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Biology 11- Lecture 4- Photosynthesis

Photosynthesis

This handout is for lecture use only and not


for commercial reproduction and
distribution.

Autotrophs

Heterotrophs

producers of the biosphere, producing organic


molecules from CO2 and other inorganic
molecules
photoautotrophs - use the energy of sunlight to
make organic molecules from H2O and CO2
feed not only themselves but also most of the
living world

obtain their organic material from other


organisms
consumers of the biosphere
almost all heterotrophs depend on
photoautotrophs for food and O2

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Photosynthesis
process that converts solar energy into chemical
energy
directly or indirectly nourishes almost the entire
living world
occurs in plants, algae, certain other protists,
and some prokaryotes

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Biology 11- Lecture 4- Photosynthesis

Chloroplasts: The Sites of


Photosynthesis in Plants
leaves - major locations of photosynthesis
chlorophyll - 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

(a) Plants

(c) Unicellular protist

10 m

(e) Purple sulfur


bacteria

(b) Multicellular alga

(d) Cyanobacteria

1.5 m

40 m
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Chloroplast

Leaf cross section


Vein

chlorophyll is in
the membranes
of thylakoids

Mesophyll

Stomata

Chloroplast

CO2

Outer
membrane

chloroplasts also
contain stroma,
a dense fluid

O2

Mesophyll cell

Thylakoid
Stroma

Granum

Thylakoid
space

1 m

5 m

Photosynthesis
summary equation:

Intermembrane
space
Inner
membrane

Photosynthesis
summary equation:
6 CO2 + 12 H2O + Light energy C6H12O6 + 6 O2 + 6 H2O

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

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Word level
Carbon Dioxide + Water + Light Energy Glucose + Oxygen + Water

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Biology 11- Lecture 4- Photosynthesis

Two Stages of Photosynthesis


Reactants:

Products:

12 H2O

6 CO2

C6H12O6

6 H2O

6 O2

chloroplasts split H2O into hydrogen and oxygen,


incorporating the electrons of hydrogen into sugar
molecules

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

photosynthesis is a redox process in which H2O is


oxidized and CO2 is reduced
Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Two Stages of Photosynthesis


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

CO2

H 2O

Calvin cycle begins with


carbon fixation,
incorporating CO2 into
organic molecules

Light
NADP+
ADP
+ P

Light
Reactions

Calvin
Cycle

ATP
NADPH

Chloroplast
O2

[CH2O]
(sugar)

Calvin cycle (in the


stroma) forms
sugar from CO2,
using ATP and
NADPH

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Nature of Sunlight

The Nature of Sunlight

light is a form of electromagnetic energy, also


called electromagnetic radiation (travels in
rhythmic waves)
wavelength determines the type of
electromagnetic energy

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 photons

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Biology 11- Lecture 4- Photosynthesis

105 nm 103 nm

103 nm

1 nm

Gamma
X-rays
rays

UV

106 nm

1m
(109 nm)

Microwaves

Infrared

103 m

Radio
waves

Photosynthetic Pigments

substances that absorb visible light


different pigments absorb different wavelengths
chlorophyll a - main photosynthetic pigment
accessory pigments (chlorophyll b), broaden the
spectrum used for photosynthesis

Visible light

380

450

500

550

600

650

700

750 nm

Longer wavelength
Lower energy

Shorter wavelength
Higher energy
Electromagnetic Spectrum

The chlorophyll molecules


of chloroplasts absorb
VioIetblue and red light
(the colors most effective
in driving
Chloroplast
photosynthesis) and
reflect or transmit light.
This is why leaves
appear green.

wavelengths that
are not absorbed
are reflected or
transmitted

carotenoids absorb excessive light that would


damage chlorophyll

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Light
Reflected
light

Absorbed
light

Granum

Transmitted
light

CH3
CHO

(methyl) in chlorophyll a
(aldehyde/carbonyl) in chlorophyll b

an absorption spectrum is a graph plotting a pigments light


absorption versus wavelength
the absorption spectrum of chlorophyll a suggests that violetblue and red light work best for photosynthesis

Photosystem

Porphyrin ring:
light-absorbing
head of molecule;
note magnesium
atom at center

Structure of
chlorophyll molecules
in chloroplasts of plants

Magnesium is bound as the


central atom of the porphyrin
ring of the green plant pigment
chlorophyll. Magnesium is the
element that causes plants to be
able to convert light into energy.

Hydrocarbon tail:
interacts with hydrophobic
regions of proteins inside
thylakoid membranes of
chloroplasts; H atoms not
shown

Biology 11- Lecture 4- Photosynthesis

Photosystem

Photosystem

consists of a reaction-center complex


surrounded by light-harvesting complexes
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

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Photosystems

Linear Electron Flow

PS II functions first and is best at absorbing a


wavelength of 680 nm
the reaction-center chlorophyll a of PS II is called
P680

PS I is best at absorbing a wavelength of 700 nm


the reaction-center chlorophyll a of PS I is called
P700

Each photoexcited electron


passes from the primary
electron acceptor of PS II to PS I
via an electron transport chain
[electron carrier plastoquinone
(Pq), a cytochrome complex,
and a protein called
plastocyanin (Pc).]

Electrons from water


split are supplied one
by one to the P680+
pair, each electron
replacing one
transferred to
the primary electron
acceptor

2 H+
+
1/ O
2
2

Primary
acceptor

3
e
e

Electron is
transferred to
primary
electron
acceptor

Enzyme NADP+ reductase


catalyzes the transfer of
electrons from Fd to
NADP+ for reduction to
NADPH to be used in Calvin
cycle.

NADP+
reductase

Pc
5

P680

Electrons at a lower
energy level
provide energy for
the synthesis of ATP

Pigment
molecules

NADP+
+ H+
NADPH

P700
Light
6

ATP

Photosystem II
(PS II)

Fd

Cytochrome
complex

1 Light
Photon
energizes
pigment
molecule
and
electrons
and then
relayed up
to P680

Primary
acceptor
e

Pq

2
e

H2O

Photoexcited electrons are


passed to a second
electron transport chain
through the protein
ferredoxin (Fd).

Electron is
photoexcited and
transferred to PSI's
primary electron
acceptor

Photosystem I
(PS I)

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Cyclic Electron Flow


uses only photosystem I and produces ATP, but not NADPH
generates surplus ATP, satisfying the higher demand in the Calvin
cycle

Biology 11- Lecture 4- Photosynthesis

Chemiosmosis: Chloroplasts vs Mitochondria

Chemiosmosis - generation of ATP in


chloroplasts and mitochondria
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

Comparison of
chemiosmosis in
mitochondria and
chloroplasts. In both
kinds of organelles,
electron transport
chains pump protons
(H+) across a membrane
from a
region of low H+
concentration to one of
high H+ concentration.
The protons then diffuse
back across
the membrane through
ATP synthase, driving
the synthesis of ATP.

Mitochondrion

Chloroplast

MITOCHONDRION
STRUCTURE

CHLOROPLAST
STRUCTURE
H+

Intermembrane
space
Inner
membrane

Diffusion

Electron
transport
chain

Thylakoid
space
Thylakoid
membrane

ATP
synthase
Key

Stroma

Matrix
ADP + P

Higher [H+]
Lower [H+]

H+

ATP

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Calvin cycle
STROMA
(low H+ concentration)

Cytochrome
complex

Photosystem II
4 H+

Light

Photosystem I
Light
Fd

NADP+
reductase

H2O

e
1

THYLAKOID SPACE
(high H+ concentration)

e
2

carbon enters the cycle as CO2 and leaves as a sugar


named glyceraldehyde-3-phospate (G3P)

Pc

2
1/

NADP+ + H+

NADPH

Pq

O2
4 H+

+2 H+

To
Calvin
Cycle
Thylakoid
membrane

three phases:
Carbon fixation (catalyzed by rubisco)

ATP
synthase

STROMA
(low H+ concentration)

builds sugar from smaller molecules by using


ATP and the reducing power of electrons carried
by NADPH

ADP
+
Pi

Reduction

ATP
H+

Light reactions and chemiosmosis: the organization of the thylakoid membrane. Storing energy as a proton-motive force (H+
gradient , attributed by 1) Water is split by photosystem II on the side of the membrane facing tile thylakoid space; 2) as plastoquinone (Pq), a
mobile carrier, transfers electrons to the cytochrome complex, four protons are translocated across the membrane into the thylakoid space; 3)
a hydrogen ion is removed from the stroma when it is taken up by NADP+.

Regeneration of the CO2 acceptor (RuBP)


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Input

The Calvin cycle

3
CO2

(Entering one
at a time)

Incorporation of each CO2


molecule, one at a time, by
attaching it to a five carbon sugar
named ribulose bisphosphate
(abbreviated RuBP).RuBP
carboxylase, or rubisco catalyzes
this first step. Product of the
reaction is a six-carbon
intermediate that splits in half,
forming two molecules of 3phosphoglycerate

Phase 1: Carbon fixation


Carbon skeletons of five
molecules of G3P are
rearranged by the last steps
of the Calvin cycle into
three molecules of RuBP.
To accomplish this, the
cycle spends three more
molecules of ATP. The
RuBP is now prepared to
receive CO2 again, and the
cycle continues.

Rubisco

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

6
P
3-Phosphoglycerate

ATP

6 ADP

3 ADP
3

Calvin
Cycle

6 P
P
1,3-Bisphosphoglycerate

ATP

6 NADPH
Phase 3:
Regeneration of
the CO2 acceptor
(RuBP)

6 NADP+
6 Pi
P

5
G3P

6
P
Glyceraldehyde-3-phosphate
(G3P)

1
Output

P
G3P
(a sugar)

Phase 2:
Reduction

Each molecule of 3phosphoglycerate receives an


additional phosphate group
from ATP, becoming
1,3bisphosphoglycerate. Next, a
pair of electrons donated from
NADPH reduces 1,3bisphosphoglycerate, which also
loses a phosphate group,
becoming G3P. One molecule
exits the cycle to be used by the
plant cell, but the other five
molecules must be recycled to
regenerate the three molecules
of RuB.

Glucose and
other organic
compounds

Copyright 2002 Pearson Education, Inc., publishing as Benjamin Cummings

Biology 11- Lecture 4- Photosynthesis

For the net synthesis of one G3P molecule, the


Calvin recycle consumes nine ATP and six
NAPDH.
The G3P from the Calvin cycle is the starting
material for metabolic pathways that
synthesize other organic compounds, including
glucose and other carbohydrates.

Copyright 2002 Pearson Education, Inc., publishing as Benjamin Cummings

In most plants (C3 plants) initial fixation of CO2


occurs via rubisco and results in a three-carbon
compound, 3-phosphoglycerate.
These plants include rice, wheat, and soybeans.

While rubisco normally accepts CO2, when the


O2/CO2 ratio increases (on a hot, dry day with
closed stomata), rubisco can add O2 to RuBP.

Alternative mechanisms of carbon


fixation have evolved in hot, arid climates
One of the major problems facing terrestrial plants is
dehydration.
The stomata are not only the major route for gas
exchange (CO2 in and O2 out), but also for the
evaporative loss of water.
On hot, dry days plants close the stomata to conserve
water, but this causes problems for photosynthesis.

Copyright 2002 Pearson Education, Inc., publishing as Benjamin Cummings

When rubisco adds O2 to RuBP, RuBP splits into


a three-carbon piece and a two-carbon piece in
a process called photorespiration.
The two-carbon fragment is exported from the
chloroplast and degraded to CO2 by mitochondria
and peroxisomes.
Unlike normal respiration, this process produces no
ATP, nor additional organic molecules.

Photorespiration decreases photosynthetic


output by siphoning organic material from the
Calvin cycle.

Copyright 2002 Pearson Education, Inc., publishing as Benjamin Cummings

Copyright 2002 Pearson Education, Inc., publishing as Benjamin Cummings

The C4 plants fix CO2 first in a four-carbon compound.


Several thousand plants, including sugarcane and corn,
use this pathway.
In C4 plants, mesophyll cells incorporate CO2 into organic
molecules.
The key enzyme, phosphoenolpyruvate carboxylase,
adds CO2 to phosphoenolpyruvate (PEP) to form
oxaloacetetate.
PEP carboxylase has a very high affinity for CO2 and can
fix CO2 efficiently when rubisco cannot - on hot, dry
days with the stomata closed.

Copyright 2002 Pearson Education, Inc., publishing as Benjamin Cummings

Copyright 2002 Pearson Education, Inc., publishing as Benjamin Cummings

Biology 11- Lecture 4- Photosynthesis

In effect, the mesophyll cells pump CO2 into the


bundle sheath cells, keeping CO2 levels high
enough for rubisco to accept CO2 and not O2.
C4 photosynthesis minimizes photorespiration
and enhances sugar production.
C4 plants thrive in hot regions with intense
sunlight.

Copyright 2002 Pearson Education, Inc., publishing as Benjamin Cummings

A second strategy to minimize photorespiration is found in


succulent plants, cacti, pineapples, and several other plant
families.
These plants, known as CAM plants for crassulacean
acid metabolism (CAM), open stomata during the night
and close them during the day.
During the night, these plants fix CO2 into a variety of
organic acids in mesophyll cells.
During the day, the light reactions supply ATP and
NADPH to the Calvin cycle and CO2 is released from the
organic acids.

Copyright 2002 Pearson Education, Inc., publishing as Benjamin Cummings

Photosynthesis is the biospheres


metabolic foundation
In photosynthesis, the energy that enters the
chloroplasts as sunlight becomes stored as
chemical energy in
organic
compounds.

Copyright 2002 Pearson Education, Inc., publishing as Benjamin Cummings

Copyright 2002 Pearson Education, Inc., publishing as Benjamin Cummings