Potential vs. Kinetic Energy in the Chloroplast
- Pumping protons (H⁺) into the thylakoid lumen stores potential energy.
- Comparable to pumping water up a hill in a hydro-electric dam.
- When protons flow back through ATP synthase, potential → kinetic energy (rotation of the rotor).
- Hydroelectric analogy: water potential drives turbines → electrical energy; proton potential drives ATP synthase → chemical energy (ATP).
Light-Dependent Reactions Overview
- Occur on the thylakoid membrane; include Photosystem II (PS II), cytochrome b6f complex, Photosystem I (PS I), ferredoxin, and ATP synthase.
- Primary outputs: \text{ATP} and \text{NADPH} (energy & reducing power for the Calvin cycle).
- Overall energy flow:
- Photons excite pigments.
- Electron transport chain (ETC) pumps H⁺ into lumen.
- Proton-motive force drives ATP synthase.
- Electrons eventually reduce \text{NADP}^+ \;\rightarrow\; \text{NADPH}.
Photosystem II (PS II)
- First complex hit by light (though discovered second).
- Only biological system able to oxidize water.
- Splits \text{H}2\text{O} → 4\,e^- + 4\,H^+ + O2.
- Special reaction-center pair: P680 (absorbs 680\,\text{nm}).
- Primary electron acceptor: pheophytin → plastoquinone (PQ).
- PQ shuttles electrons to cytochrome b6f and simultaneously pumps H⁺.
Cytochrome b6f Complex & Plastoquinone (PQ)
- Acts like mitochondrial Complex III.
- PQ cycle translocates additional H⁺ from stroma to lumen (builds gradient).
Plastocyanin (PC)
- Soluble Cu-protein in lumen; transfers electrons from cytochrome b6f to PS I.
- Physical bridge linking PS II and PS I.
Photosystem I (PS I)
- Special pair P700 (absorbs 700\,\text{nm}).
- Photons re-energize incoming electrons (they lost energy traveling downhill from PS II).
- Electrons passed to ferredoxin (Fd) → NADP⁺ reductase (FNR) → \text{NADPH}.
Z-Scheme (Energy Diagram)
- Illustrates the two “ups” in redox potential: one at PS II, one at PS I.
- Older view treated systems as separate; modern “Z-scheme” shows simultaneous, continuous flow.
Cyclic vs. Non-Cyclic Electron Flow
- Non-cyclic (described above): net production of \text{ATP}, \text{NADPH}, O_2.
- Cyclic (PS I only): electrons cycle back to cytochrome b6f → extra \text{ATP}, no \text{NADPH}, no O_2; helps adjust ATP/NADPH ratio.
ATP Synthase & Proton Gradient
- ~3\,H^+ moving down gradient → rotation → phosphorylation \text{ADP} + P_i \;\rightarrow\; \text{ATP}.
- Same rotary mechanism as mitochondrial ATP synthase.
Why Make ATP & NADPH?
- Fuel the Calvin cycle (light-independent reactions).
- PS II → majority of ATP; PS I → all NADPH.
Calvin Cycle (C₃ Pathway)
- Location: chloroplast stroma.
- Discovered by Melvin Calvin; initial acceptor: ribulose-1,5-bisphosphate (RuBP) (5 C).
- Enzyme: RuBisCO (ribulose bisphosphate carboxylase/oxygenase).
- Most abundant protein on Earth; slow, large, smells “plant-y”.
- Catalyzes addition of CO2 or O2 (photorespiration).
Three Phases
- Carbon Fixation
- \text{RuBP} (5\,C) + CO_2 \xrightarrow[{RuBisCO}]{} 2\,\times\,3\text{PGA} (3\,C).
- Reduction
- Uses 6\,ATP + 6\,NADPH to convert 3\text{PGA} → glyceraldehyde-3-phosphate (G3P).
- Regeneration
- 5\,\text{G3P} + 3\,ATP → regenerate 3\,\text{RuBP}.
- For every 1 G3P exported, 5 are recycled.
Net Stoichiometry (per G3P exported)
3\,CO2 + 9\,ATP + 6\,NADPH \;\rightarrow\; G3P + 9\,ADP + 8\,Pi + 6\,NADP^+
Fate of G3P
- 1 G3P (3 C) ⇒ combine two G3P → glucose/fructose (6 C).
- Glucose + fructose → sucrose; storage as starch in chloroplast.
Stomata, Gas Exchange & Water Loss
- Leaf surface coated by lipid cuticle to limit evaporation.
- Stoma (sing.) / Stomata (pl.): pore formed by two guard cells.
- Open → CO2 in, O2 + water vapor out.
- Close to conserve water or when CO_2 sufficient.
- Guard-cell dynamics regulate photosynthesis vs. transpiration.
Photorespiration
- RuBisCO oxygenase activity: RuBP + O_2 \rightarrow 3\text{PGA} + 2\text{C glycolate}.
- Consumes O2, releases previously fixed CO2; wastes ATP & NADPH.
- Hypothesized regulatory/photoprotective role under high light, low CO_2.
Regulation of Photosynthesis
- Light triggers synthesis/activation of photosynthetic proteins.
- High sucrose/glucose in cell → repress photosynthetic genes, induce storage pathways.
- RuBisCO activated by light-linked signals; inhibited when stromal CO_2 low.
- Balance of ATP/NADPH fine-tuned by switching between cyclic & non-cyclic flow.
Global Significance
- Carbon fixation in Calvin cycle supplies nearly all organic carbon for ecosystems.
- Every carbon atom in your body once passed through RuBisCO.
Transition to Cellular Respiration
- Plants: perform both photosynthesis (chloroplast) and cellular respiration (mitochondria).
- Animals & other heterotrophs: rely solely on respiration; glucose ultimately comes from plants (or organisms that ate plants).
Intro to Cellular Respiration (Preview)
- Overall equation (aerobic):
C6H{12}O6 + 6\,O2 \;\rightarrow\; 6\,CO2 + 6\,H2O + \text{energy (ATP + heat)} - Sequence of pathways:
- Glycolysis (cytosol) → 2 pyruvate, small ATP, NADH.
- Pyruvate oxidation (mitochondrial matrix).
- Citric Acid (TCA) Cycle → more NADH, FADH2, CO2.
- Electron Transport Chain & Oxidative Phosphorylation → large ATP yield via proton gradient (chemiosmosis).
- Analogy to automobile engine: glucose ≈ gasoline; O_2 as oxidant; ATP ≈ mechanical work; heat lost.
- Fermentation: backup ATP production without O_2; lower energy yield; end products retain potential energy (e.g., lactate, ethanol).
Hydroelectric Dam Analogy (Revisited)
- Upper reservoir water ≈ proton gradient.
- Turbine rotation ≈ ATP synthase.
- Pumping water uphill ≈ ETC-driven H⁺ pumping (respiration or photosynthesis).
Study & Exam Tips Mentioned by Instructor
- Redraw diagrams yourself (track carbons/electrons).
- Understand, don’t just memorize; focus on inputs/outputs of each phase.
- Expect quiz on energetics & photosynthesis; final exam soon after.
Key Equations & Numbers (LaTeX Style)
- Photosynthesis summary:
6\,CO2 + 6\,H2O + \text{light} \;\rightarrow\; C6H{12}O6 + 6\,O2 - Cellular respiration summary:
C6H{12}O6 + 6\,O2 \;\rightarrow\; 6\,CO2 + 6\,H2O + \text{ATP} - Proton-to-ATP ratio (chloroplast): \sim 3\,H^+ / \text{ATP}.
Connections to Previous & Future Topics
- Chemiosmosis concept shared across chloroplasts and mitochondria.
- Redox coupling: exergonic electron flow drives endergonic ATP/NADPH synthesis.
- Calvin cycle parallels gluconeogenesis & glycogenesis in animals (anabolic sugar formation).
- Cellular respiration will mirror photosynthesis in reverse redox flow.
Ethical / Ecological Implications
- Plant carbon fixation mitigates atmospheric CO_2—critical for climate regulation.
- Blocking RuBP regeneration would halt global food webs (comic-villain thought experiment).
- Understanding photorespiration aids crop engineering for higher yields under climate stress.
- Hydroelectric dam = thylakoid lumen gradient.
- Breaking Hershey bar into two = glycolysis splitting glucose.
- Money analogy: spend 1 G3P (dinner), bank 5 G3P (savings) to keep cycle running.
- NADPH = rechargeable battery; ATP = arcade token (spendable currency).
Numerical Highlights
- Calvin cycle cost per G3P: 9\,ATP + 6\,NADPH.
- Light reactions ATP/NADPH ratio adjustable via cyclic flow.
- Cells store only \approx 30!\text{–}!60\,\text{s} worth of ATP – must regenerate continuously.
Vocabulary Recap
- Potential vs. kinetic energy, chemiosmosis, proton-motive force, resonance energy transfer, redox, oxidized vs. reduced, electron carrier, photorespiration, cuticle, guard cells, stoma/stomata, Z-scheme, RuBP, RuBisCO, G3P, 3-PGA, ferredoxin, plastocyanin, plastoquinone, cytochrome b6f, ATP synthase, cyclic electron flow, non-cyclic electron flow, Calvin cycle phases, glycolysis, citric acid cycle, oxidative phosphorylation.