ENERGY CHAPTER 5

Norlizayati Ramlan

Learning Outcomes
1. Differentiate between autotrophy and heterotrophy based on their energy and carbon sources. 2. Discuss the relationship between an action spectrum and an absorption spectrum 3. Describe the photosynthetic pigments including the structure of a chloroplast.

Learning Outcomes
5. Explain the photoactivation of chlorophyll that converts light energy into ATP and NADP+. 6. Trace the movement of electrons in linear electron flow and cyclic electron flow. 7. Describe Calvin cycle that involves three stages.

Learning Outcomes
5. Discuss the role of ATP and NADPH. 6. Explain carbon fixation in C3, C4 and CAM plants. 7. Discuss the major consequences of photorespiration. 8. Describe two important photosynthetic adaptations that minimize photorespiration.

1. INTRODUCTION
ENERGY

Energy powers all life processes Photosynthesis The biological process that captures light energy and transforms it into chemical energy of organic molecule (e.g. carbohydrates) which are manufactured from CO2 and H2O. Cells use chemical energy stored in organic molecules to regenerate ATP, which powers work. Organism obtain organic compounds by one or two major modes: autotrophs & heterotrophs.

Light energy ECOSYSTEM

Photosynthesis in chloroplasts CO2 + H2O Cellular respiration in mitochondria

Organic +O molecules 2

ATP ATP powers most cellular work Heat energy

1. INTRODUCTION
AUTOTROPH

Autotroph auto self and trophos, nourishing. Definition: An organism that obtains organic food molecules without eating other organisms or substances derived from other organism . Produce their organic molecules from CO2 and other inorganic raw materials obtained from the environment.

1. INTRODUCTION
AUTOTROPH

Autotrophs can be separated by the source of energy that drives their metabolism. Photoautotrophs use light as a source of energy to synthesize organic compounds. Photosynthesis occurs in plants, algae, some other protists, and some prokaryotes. Chemoautotrophs harvest energy from oxidizing in organic substances, such as sulfur and ammonia. Chemoautotrophy is unique to prokaryotes.

1. INTRODUCTION
HETEROTROPH

Heterotrophs - heter, other and trophs, nourishing. Definition: an organism that obtains organic food molecules by eating other organisms or their byproducts. Cannot fix carbon; they use organic molecule produced by other organisms as the building blocks from which they synthesize the carbon compound they need.

1. INTRODUCTION
HETEROTROPH

These organisms are the consumers of the biosphere-completely dependent on photoautotrophs for food and for oxygen, a byproduct of photosynthesis. E.g animal feeds on other organism Other heterotrophs decompose and feed on dead organisms or on organic litter, like feces and fallen leaves.

2. PHOTOSYNTHESIS
IMPORTANCE OF PHOTOSYNTHESIS

The energy entering chloroplasts as sunlight gets stored as chemical energy in organic compounds. Sugar made in the chloroplasts supplies chemical energy and carbon skeletons to synthesize the organic molecules of cells. Plants store excess sugar as starch in structures such as roots, tubers, seeds, and fruits. In addition to food production, photosynthesis produces the O2 in our atmosphere

2. PHOTOSYNTHESIS
ABSORPTION SPECTRUM OF PHOTOSYNTHETIC PIGMENTS

Light is a form of electromagnetic energy, also called electromagnetic radiation. Like other electromagnetic energy, light travels in rhythmic waves. Wavelength is the distance between crests of waves. Wavelength determines the type of electromagnetic energy.

2. PHOTOSYNTHESIS
ABSORPTION SPECTRUM OF PHOTOSYNTHETIC PIGMENTS

The electromagnetic spectrum is the entire range of electromagnetic energy, or radiation Visible light consists of wavelengths (including those that drive photosynthesis) that produce colors we can see Light also behaves as though it consists of discrete particles, called photon

105 nm 103 nm
Gamma X-rays rays

1 nm UV

103 nm

106 nm

1m (109 nm)

103 m

Infrared

Microwaves

Radio waves

Visible light

380

450

500

550

600

650

700

750 nm

Shorter wavelength Higher energy

Longer wavelength Lower energy

2. PHOTOSYNTHESIS
THE PHOTOSYNTHETIC PIGMENTS

Pigments (chlorophyll a, b and carotenoids) are substances that absorb visible light. Different pigments absorb different wavelengths. Wavelengths that are not absorbed are reflected or transmitted. Leaves appear green because chlorophyll reflects and transmits green.

Light Reflected light Chloroplast

Absorbed light

Granum

Transmitted light

Absorption spectra = a graph plotting absorption vs spectra The three curves show the wavelengths of light best absorbed by three types of chloroplast pigments. The absorption of violet and red is maximum for chlorophyll a. Green is the least effective color.

2. PHOTOSYNTHESIS
THE PHOTOSYNTHETIC PIGMENTS

Chlorophyll a (blue-green), the dominant pigment, absorbs best in the red and violet-blue wavelengths and least in the green. Only chlorophyll a participates directly in the light reaction, but accessory photosynthetic pigments absorb light and transfer energy to chlorophyll a. Other pigments with different structures have different absorption spectra. Collectively, these photosynthetic pigments determine an overall action spectrum for photosynthesis.

2. PHOTOSYNTHESIS
THE PHOTOSYNTHETIC PIGMENTS

Chlorophyll b (yellow-green), with a slightly different structure than chlorophyll a, has a slightly different absorption spectrum and funnels the energy from these wavelengths to chlorophyll a. Carotenoids (yellow-orange) can funnel the energy from other wavelengths to chlorophyll a and also participate in photoprotection against excessive light. These compounds absorb and dissipate excessive light energy that would other wise damage chlorophyll. They also interact with oxygen to form reactive oxidative molecules that could damage the cell.

2. PHOTOSYNTHESIS
THE PHOTOSYNTHETIC PIGMENTS

2. PHOTOSYNTHESIS
THE PHOTOSYNTHETIC PIGMENTS

2. PHOTOSYNTHESIS
THE PHOTOSYNTHETIC PIGMENTS

When a pigment absorbs light, it goes from a ground state to an excited state, which is unstable. When excited electrons fall back to the ground state, photons are given off, an afterglow called fluorescence. an isolated solution of chlorophyll will fluoresce, giving off light and heat.
Note: photon is a quantum or discrete amount of light energy.

2. PHOTOSYNTHESIS
THE PHOTOSYNTHETIC PIGMENTS

2. PHOTOSYNTHESIS
CHLOROPLAST

Leaves are the major locations of photosynthesis. Their green color is from chlorophyll, the green pigment within chloroplasts. Light energy absorbed by chlorophyll drives the synthesis of organic molecules in the chloroplast. CO2 enters and O2 exits the leaf through microscopic pores called stomata.

2. PHOTOSYNTHESIS
CHLOROPLAST

Chloroplasts are found mainly in cells of the mesophyll, the interior tissue of the leaf. A typical mesophyll cell has 3040 chloroplasts. The chlorophyll is in the membranes of thylakoids (connected sacs in the chloroplast); thylakoids may be stacked in columns called grana. Chloroplasts also contain stroma, a dense fluid.

6 CO2 + 12 H2O + Light energy C6H12O6 + 6 O2 + 6 H2O

Chloroplasts split H2O into hydrogen and oxygen, incorporating the electrons of hydrogen into sugar molecules. Photosynthesis is a redox process in which H2O is oxidized and CO2 is reduced.

2. PHOTOSYNTHESIS
CHLOROPLAST

Photosynthesis consists of the light reactions (the photo part) and Calvin cycle (the synthesis part). The light reactions (in the thylakoids): Absorption of light energy and photoactivation of chlorophyll. ATPsynthesis through photophosphorylation Production ofNADPH (reduce NADP+ to NADPH) Release O2(from photolysis of water)-by product Thus, in light reaction two compounds will be produced: ATP & NADPH (will be used in Calvin Cycle)

2. PHOTOSYNTHESIS
CHLOROPLAST

Calvin cycle (the synthesis part) or light independent reaction (occur in stroma) none of the steps require light directly. The Calvin cycle: Forms sugar from CO2, using ATP and NADPH. The Calvin cycle begins with carbon fixation, incorporating CO2 into organic molecules. Thus, it is the Calvin cycle that makes sugar, but only with the help of ATP and NADPH form the light reaction.

Cooperation of the Light Reactions and the Calvin cycle

THE PHOTOSYSTEM

2. PHOTOSYNTHESIS
LIGHT REACTION

THE PHOTOSYSTEM

2. PHOTOSYNTHESIS
LIGHT REACTION

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) funnel the energy of photons to the reaction center. A primary electron acceptor in the reaction center accepts an excited electron from chlorophyll a.

THE PHOTOSYSTEM

2. PHOTOSYNTHESIS
LIGHT REACTION

Solar-powered transfer of an electron from a chlorophyll a molecule to the primary electron acceptor is the first step of the light reactions. Together, these light-harvesting complexes act like light- gathering antenna complexes for the reaction center.

THE PHOTOSYSTEM

2. PHOTOSYNTHESIS
LIGHT REACTION

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

LINEAR ELECTRON FLOW

2. PHOTOSYNTHESIS
LIGHT REACTION

The primary pathway. Involves both photosystems . produces ATP and NADPH by using light energy.

LINEAR ELECTRON FLOW

2. PHOTOSYNTHESIS
LIGHT REACTION

Primary acceptor
2 e

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.
Pigment molecules

P680 1 Light

Photosystem II (PS II)

LINEAR ELECTRON FLOW

2. PHOTOSYNTHESIS
LIGHT REACTION

Primary acceptor
H2O 3 e e P680 1 Light e 2

P680+ (P680 that is missing an electron) 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 Pigment molecules reaction.

2 H+ + 1/ O 2 2

Photosystem II (PS II)

LINEAR ELECTRON FLOW

2. PHOTOSYNTHESIS
LIGHT REACTION

Primary acceptor
H2O 3 e e P680 1 Light ATP 5 e 2

Pq
Cytochrome complex

2 H+
1/ 2

+ O2

Pc

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. 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.

LINEAR ELECTRON FLOW

2. PHOTOSYNTHESIS
LIGHT REACTION

Primary acceptor
H2O 3 e e P680 1 Light ATP 5 e 2

Primary acceptor
e

Pq
Cytochrome complex

2 H+
1/ 2

+ O2

Pc
P700 Light 6

Pigment molecules Photosystem II (PS II)

Photosystem I (PS I)

LINEAR ELECTRON FLOW

2. PHOTOSYNTHESIS
LIGHT REACTION

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.

LINEAR ELECTRON FLOW

2. PHOTOSYNTHESIS
LIGHT REACTION

Primary acceptor
H2O 3 e e P680 1 Light ATP 5 e 2

Primary acceptor
e

Pq
Cytochrome complex

Fd
e e 8 NADP+ reductase NADP+ + H+ NADPH P700 Light 6

2 H+ + 1/ O 2 2

Pc

Pigment molecules Photosystem II (PS II)

Photosystem I (PS I)

LINEAR ELECTRON FLOW

2. PHOTOSYNTHESIS
LIGHT REACTION

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. The electrons of NADPH are available for the reactions of the Calvin cycle.

ATP
e

e e

NADPH
e

Mill makes ATP

e

Photosystem II

Photosystem I

CYCLIC ELECTRON FLOW

2. PHOTOSYNTHESIS
LIGHT REACTION

PSI also absorbs light energy and excited electron are transport through an electron transport chain between PSI and PSII. Energy released is used for ATP synthesis. Electron pass along the electron carrier chain back to PSI. Cyclic electron flow uses only photosystem I and produces ATP, but not NADPH. Cyclic electron flow generates surplus ATP, satisfying the higher demand in the Calvin cycle.

CYCLIC ELECTRON FLOW

2. PHOTOSYNTHESIS
LIGHT REACTION

Primary acceptor Pq

Fd

Primary acceptor Fd NADP+ reductase NADP+ + H+ NADPH

Cytochrome complex Pc

Photosystem I Photosystem II ATP

CYCLIC ELECTRON FLOW

2. PHOTOSYNTHESIS
LIGHT REACTION

Photoexcited electrons from PS I are ocassionally shunted back from Ferredoxin (Fd) to chlorophyll via cytochrome complex and Plastocyanin (Pc). These electron shunt supplements the supply of ATP (via chemiosmosis) but produces no NADPH.

CHEMIOSMOSIS

2. PHOTOSYNTHESIS
LIGHT REACTION

Chemiosmosis: an energy coupling mechanism that uses energy that stored in the form of a H+ ion gradient across a membrane to drive cellular work such as synthesis of ATP. Osmosis refers to the flow of H+ across membrane. ATP synthase complex coupled the diffusion of H+ ion down their concentration gradient to the phosphorylation of ADP.

CHEMIOSMOSIS

2. PHOTOSYNTHESIS
LIGHT REACTION

CHEMIOSMOSIS

2. PHOTOSYNTHESIS
LIGHT REACTION

1. Water is split by PS II on the side facing thylakoid space. 2. As the plastoquinone (Pq), a mobile carriers, transfers electron to cytochrome complex, protons are translocated across the membrane into the thylakoid space. 3. H+ ion is removed from the stroma when it is taken up by NADP+

ATP SYNTHASE

2. PHOTOSYNTHESIS
LIGHT REACTION

An enzyme that makes ATP from ADP and inorganic phosphate. Uses energy from existing ion gradient to power ATP synthesis. Consists of 4 main parts A rotor: located in thylakoid membrane (inner membrane) protrudes into thylakoid space (mitochondrial matrix). A stator: anchor next to the rotor; holds knob stationary. A rod: extending from rotor into the knob. A knob: contain 3 catalytic sites.

ATP SYNTHASE

2. PHOTOSYNTHESIS
LIGHT REACTION

H+ flow down a narrow space between the stator and the rotor. It cause the rotor and rod to rotate (like waterwheel) The spinning rod cause the conformational change in the stationary knob. Three catalytic site in the knob is activated. ADP and inorganic phosphate is combined.

2. PHOTOSYNTHESIS
DARK REACTION

Where does it occur? What are the compounds needed from the light reaction process to be used in this cycle? What is the important gas to begin the process in Calvin cycle? What is produced in Calvin Cycle?

2. PHOTOSYNTHESIS
DARK REACTION

Most plant use the Calvin cycle to fix carbon. The Calvin Cycle is also known as C3pathway the 1st product is a 3 carbon sugar Carbon fixation occurs in stroma of chloroplast. The cycle spends thee energy of ATP and the reducing power of electrons carried by NADPH to make sugar. The Calvin cycle is anabolic, using energy to build sugar from smaller molecules.

2. PHOTOSYNTHESIS
DARK REACTION

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: 1. Carbon fixation (catalyzed by rubisco) 2. Reduction 3. Regeneration of the CO2 acceptor (RuBP)

PHASE 1: CARBON FIXATION

2. PHOTOSYNTHESIS
DARK REACTION

The Calvin cycle in corporates each CO2 molecule, one at a time by attaching it to a five carbon sugar named ribulose bisphosphate (RuBP). The reaction is catalyzed by Rubisco/RuBP carboxylase.

PHASE 1: CARBON FIXATION

2. PHOTOSYNTHESIS
DARK REACTION

The product of this reaction is an unstable six carbon intermediate, which immediately breaks down into two molecules of 3-phosphoglycerate (PGA) with three carbon each. The carbon that was originally part of CO2 molecule is now part of a carbon skeleton; the carbon has beenfixed. NOTE: glycerate 3-phosphate (GP)

Input

3 CO2

(Entering one at a time)

Phase 1: Carbon fixation

Rubisco

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

6 P 3-Phosphoglycerate

PHASE 2: REDUCTION

2. PHOTOSYNTHESIS
DARK REACTION

Each molecule of 3-phosphoglycerate receives an additional phosphate group from ATP, becoming 1,3-bisphosphoglycerate. A pair of electrons donated from NADPH reduces 1,3-bisphosphoglycerate (by losing a phosphate group), becoming glyceraldehyde-3phosphate(G3P). Specifically, electrons fron NADPH reduce carboxyl group of 3-phosphoglycerate to aldehyde group of G3P NOTE: Phosphoglyceraldehyde (PGAL)

Input

3 CO2

(Entering one at a time)

Phase 1: Carbon fixation

Rubisco

3 P Short-lived intermediate 3 P 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 Output G3P (a sugar)

Glucose and other organic compounds

PHASE 2: REDUCTION

2. PHOTOSYNTHESIS
DARK REACTION

From 6 molecules of G3P formed: - 1 molecule of G3P exits the cells to be used by plant cells (a net gain of carbohydrate). - another 5 molecules must be recycled to regenerate the 3 molecules of RuBP.

PHASE 3: REGENERATION OF RuBP

2. PHOTOSYNTHESIS
DARK REACTION

Five molecules of G3P are rearranged by the last steps of the Calvin Cycle into 3 molecules of RuBP. Another 3 ATP is used in this phase. The RuBP molecules can now begin the process of CO2 fixation.

Input

3 CO2

(Entering one at a time)

Phase 1: Carbon fixation

Rubisco

3 P Short-lived intermediate 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 Output G3P (a sugar)

Glucose and other organic compounds

PHASE 3: REGENERATION OF RuBP

2. PHOTOSYNTHESIS
DARK REACTION

For the net synthesis of G3P molecules (3 Calvin Cycle), the Calvin cycle consumes a total of: 3 molecules of CO2 3 times cycle 9 molecules of ATP 6 molecules of NADPH

2. PHOTOSYNTHESIS
C3 PLANTS

Most plants fix carbon dioxide via rubisco in Calvin Cycle (C3 pathway). Such plants are called C3 because the first organic product of carbon fixation is a 3 carbon compound (3 phosphoglycerate). E.g. Rice, wheat, soybeans Many C3 plants do not yield as much carbohydrates as might be expected especially during hot, dry days.

2. PHOTOSYNTHESIS
PHOTORESPIRATION

Photorespiration is a metabolic pathway that consumes oxygen, releases carbon dioxide, generates no ATP and decreases photosynthetic output. On hot, dry days, plants close their stomata to conserve water. Once the stomata close, photosynthesis rapidly uses up the CO2 remaining in the leaf and produces O2, which accumulates in chloroplast.

2. PHOTOSYNTHESIS
PHOTORESPIRATION

When chloroplast O2 level is higher than CO2, rubisco is more likely to catalyze the reaction of O2 with RuBP instead of CO2 (O2 and CO2 compete for active site of rubisco). When this occurs, some of the intermediates involved in the Calvin Cycle are degraded to CO2 and H2O.

2. PHOTOSYNTHESIS
PHOTORESPIRATION

This process is called photorespiration because: It occurs in the presence of light (photo) It requires O2, like aerobic respiration It produces CO2and H2O (respiration) Unlike normal respiration, this process produces no ATP; In fact, photorespiration consumes ATP. Unlike photosynthesis, photorespiration does not produce sugar; they decrease photosynthetic output.

2. PHOTOSYNTHESIS
PHOTORESPIRATION

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.

2. PHOTOSYNTHESIS
C4 PLANTS

C4 plants are so named because they form four carbon compound as its first product. Example: sugar cane, corn. Two type of photosynthetic cells: Bundle sheath cells: arranged into tightly packed sheaths around the vein of leaves. (Calvin cycle) Mesophyll cells: loosely arranged cells. (incorporating CO2 into PEP)

2. PHOTOSYNTHESIS
C4 PLANTS

This step requires the enzyme PEP carboxylase. PEP carboxylase has a higher affinity for CO2 than rubisco does. It has no affinity for O2 . it can fix CO2 even when CO2 concentrations are low (usually during hot and dry surrounding; stomata partially open)

2. PHOTOSYNTHESIS
C4 PLANTS

1. PEP carboxyase will add CO2 to phosphoenolpyruvate forming oxaloacetate (4C). 2. Oxaloacetate will be converted to malate. 3. Malate will be exported to bundle sheath cell through plasmodesmata. 4. Malate will release CO2 which will be fix to RuBP by Rubisco. 5. Pyruvate will be regenerated for conversion to PEP in the mesophyll cells.

1. In mesophyll cells, the enzyme PEP carboxylase adds carbon dioxide to PEP. 2. A four-carbon compound conveys the atoms of the CO2 into a bundle sheath cell via plasmodesmata

3. In bundle-sheath cells, CO2 is released and enters the Calvin cycle

2. PHOTOSYNTHESIS
CAM PLANTS

Crassulacean acid metabolism (CAM) is named after the family crassulaceae, the succulents in which the process was first discovered. Example: cactus, pineapples. They open stomata during night and closed them during day to conserve water; CO2 will be prevented from entering the leaves.

2. PHOTOSYNTHESIS
CAM PLANTS

1. The CO2 will be incorporated to PEP by PEP carboxylase producing Malate. The process occur in mesophyll cells during night. 2. malate will be stored in the vacuole in the form of organic acid (malic acid) until morning. 3. During the day, CO2 is released from organic acids to be used in Calvin cycle.

Sugarcane

C4
CO2

Pineapple

CAM
CO2 Night Organic acid

Mesophyll cell Organic acid

1 CO2 incorporated into four-carbon organic acids (carbon fixation)

Bundlesheath cell

CO2 Calvin Cycle Sugar 2 Organic acids release CO2 to Calvin cycle

CO2 Calvin Cycle Sugar

Day

(a) Spatial separation of steps

(b) Temporal separation of steps

Occur in the same time but different type of cells

Occur in the same cell but different times.