Lecture 6 - Photosynthesis: The Light ReactionsLecture 6 - Photosynthesis: The Light Reactions. 2...

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Chem 454: Regulatory Mechanisms in Biochemistry University of Wisconsin-Eau Claire Lecture 6 - Photosynthesis: The Light Reactions

Transcript of Lecture 6 - Photosynthesis: The Light ReactionsLecture 6 - Photosynthesis: The Light Reactions. 2...

Page 1: Lecture 6 - Photosynthesis: The Light ReactionsLecture 6 - Photosynthesis: The Light Reactions. 2 Text Photosynthesis represents for the biosphere the major source of Free energy (1017

Chem 454: Regulatory Mechanisms in BiochemistryUniversity of Wisconsin-Eau Claire

Lecture 6 - Photosynthesis:The Light Reactions

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TextPhotosynthesis represents for the biosphere the major source of

Free energy (1017 kcal/year stored)

Carbon (1010 tons/year assimilated)

Oxygen

Introduction

CO2 + H2O (CH2O) + O2Light

carbohydrate

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Analogous to the combination of oxidative phosphorylation and the citric acid cycle.

Overview

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TextLike oxidative phosphorylation, photosynthesis is compartmentalized

1. Chloroplasts

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Mitochondria are bound by a double membrane

1 Mitochondria

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TextLight absorption by chlorophyll induces electron transfer

2. Electron Transfer

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TextAbsorption of light leads to photoinduced charge separation

2. Electron Transfer

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TextAbsorption of light leads to photoinduced charge separation

2. Electron Transfer

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Plants have two photosystemsPhotosystem I13 polypeptides chains

60 Chlorophylls

3 4Fe-4S centers

1 Quinone

Photosystem II10 polypeptide chains

30 Chlorophylls

1 Non-heme Fe

4 Mn+2 (Mn+2, Mn+3, Mn+4 , Mn+5)

2.1 Bacteria vs. Green Plants

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Bacteria have a single system4 Bacteriochlorophylls b2 Bacteriopheophytin b2 Quinones1 Fe2+

2.1 Bacteria vs. Green Plants

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Bacteria have a single system

2.1 Bacteria vs. Green Plants

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2.2 The Bacterial Photosynthetic Reaction Center

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The rate of electron transfer is dependent up two factors:

2.3 Electron Transfer

Distance Driving Force

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3.3 Q-Cytochrome c Oxidoreductase

The Q cycle:

QH2 + 2 Cyt cox 2 H+matrix Q + 2 Cyt cred + 4 H+cytosol+

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TextIn green plants there are two photosynthetic reaction centers.

3. Two Photosystems of Plants

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The core of PSII is similar to the bacterial system.

3.1 Photosystem II

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3.1 Photosystem II

2 Q + 2 H2O O2 + 2 QH2

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3.1 Photosystem II

Four photons are required to generate one oxygen molecule

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3.1 Photosystem II

The 4 Manganese center is where the electrons are extracted from the water.

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3.1 Photosystem II

An equivalent of 4 H+ are moved across the thylakoid membrane by PSII

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The cytochrome bf complex transfers the electrons from plastoquinone to platocyanin:

3.2 Cytochrome bf

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3.2 Cytochrome bf

2 QH2 + 2 Pc(Cu2+) Q + 2 Pc(Cu+) 2 H+thylakoid lumen+

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3.3 Q-Cytochrome c Oxidoreductase

The Q cycle:

QH2 + 2 Cyt cox 2 H+matrix Q + 2 Cyt cred + 4 H+cytosol+

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Though larger, the core of PSI is also similar to the bacterial system.

3.3 Photosystem I

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3.3 Photosystem I

The receptor of the the electrons from PSI is the small one-electron redox protein, Ferredoxin.

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3.3 Photosystem I

Pc(Cu+) + Fdox Pc(Cu2+) + Fdred

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The Z-scheme of photosynthesis

3.3 Photosystem I

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Ferredoxin-NADP+ reductase

3.4 Ferredoxin-NADP+ Reductase

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Ferredoxin-NADP+ reductase contains the coenzyme FAD.

3.4 Ferredoxin-NADP+ Reductase

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Ferredoxin-NADP+ reductase contains the coenzyme FAD.

3.4 Ferredoxin-NADP+ Reductase

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TextPSII, Cytochrome bf and ferredoxin-NADP+ reductase each contribute to the proton gradient.

4. The Proton Gradient

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TextAndré Jagendorf's demostration

1966, was one of the earliest pieces of evidence to support Peter Mitchell's chemiosmotic hypothesis.

4. The Proton Gradient

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The ATP synthase closely resembles the ATP synthase of mitochondria

4.1 ATP Synthase

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Comparing photosynthesis to oxidative phosphorylation:

4.1 ATP Synthase

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Normally photosynthesis produces both ATP and NADPH.

When there is not NADP+ available, cyclic flow of electrons through PSI can be used to produce ATP without reducing NADP+

4.2 Flow of Electrons Through Photosystem I

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Cyclic flow of electrons through PSI:

4.2 Flow of Electrons Through Photosystem I

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12 protons are expected to be used for one turn of the ATP synthase, therefore, producing 3 AtP's from 3ADP's and 3Pi's

4.3 Stoichiometry of Photosynthesis

2 H2O + 2 NADP+ + 10 H+stroma

O2 + 2 NADPH + 12 H+lumen