lumen.
Reduction of $NADP^{+}$:
$(c)$ The $NADP$ reductase enzyme is located on the stroma side. Along with electrons from $PS I$,protons are required for the reduction of $NADP^{+}$ to $NADPH + H^{+}$. These protons are also removed from the stroma.
$NADP^{+} + 2H^{+} \longrightarrow NADPH + H^{+}$
- Consequently,protons in the stroma decrease,while they accumulate in the lumen,creating a proton gradient and a measurable decrease in $pH$ in the lumen.
Importance: The breakdown of this gradient releases energy. Protons move back to the stroma through the transmembrane channel of the $ATPase$ enzyme.
Structure of $ATPase$ enzyme:
- $F_{0}$: This part is embedded in the membrane and forms a transmembrane channel that facilitates the diffusion of protons.
- $F_{1}$: This portion protrudes on the outer surface of the thylakoid membrane facing the stroma. The energy released from the gradient breakdown causes a conformational change in $F_{1}$,enabling the synthesis of $ATP$.
Requirements of Chemiosmosis:
$(a)$ $A$ membrane,a proton pump,a proton gradient,and $ATPase$.
$(b)$ Energy is used to pump protons across the membrane to create a high concentration within the thylakoid lumen.
$(c)$ $ATPase$ allows proton diffusion back to the stroma,releasing energy to catalyze $ATP$ formation.
Usage: $ATP$ and $NADPH$ produced are used immediately in the biosynthetic reactions in the stroma for $CO_{2}$ fixation and sugar synthesis ($C_{3}$ pathway).