CHAPTER 8 PHOTOSYNTHESIS

Prepared by

Brenda Leady, University of Toledo

Copyright (c) The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

PHOTOSYNTHESIS
SWBAT:

Describe overall equation of photosynthesis Distinguish: autotrophs, heterotrophs, photoautotrophs and chemoautotrophs Describe stages of photosynthesis (light-dependent reactions and Calvin Cycle) Outline chemiosmosis in photophosphorilation Describe alternative ways of Carbon fixation (C4 and CAM plants)

PHOTOSYNTHESIS
SWBAT:

Describe overall equation of photosynthesis Distinguish: autotrophs, heterotrophs, photoautotrophs and chemoautotrophs Describe stages of photosynthesis (light-dependent reactions and Calvin Cycle) Outline chemiosmosis in photophosphorilation Describe alternative ways of Carbon fixation (C4 and CAM plants)

Trophic organization

Heterotroph
Must

eat food, organic molecules from their environment, to sustain life organic molecules from inorganic sources

Autotroph
Make

Photoautotroph
Use light as a source of energy Green plants, algae, cyanobacteria

Photosynthesis

Energy within light is captured and used to synthesize carbohydrates

CO2 + H2O + light energy C6H12O6 + O2 + H2O

CO2 is reduced H2O is oxidized Energy from light drives this endergonic reaction

Link between Photosynthesis and Respiration:

Who does it?

Plants Algae

Cyanobacteria
Photosynthetic bacteria

Biosphere
Regions on the surface of the Earth and in the atmosphere where living organisms exist Largely driven by the photosynthetic power of green plants Cycle where cells use organic molecules for energy and plants replenish those molecules using photosynthesis

Plants

also produce oxygen

Chloroplast
Organelles in plants and algae that carry out photosynthesis Chlorophyll- green pigment Majority of photosynthesis occurs in leaves in central mesophyll Stomata- carbon dioxide enters and oxygen exits leaf

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Chloroplast anatomy
Outer and inner membrane Intermembrane space Thylakoid membrane contains pigment molecules Thylakoid membrane forms thylakoids Enclose thylakoid lumen Granum- stack of thylakoids Stroma- fluid filled region between thylakoid membrane and inner membrane

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2 stages of photosynthesis

Light reactions
Take

place in thylakoid membranes Produce ATP, NADPH and O2

Calvin cycle
Occurs

in stroma Uses ATP and NADPH to incorporate CO2 into organic molecules
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Light energy
Type of electromagnetic radiation Travels as waves

Short

to long wavelengths wavelengths have more energy

Also behaves as particles- photons

Shorter

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Photosynthetic pigments absorb some light energy and reflect others

Leaves

are green because they reflect green wavelengths

Absorption boosts electrons to higher energy levels Wavelength of light that a pigment absorbs depends on the amount of energy needed to boost an electron to a higher orbital

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After an electron absorbs energy, it is an excited state and usually unstable Releases energy as

Heat Light

Excited electrons in pigments can be transferred to another molecule or captured Captured light energy can be transferred to other molecules to ultimately produce energy intermediates for cellular work

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Pigments

Chlorophyll a Chlorophyll b Carotenoids

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Spectrophotometers are used to measure light absorption

White light

Refracting prism

Chlorophyll solution Photoelectric tube

Slit moves to pass light of selected wavelength

Blue light

The low transmittance (high absorption) reading indicates that chlorophyll absorbs most blue light.

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Absorption vs. action spectrum

Absorption spectrum
Wavelengths that are absorbed by different pigments in the plant

Action

spectrum

Rate of photosynthesis by whole plant at specific wavelengths

The color of the pigment comes from the wavelengths of light reflected (in other words, those not absorbed). Chlorophyll, the green pigment common to all photosynthetic cells, absorbs all wavelengths of visible light except green
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The color of the pigment comes from the wavelengths of light reflected (in other words, those not absorbed). Chlorophyll, the green pigment common to all photosynthetic cells, absorbs all wavelengths of visible light except green

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Photosystems

Thylakoid membrane:

Photosystem I (PSI) o Photosystem II (PSII) Each of them has a light harvesting complex (antenna complex) and a reaction center
o

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Photosytem II (PSII)

2 main components
Light-harvesting complex or antenna complex Directly absorbs photons Energy transferred via resonance energy transfer to P680 in the reaction center Reaction center P680 P680* (Relatively unstable) P680* actually releases its high-energy electron to the primary electron acceptor and becomes P680+(more stable). P680 has to be regenerated, by replacing the electron so P680 can work again: : This missing electron of P680+ is replaced with a low-energy electron from water which yields oxygen gas

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Photosystem II (PSII)
Redox machine Recent research in biochemical composition of protein complex and role of components 3 dimensional structure determined in 2004 using x-ray crystallography

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D1 and D2, contain the reaction center that carries out the redox reactions CP43 and CP47, bind the pigment molecules that form the light-harvesting complex wrap around D1 and D2 so that pigments in CP43 and CP47 can transfer energy to P680 by resonance energy transfer. Pp: primary electron acceptor: a chlorophyll molecule lacking Mg2+, called pheophytin (Pp). QA:plastoquinone molecule, designated QA QB another plastoquinone molecule which can accept two high-energy electrons and bind two H+. QB can diffuse away from the reaction center.
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The oxidation of water occurs in a region called the manganese cluster. This site is located on the side of D1 that faces the thylakoid lumen. The manganese cluster has four Mn2+, one Ca2+, and one Cl. Two water molecules bind to this site. D1 catalyzes the removal of four low-energy electrons from the two water molecules to create four H+ and O2. Each low-energy electron is transferred, one at a time, to an amino acid in D1 (a tyrosine, Tyr) and then to P680+ to produce P680. 30

Electrons accepted by primary electron acceptor in PSII are transferred to a pigment molecule in the reaction center of PSI Electron releases some of its energy along the way
H+ electrochemical gradient ATP synthesis uses chemiosmotic mechanism similar to mitochondria
Establishes

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Photosystem I (PSI)
Key role to make NADPH Light striking light-harvesting complex of PSI transfers energy to a reaction center High energy electron removed from P700 and transferred to a primary electron acceptor NADP+ reductase

NADP+ + 2 electrons + H + P700+ replaces its electrons

NADPH

from
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plastocyanin
No

splitting water, no oxygen gas formed

Summary
1. 2.

3.

O2 produced in thylakoid lumen by oxidation of H2O by PSII 2 electrons transferred to P680+ ATP produced in stroma by H+ electrochemical gradient (chemiosmosis) due to: 1. Splitting of water places H+ in the lumen 2. High-energy electrons move from PSII to PSI, pumping H+ into the lumen 3. Formation of NADPH consumes H+ in the stroma NADPH produced in the stroma from high-energy electrons that start in PSII and boosted in PSI NADP+ + 2 electrons + H + NADPH
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From FSII to FSI

QB can diffuse away from the reaction center PSII carrying a pair of electrons Each electron enters an electron transport chain: a series of electron carriers located in the thylakoid membrane. The electron transport chain functions similarly to the one found in mitochondria. The electrons go from QB, to a cytochrome complex, then to plastocyanin (Pc), a small protein; and then to PSI Along its journey from PSII to PSI, the electron releases some of its energy at particular steps and is transferred to the next component that has a higher electronegativity. In the cyochrome complex the energy released is harnessed to pump H+ into the thylakoid lumen. One result of the electron movement is to establish a H+ electrochemical gradient
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FSI
When light strikes the light-harvesting complex of PSI, this energy is also transferred to a reaction center, where a high-energy electron is removed from a pigment molecule, designated P700, and transferred to a primary electron acceptor. A protein called ferredoxin (Fd) can accept two high-energy electrons, one at a time, from the primary electron acceptor. Fd then transfers the two electrons to the enzyme NADP+ reductase. This enzyme transfers the two electrons to NADP+ and together with a H+ creates NADPH. The formation of NADPH results in fewer H+ in the stroma and thereby contributes to the formation of a H + electrochemical gradient across the 35 thylakoid membrane.

Chemiosmosis
Synthesis of ATP in chloroplasts is achieved by a chemiosmotic mechanism similar to that used to make ATP in mitochondria. ATP synthesis is driven by the flow of H+ from the thylakoid lumen into the stroma via ATP synthase + A H gradient is generated in three ways: (1) the splitting of water, which places H+ in the thylakoid lumen; (2) the movement of high-energy electrons from photosystem II to photosystem I, which pumps H + into the thylakoid lumen 36 (3) the formation of NADPH, which consumes H+ in the stroma.

Cyclic and noncyclic electron flow

Noncyclic
Electrons

begin at PSII and eventually transfer to NADPH Linear process produces ATP and NADPH in equal amounts

Cyclic photophosphorylation
Electron

cycling releases energy to transport H+ into lumen driving synthesis of ATP

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Cyclic

photophosphorylation

cycling releases energy to transport H+ into lumen driving synthesis of ATP. NADPH IS NOT produced H2O is not splitter and O2 is not produced 38
Electron

Photosystems II and I work together to produce ATP and NADPH

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Z scheme (energy curve)

Robin Hill and Fay Bendall

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Photosynthesis animations

Photosynthetic Electron Transport and ATP Synthesis

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The cytochrome complexes of mitochondria and chloroplasts have evolutionarily related proteins in common

Homologous genes are similar because they are derived from a common ancestor Comparing the electron transport chains of mitochondria and chloroplasts reveals homologous genes Family of cytochrome b-type proteins plays similar but specialized roles

Calvin cycle
ATP and NADPH used to make carbohydrates Somewhat similar to citric acid cycle CO2 incorporated into carbohydrates

Precursors

to all organic molecules Energy storage

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Overview of Calvin Cycle

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CO2 incorporation

Also called Calvin-Benson cycle Requires massive input of energy: For every 6 CO2 incorporated, 18 ATP and 12 NADPH used Glucose is not directly made. Instead, molecules of glyceraldehyde-3-phosphate, which are products of the Calvin cycle, are used as starting materials for the synthesis of glucose and other molecules, including sucrose. After glucose molecules are
made, they may be linked together to form a polymer of glucose called starch, which is stored in the chloroplast for later use. Alternatively, the disaccharide sucrose may be made and transported out of the leaf to other parts of the plant.

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3 phases
1.

Carbon fixation

CO2 incorporated in ribulose bisphosphate (RuBP) using RuBP carboxylase/oxygenase (rubisco) 6 carbon intermediate splits into 2 3PG

2.

Reduction and carbohydrate production

ATP is used to convert 3 phosphoglycerate (3PG) into 1,3-bisphosphoglycerate (1,3 BPG) NADPH electrons reduce it to glyceraldehyde 3 P (G3P) 6 CO2 12 G3P

2 for carbohydrates 10 for regeneration

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3.

Regeneration of RuBP

10 G3P converted into 6 RuBP using 6 ATP

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Photosynthesis animations

calvinCycle.swf

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The Calvin cycle was determined by isotope labeling methods

14C-labeled

algae Allowed to incubate different lengths of time Separated newly made radiolabeled molecules using two-dimensional paper chromatography Autoradiography- radiation from 14C-labeled molecules makes dark spots on the film Identified 14C-labeled spots and the order they appeared Calvin awarded Nobel Prize in 1961

CO2 injected into cultures of green

Variations in photosynthesis

Certain environmental conditions can influence both the efficiency and way the Calvin cycle works
Light

intensity Temperature Water availability

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Photorespiration
Rubisco: RuBP carboxylase/oxygenase

RuBP + CO2 2 3PG

Rubisco functions as C3 plants make 3PG

a carboxylase

Rubisco can also be an oxygenase

Adds O2 to RuBP eventually releasing CO2 Photorespiration Using O2 and liberating CO2 is wasteful

More likely in hot and dry environment Favored when CO2 low and O2 high

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C3 plants
90% of plants are C3

Wheat plants

Oak leaves

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C4 plants: To avoid photorespiration

C4 plants make a 4 carbon compound in the first step of carbon fixation Hatch-Slack pathway Leaves have 2-cell layer organization

Mesophyll

cells

CO2 enters via stomata and 4 carbon compound formed (PEP carboxylase does not promote photorespiration)

Bundle-sheath

cells

4 carbon compound transferred that releases steady supply CO2

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C4 plants

In warm and dry climates, C4 plants have an advantage. During the day, they can keep their stomata partially closed to conserve water. Furthermore, they can avoid photorespiration. C4 plants are well adapted to habitats with high daytime temperatures and intense sunlight. 56 Examples of C4 plants are sugarcane, crabgrass, and corn.

CAM plants

Some C4 plants separate processes using time Crassulacean Acid Metabolism CAM plants open their stomata at night CO2 enters and is converted to malate Stomata close during the day to conserve water Malate broken down into CO2 to drive Calvin cycle

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C4

CAM

Process in separated cells

Process at different times

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Adaptations for hot weather: C4 and CAM plants

C4: corn, sugarcane and sorghum

CAM: succulents (aloe, jade), pineapple, cacti CAM = crassulasean acid metabolism
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C4 and CAM compared

Both fix CO2 into a C4 compound Then CO2 is transferred to the Calvin cycle

In C4 plants there is a spatial separation (2 cell types)

In CAM plants there is a temporal separation (C4 accumulates at night, Calvin cycle during the day)
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What is better? C3 or C4 strategy?

In warm dry climates C4 plants have the advantage in conserving water and preventing photorespiration

In cooler climates, C3 plants use less energy to fix CO2 90% of plants are C3

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