BIO 150 Metabolism and cell division Photosynthesis

LEARNING OBJECTIVES
By the end of this lecture you will be able to:
1. Understand that ENERGY can be transformed from one form to another. 2. Know that energy exist in two forms; free energy - available for doing work or as heat - a form unavailable for doing work. 3. Appreciate that the Sun provides most of the energy needed for life on Earth. 4. Explain why photosynthesis is so important to energy and material flow for life on earth. 5. Know why plants tend to be green in appearance. 6. Equate the organelle of photosynthesis in eukaryotes with the chloroplast. 7. Describe the organization of the chloroplast. 8. Understand that photosynthesis is a two fold process composed of the light-dependent reactions (i.e., light reactions) and the light independent reactions (i.e. Calvin Cycle or Dark Reactions). 9. Tell where the light reactions and the CO2 fixation reactions occur in the chloroplast. 10. Define chlorophylls giving their basic composition and structure. 11. Draw the absorption spectrum of chlorophyll and compare it to the action spectrum of photosynthesis. 12. Define the Reaction Centers and Antennae and describe how it operates. 13. Describe cyclic photophosphorylation of photosynthesis. 14. Describe noncyclic photophosphorylation of photosynthesis.

Energy can be transformed from one form to another

FREE ENERGY
(available for work)

vs. HEAT
(not available for work)

THE SUN: MAIN SOURCE OF ENERGY FOR LIFE ON EARTH

THE BASICS OF PHOTOSYNTHESIS

Almost all plants are photosynthetic autotrophs, as are some bacteria and protists
Autotrophs generate their own organic matter through photosynthesis Sunlight energy is transformed to energy stored in the form of chemical bonds

(c) Euglena (b) Kelp (a) Mosses, ferns, and flowering plants

(d) Cyanobacteria

Light Energy Harvested by Plants & Other Photosynthetic Autotrophs

6 CO2 + 6 H2O + light energy C6H12O6 + 6 O2

Light use in photosynthesis

Sun release energy in a broad spectrum of electromagnetic radiation. The energy of photon relates to its wavelength. Longer wavelength photons have low energy. Short wavelength photons contain high energy. Photosynthesis use 2 wavelength 450nm & 750nm

Electromagnetic Spectrum and Visible Light

Gamma rays X-rays UV Infrared & Microwaves Radio waves

Visible light

Wavelength (nm)

 When light strikes a leaf, 3 processes occur: Light is absorbed - use to drive photosynthesis., Light is reflected & light is transmitted give color to the object.

Why are plants green?

Transmitted light

WHY ARE PLANTS GREEN?

Different wavelengths of visible light are seen by the human eye as different colors.

Gamma rays

X-rays

UV

Infrared

Microwaves

Radio waves

Visible light

Wavelength (nm)

Pigments in chlorophyll
Chlorophyll is the main photosynthetic pigment in tylakoid membranes. Chlorophyll a a primary pigment absorbs blue violet and red wavelength. It initiates the light dependent reaction of photosynthesis. Chlorophyll b an accessory pigment absorbs blue and red orange wavelength.

 Carotenoids absorbs mainly blue green light and reflect mostly yellow, orange or red. Phycocyanin absorbs green and yellow and reflect blue or purple wavelength.

THE COLOR OF LIGHT SEEN IS THE COLOR NOT ABSORBED

Chloroplasts absorb light energy and convert it to chemical energy

Light

Reflected light

Absorbed light Transmitted light

Chloroplast

AN OVERVIEW OF PHOTOSYNTHESIS
Photosynthesis is the process by which autotrophic organisms use light energy to make sugar and oxygen gas from carbon dioxide and water

Carbon dioxide

Water PHOTOSYNTHESIS

Glucose

Oxygen gas

AN OVERVIEW OF PHOTOSYNTHESIS
The light reactions convert solar energy to chemical energy Produce ATP & NADPH
Light
Chloroplast NADP ADP +P Light reactions Calvin cycle

The Calvin cycle makes sugar from carbon dioxide ATP generated by the light reactions provides the energy for sugar synthesis The NADPH produced by the light reactions provides the electrons for the reduction of carbon dioxide to glucose

Chloroplasts: Sites of Photosynthesis

Photosynthesis
Occurs in chloroplasts, organelles in certain plants All green plant parts have chloroplasts and carry out photosynthesis
The leaves have the most chloroplasts The green color comes from chlorophyll in the chloroplasts The pigments absorb light energy

Photosynthesis occurs in chloroplasts In most plants, photosynthesis occurs primarily in the leaves, in the chloroplasts A chloroplast contains:
stroma, a fluid grana, stacks of thylakoids

The thylakoids contain chlorophyll

Chlorophyll is the green pigment that captures light for photosynthesis

Figure 10.4a

Leaf cross section Chloroplasts Vein

Mesophyll

Stomata Chloroplast

CO2

O2

Mesophyll cell

20 m

Figure 10.4b

Chloroplast

Outer membrane

Stroma

Thylakoid Granum Thylakoid space

Intermembrane space Inner membrane

1 m

The Two Stages of Photosynthesis: A Preview

Photosynthesis consists of the light reactions (the photo part) and Calvin cycle (the synthesis part) The light reactions (in the thylakoids)
Split H2O Release O2 Reduce NADP+ to NADPH Generate ATP from ADP by photophosphorylation

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 The Calvin cycle (in the stroma) forms sugar from CO2, using ATP and NADPH The Calvin cycle begins with carbon fixation, incorporating CO2 into organic molecules

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Figure 10.6-1

H2O

Light NADP ADP + Pi Light Reactions

Chloroplast

Figure 10.6-2

H2O

Light NADP ADP + Pi Light Reactions ATP

NADPH

Chloroplast O2

Figure 10.6-3

H2O

CO2

Light NADP ADP + Pi Light Reactions ATP

Calvin Cycle

NADPH

Chloroplast O2

Figure 10.6-4

H2O

CO2

Light NADP ADP + Pi Light Reactions ATP

Calvin Cycle

NADPH

Chloroplast O2 [CH2O] (sugar)

Stages in photosynthesis
Light reaction and dark reaction. Light reactions requires direct energy of light to make energy carrier molecules used in dark reaction. Light reaction occur in the grana and dark reaction take place in the stroma.

A Photosystem: A Reaction-Center Complex Associated with Light-Harvesting Complexes

A photosystem consists of a reaction-center complex (a type of protein complex) surrounded by light-harvesting complexes The light-harvesting complexes (pigment molecules bound to proteins) transfer the energy of photons to the reaction center

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Figure 10.13

Photosystem Photon Lightharvesting complexes Reactioncenter complex

STROMA Primary electron acceptor

Thylakoid membrane

Transfer of energy

Pigment Special pair of molecules chlorophyll a molecules THYLAKOID SPACE (INTERIOR OF THYLAKOID)

Thylakoid membrane

Chlorophyll

STROMA

Protein subunits (b) Structure of photosystem II

THYLAKOID SPACE

(a) How a photosystem harvests light

Figure 10.13a

Photosystem Photon Lightharvesting complexes Reactioncenter complex

STROMA Primary electron acceptor

Thylakoid membrane

Transfer of energy

Pigment Special pair of molecules chlorophyll a molecules THYLAKOID SPACE (INTERIOR OF THYLAKOID)

(a) How a photosystem harvests light

 A primary electron acceptor in the reaction center accepts excited electron and is reduced as a result Solar-powered transfer of an electron from a chlorophyll a molecule to the primary electron acceptor is the first step of the light reactions

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 There are two types of photosystems in the thylakoid membrane Photosystem II (PS II) functions first (the numbers reflect order of discovery) and is best at absorbing a wavelength of 680 nm The reaction-center chlorophyll a of PS II is called P680

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 Photosystem I (PS I) is best at absorbing a wavelength of 700 nm The reaction-center chlorophyll a of PS I is called P700

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Linear Electron Flow

During the light reactions, there are two possible routes for electron flow: cyclic and linear Linear electron flow, the primary pathway, involves both photosystems and produces ATP and NADPH using light energy

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 A photon hits a pigment and its energy is passed among pigment molecules until it excites P680 An excited electron from P680 is transferred to the primary electron acceptor (we now call it P680+)

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Figure 10.14-1

Primary acceptor e
2

P680
1 Light

Pigment molecules
Photosystem II (PS II)

 P680+ is a very strong oxidizing agent H2O is split by enzymes, and the electrons are transferred from the hydrogen atoms to P680+, thus reducing it to P680 O2 is released as a by-product of this reaction

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Figure 10.14-2

Primary acceptor H2O

3

2 H + 1/ O 2 2

e e P680

1 Light

Pigment molecules
Photosystem II (PS II)

 Each electron falls down an electron transport chain from the primary electron acceptor of PS II to PS I The electron transport chain consists of plastoquinone (Pq), cytochrome complex and plastocyanin (Pc). Energy released by the fall drives the creation of a proton gradient across the thylakoid membrane Diffusion of H+ (protons) across the membrane drives ATP synthesis
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Photophosphorylation
The released energy is harnessed by the thylakoids membrane to produce ATP. The ATP synthesis is called photophosporylation because it is driven by light energy.

Figure 10.14-3

Primary acceptor H2O

3

2
1/ 2

H + O2

Pq Cytochrome complex Pc
5

e e P680

1 Light

ATP

Pigment molecules
Photosystem II (PS II)

 In PS I (like PS II), transferred light energy excites P700, which loses an electron to an electron acceptor P700+ (P700 that is missing an electron) accepts an electron passed down from PS II via the electron transport chain

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Figure 10.14-4

Primary acceptor H2O

3

Primary acceptor e

2
1/ 2

H + O2

Pq Cytochrome complex Pc

e e P680
5

P700
Light
6

1 Light

ATP

Pigment molecules
Photosystem II (PS II)

Photosystem I (PS I)

 Each electron falls down an electron transport chain from the primary electron acceptor of PS I to the protein ferredoxin (Fd) The electrons are then transferred to NADP+ and reduce it to NADPH by NADP+ reductase. The electrons of NADPH are available for the reactions of the Calvin cycle This process also removes an H+ from the stroma

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Figure 10.14-5

Primary acceptor H2O

3

Primary acceptor e Fd e e

Pq Cytochrome complex Pc

+ 1/ O 2 2

NADP reductase

NADP + H NADPH

e e P680
5

P700
Light
6

1 Light

ATP

Pigment molecules
Photosystem II (PS II)

Photosystem I (PS I)

Figure 10.15

Mill makes ATP

e

NADPH
e

ATP Photosystem II Photosystem I

Cyclic Electron Flow

Cyclic electron flow uses only photosystem I (P700) and produces only ATP, but not NADPH. Occurs when cells require additional ATP and thre is no NADP+ to reduce to NADPH. No oxygen is released during this reaction. Cyclic electron flow generates surplus ATP, satisfying the higher demand in the Calvin cycle

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 Non-cyclic electron flow produces ATP and NADPH roughly in equal quantities but Calvin cycle consumes more ATP than NADPH. If ATP runs low, the Calvin cycle will slow down and NADPH will accumulate. The rise of NADPH will stimulate a temporary shift from noncyclic to cyclic electron flow until ATP supply is enough.

Figure 10.16

Primary acceptor

Fd

Primary acceptor Fd NADP + H NADPH Pc

Pq
Cytochrome complex

NADP reductase

Photosystem I Photosystem II ATP

A Comparison of Chemiosmosis in Chloroplasts and Mitochondria

Chloroplasts and mitochondria generate ATP by chemiosmosis, but use different sources of energy Mitochondria transfer chemical energy from food to ATP; chloroplasts transform light energy into the chemical energy of ATP Spatial organization of chemiosmosis differs between chloroplasts and mitochondria but also shows similarities
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 In mitochondria, protons are pumped to the intermembrane space and drive ATP synthesis as they diffuse back into the mitochondrial matrix In chloroplasts, protons are pumped into the thylakoid space and drive ATP synthesis as they diffuse back into the stroma

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Figure 10.17

Mitochondrion

Chloroplast

MITOCHONDRION STRUCTURE

CHLOROPLAST STRUCTURE H Electron transport chain ATP synthase Diffusion Thylakoid space Thylakoid membrane

Intermembrane space
Inner membrane

Matrix
ADP P i Key Higher Lower [H ] [H ] H ATP

Stroma

 ATP and NADPH are produced on the side facing the stroma, where the Calvin cycle takes place In summary, light reactions generate ATP and increase the potential energy of electrons by moving them from H2O to NADPH

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Figure 10.18

STROMA (low H concentration) Photosystem II Light

4 H+

Cytochrome complex Photosystem I Light

Fd Pq

NADP reductase 3

NADP + H

NADPH

H2O

2 1
1/ 2

Pc

O2 4 H+

THYLAKOID SPACE (high H concentration)

+2 H+

To Calvin Cycle
Thylakoid membrane STROMA (low H concentration)

ATP synthase

ADP + Pi

ATP H+

Concept 10.3: The Calvin cycle uses the chemical energy of ATP and NADPH to reduce CO2 to sugar
The Calvin cycle, like the citric acid cycle, regenerates its starting material after molecules enter and leave the cycle The cycle builds sugar from smaller molecules by using ATP and the reducing power of electrons carried by NADPH

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 Dark reaction occurs in stroma of the chloroplast and involves many reaction. Each reaction are catalyzed by different enzymes. This reaction do not require light. Both light reaction and dark reaction occur at the same time in chloroplasts during daylight. Each dependent upon each other.

 Carbon enters the cycle as CO2 and leaves as a sugar named glyceraldehyde 3-phospate (G3P) For net synthesis of 1 G3P, the cycle must take place three times, fixing 3 molecules of CO2 The Calvin cycle has three phases
Carbon fixation (catalyzed by rubisco) Reduction Regeneration of the CO2 acceptor (RuBP)

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Figure 10.19-1

Input 3 (Entering one CO2 at a time)

Phase 1: Carbon fixation

Rubisco 3 P Short-lived intermediate P 3P Ribulose bisphosphate (RuBP) P

6 P 3-Phosphoglycerate

Figure 10.19-2

Input 3 (Entering one CO2 at a time)

Phase 1: Carbon fixation

Rubisco 3 P Short-lived intermediate P 3P Ribulose bisphosphate (RuBP) P

6 P 3-Phosphoglycerate

6 6 ADP

ATP

Calvin Cycle

6 P P 1,3-Bisphosphoglycerate 6 NADPH 6 NADP 6 Pi

6 P Glyceraldehyde 3-phosphate (G3P)

Phase 2: Reduction

1 G3P (a sugar) Output

P Glucose and other organic compounds

Figure 10.19-3

Input 3 (Entering one CO2 at a time)

Phase 1: Carbon fixation

Rubisco 3 P Short-lived intermediate P 3P Ribulose bisphosphate (RuBP) P

6 P 3-Phosphoglycerate

6 6 ADP

ATP

3 ADP 3 ATP

Calvin Cycle

6 P P 1,3-Bisphosphoglycerate 6 NADPH

Phase 3: Regeneration of the CO2 acceptor (RuBP)

6 NADP 6 Pi 5 G3P P 6 P Glyceraldehyde 3-phosphate (G3P) Phase 2: Reduction

1 G3P (a sugar) Output

P Glucose and other organic compounds

Concept 10.4: Alternative mechanisms of carbon fixation have evolved in hot, arid climates
Dehydration is a problem for plants, sometimes requiring trade-offs with other metabolic processes, especially photosynthesis On hot, dry days, plants close stomata, which conserves H2O but also limits photosynthesis The closing of stomata reduces access to CO2 and causes O2 to build up These conditions favor an apparently wasteful process called photorespiration
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Photorespiration: An Evolutionary Relic?

In most plants (C3 plants), initial fixation of CO2, via rubisco, forms a three-carbon compound (3-phosphoglycerate) In photorespiration, rubisco adds O2 instead of CO2 in the Calvin cycle, producing a twocarbon compound Photorespiration consumes O2 and organic fuel and releases CO2 without producing ATP or sugar

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 Rubisco adds O2 to the Calvin cycle instead of CO2 The product split and two carbon compound formed called glycolic acid and exported from the chloroplast Mitochondria and perosixomes then break the glycolic acid to CO2.

 Photorespiration may be an evolutionary relic because rubisco first evolved at a time when the atmosphere had far less O2 and more CO2 Photorespiration limits damaging products of light reactions that build up in the absence of the Calvin cycle In many plants, photorespiration is a problem because on a hot, dry day it can drain as much as 50% of the carbon fixed by the Calvin cycle

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C4 Plants
C4 plants minimize the cost of photorespiration by incorporating CO2 into four-carbon compounds in mesophyll cells This step requires the enzyme PEP carboxylase PEP carboxylase has a higher affinity for CO2 than rubisco does; it can fix CO2 even when CO2 concentrations are low These four-carbon compounds are exported to bundle-sheath cells, where they release CO2 that is then used in the Calvin cycle
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 CO2 bound to the 3 carbon molecule known as phospoenol pyruvate (PEP) to form 4 carbon compounds called oxaloacetate This reaction catalyzed by enzyme PEP carboxylase Oxaloacetate then converted to the intermediate malate which was then sent to adjacent bundle sheath through plasmodesmata. Malate then is decarboxylated and dehydrogenated to form pvruvate and NADPH and CO2 is released.

 Bundle sheath cells are impermeable to CO2, result with a high concentration of CO2 within the bundle sheath cells The increase of CO2 concentration minimizes photorespiration and increase sugar production. The pyruvate goes back to the mesophyll cell and reacts with ATP to regenerate PEP and ready to accept another CO2

Figure 10.20

C4 leaf anatomy Mesophyll cell Photosynthetic cells of C4 Bundleplant leaf sheath cell Vein (vascular tissue)

The C4 pathway Mesophyll cell PEP carboxylase CO2

Oxaloacetate (4C) Malate (4C)

PEP (3C) ADP ATP

Stoma

Bundlesheath cell

Pyruvate (3C) CO2 Calvin Cycle Sugar

Vascular tissue

Figure 10.20a

C4 leaf anatomy
Photosynthetic cells of C4 plant leaf Mesophyll cell Bundlesheath cell

Vein (vascular tissue)

Stoma

Figure 10.20b

The C4 pathway Mesophyll cell PEP carboxylase CO2

Oxaloacetate (4C) Malate (4C)

PEP (3C) ADP

ATP
Pyruvate (3C)

Bundlesheath cell

CO2 Calvin Cycle Sugar

Vascular tissue

 In the last 150 years since the Industrial Revolution, CO2 levels have risen gratly Increasing levels of CO2 may affect C3 and C4 plants differently, perhaps changing relative abundance of these species The effects of such changes are unpredictable and a cause for concern

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Advantages of C4 pathway functions in addition to C3 pathway

Photosynthesis can progress even at a very low CO2 concentration. It be able to store CO2 temporarily so that it can be used later. Be able to avoid photorespiration. Photorespiration is a wasteful process and reduces their photosynthetic efficiency.

CAM Plants
Some plants, including succulents, use crassulacean acid metabolism (CAM) to fix carbon CAM plants open their stomata at night, incorporating CO2 into organic acids Stomata close during the day, and CO2 is released from organic acids and used in the Calvin cycle

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 Develop a special carbon fixation pathway known as CAM. At night when the stomata open, CAM plants use the enzyme PEP carboxylase to fix CO2 and turn to oxaloacetate. The oxaloacetate then converted to malate and stored in cell vacuole. During day time, exchange of gas cannot occur because of the closing stomata, CO2 then taken from malate by decarboxylation process.

Figure 10.21

Sugarcane
C4 Mesophyll Organic acid cell CO2 Bundlesheath cell Calvin Cycle Sugar (a) Spatial separation of steps CO2

Pineapple CAM 1 CO2 incorporated (carbon fixation) Organic acid CO2 Night

CO2 2 CO2 released to the Calvin cycle Calvin Cycle Sugar (b) Temporal separation of steps Day

Figure 10.21a

Sugarcane

Figure 10.21b

Pineapple

Features

C3 plant

C4 plant

CAM plant

Representative spp (e.g)

Efficiency Carbon fixation Site for carbon (C) fixation and C.cycle

Rice, wheat, soybean

Maize, sugarcane

Cactus, pineapple

Less than C4 and CAM Once Mesophyll cells

More than C3 Twice Initial carbon fixation mesophyll cells Calvin cycle bundle sheath cells RuBP, PEP

More than C3 Twice Both occur in mesophyll cells but at different period of time RuBP, PEP

CO2 acceptor

RuBP

CO2 fixing enzyme

Rubisco

Rubisco, PEP carboxylase

Low High High

Rubisco, PEP carboxylase

Low High High

Rate of photorespiration Rate of photosynthesis Yields

High High Low

Factors affecting photosynthesis process

Light intensity Concentration of carbon dioxide Temperature Concentration of chlorophyll Concentration of oxygen Water Chemical compounds

Autotrophic prokaryotes
Prokaryotes consist of bacteria and the cynobacteria Certain prokaryotes are autotrophic able to make their own complex organic compounds as food.

 Cynobacteria photosynthetic organisms. They use CO2, water and energy from sun to produce sugar and other substance without the presence of chloroplast but use the photosynthetic pigment located in membrane system in the cytoplasm. Autotrophic bacteria
Photosynthetic bacteria Chemosynthetic bacteria

Photosynthetic bacteria
Use pigment known as bacteriochlorophyll to capture light This pigment is similar as chlorophyll found in plants The source of hydrogen to reduce CO2 does not come from water but from other molecule. Purple sulphur bacteria use hydrogen sulphide as a source of hydogen.

Chemosynthetic bacteria
This type of bacteria produce complex organic compound from CO2 and water but the chemical energy is not coming from light but from specific chemical reaction which they able to catalyze. Nitrifying bacteria like Nitrosomonas and Nitrococcus get its energy from oxidation of ammonia to nitrite in the nitrification process in nitrogen cycle.

It's not that easy bein' green but it is essential for life on earth!