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Transcript
Photosynthesis
Chapter.7
Introduction
•  Plants use water and atmospheric carbon dioxide to
produce a simple sugar and liberate oxygen
–  Earth s plants produce 160 billion metric tons of sugar each year
through photosynthesis, a process that converts solar energy to
chemical energy
–  Sugar is food for humans and for animals that we consume
•  Scientists have suggested that plants can be used in energy
plantations to create a fuel source to replace fossil fuels
–  This would be an excellent energy solution, because air pollution, acid
precipitation, and greenhouse gases could be significantly reduced
Overview of photosynthesis
6 CO2
+6
H 2O
Carbon dioxide Water
Light
energy
C6H12O6
Photosynthesis
+ 6
O2
Glucose Oxygen gas
Autotrophs
•  Autotrophs are living things that are able to make their own
food without using organic molecules derived from any other
living thing
–  Autotrophs that use the energy of light to produce organic molecules
are called photoautotrophs
–  Most plants, algae and other protists, and some prokaryotes are photoautotrophs
–  Producers in a food chain
–  cf) Heterotrophs : the consumers
•  The ability to photosynthesize is directly related to the
structure of chloroplasts
–  Chloroplasts are organelles consisting of photosynthetic pigments,
enzymes, and other molecules grouped together in membranes
–  Chloroplasts are structurally similar to and likely evolved from photosynthetic
bacteria
(b) Multicellular
alga
(a) Plants
(d) Cyanobacteria
(c) Unicellular
protists
40 µm
10 µm
(e) Purple sulfur
1 µm
bacteria
5
Photosynthesis occurs in chloroplasts
• 
Chloroplasts are the major sites of photosynthesis in green plants
–  Chlorophyll, an important light absorbing pigment in chloroplasts, is
responsible for the green color of plants
–  Chlorophyll plays a central role in converting solar energy to chemical
energy.
• 
Chloroplasts are concentrated in the cells of the mesophyll, the green tissue in
the interior of the leaf
• 
Stomata are tiny pores in the leaf that allow carbon dioxide to enter and oxygen
to exit
• 
Veins in the leaf deliver water absorbed by roots
• 
An envelope of two membranes encloses the stroma, the dense fluid within the
chloroplast
• 
A system of interconnected membranous sacs called thylakoids segregates the
stroma from another compartment, the thylakoid space
–  Thylakoids are concentrated in stacks called grana
Anatomical structure of leaf
Mesophyll: Middle leaf in Greek
The assimilation tissue is the primary location of photosynthesis in Plant
Stomata (stoma) : gas exchange
7
Leaf cross section
Chloroplasts Vein
Mesophyll
Stomata
Chloroplast
Thylakoid
Stroma Granum Thylakoid
space
1 µm
CO2 O2
Mesophyll
cell
Outer
membrane
Intermembrane
space
Inner
membrane
20 µm
9
1.  Outer membrane
2.  Intermembrane space
3.  Inner membrane
4.  Stroma (aqueous fluid)
7  Granum (stack of thylakoids)
8  Thylakoid
9  Starch
10 Ribosome
11  Plastidial DNA
10
Plants produce O2 gas
•  Scientists have known for a long time that plants produce O2, but early on
they assumed it was extracted from CO2 taken into the plant
–  Using a heavy isotope of oxygen, 18O, they showed with tracer experiments
that O2 actually comes from H2O
Experiment 1
6 CO2 + 12 H2O
C6H12O6 + 6 H2O + 6 O2
Not
labeled
Experiment 2
6 CO2 + 12 H2O
C6H12O6 + 6 H2O + 6 O2
Labeled
Copyright © 2009 Pearson Education, Inc.
Reactants:
Products:
6 CO2
C6H12O6
12 H2O
6 H 2O
6 O2
Photosynthesis is a redox process
•  Photosynthesis, like respiration, is a redox (oxidationreduction) process
–  Water molecules are split apart by oxidation, which means that they
lose electrons along with hydrogen ions (H+)
–  Then CO2 is reduced to sugar as electrons and hydrogen ions are
added to it
•  Recall that cellular respiration uses redox reactions to
harvest the chemical energy stored in a glucose molecule
–  This is accomplished by oxidizing the sugar and reducing O2 to H2O
–  The electrons lose potential as they travel down an energy hill, the
electron transport system
–  In contrast, the food-producing redox reactions of photosynthesis
reverse the flow and involve an uphill climb
Photosynthesis (Use light energy)
Reduction
6 CO2 + 6 H2O
C6H12O6 + 6 O2
Oxidation
Cellular respiration (release chemical energy)
Oxidation
C6H12O6 + 6 O2
6 CO2 + 6 H2O
Reduction
Photosynthesis Cont d
•  In photosynthesis, electrons gain energy by being
boosted up an energy hill
–  Light energy captured by chlorophyll molecules provides
the boost for the electrons
–  As a result, light energy is converted to chemical energy,
which is stored in the chemical bonds of sugar molecules
16
Light reactions
•  Occur in thylakoids
•  light energy is converted in the thylakoid membranes to chemical energy
and O2
•  Water is split to provide the O2 as well as electrons and proton (H+)
–  O2 generation as a byproduct
–  Transfer of H+ and electron to NADP+ reducing it to NADPH
–  NADPH : electron carrier, provides reducing power to the Calvin cycle
•  ATP generation by photo-phosphorylation
Calvin cycle
•  occurs in the stroma of the chloroplast
–  It is a cyclic series of reactions that builds sugar molecules from CO2
and the products of the light reactions
–  CO2 is incorporated into organic compounds, a process called carbon
fixation
•  NADPH produced by the light reactions provides the
electrons for reducing carbon in the Calvin cycle
–  ATP from the light reactions provides chemical energy for the
Calvin cycle
–  The Calvin cycle is often called the dark (or light-independent)
reactions
•  No light is required during reaction
•  Occur during daytime due to activation of light reaction during daytime
supplying it with NADPH and ATP
CO2
H 2O
Chloroplast
Light
NADP+
ADP
! P
LIGHT
REACTIONS
CALVIN
CYCLE
(in stroma)
(in thylakoids)
ATP
NADPH
O2
Sugar
Visible radiation
•  Sunlight contains energy called electromagnetic energy or
radiation
–  Visible light is only a small part of the electromagnetic spectrum, the
full range of electromagnetic wavelengths
–  Electromagnetic energy travels in waves, and the wavelength is the
distance between the crests of two adjacent waves
•  Light behaves as discrete packets of energy called photons
–  A photon is a fixed quantity of light energy, and the shorter the
wavelength, the greater the energy
Electromagnetic spectrum
Increasing energy
10–5 nm 10–3 nm
Gamma
rays
X-rays
1 nm
103 nm
UV
1m
106 nm
Infrared
Microwaves
103 m
Radio
waves
Visible light
380 400
600
500
Wavelength (nm)
700
650
nm
750
Visible radiation for light reaction
•  Pigments, molecules that absorb light, are built into
the thylakoid membrane
–  Plant pigments absorb some wavelengths of light and
transmit others
–  We see the color of the wavelengths that are transmitted;
for example, chlorophyll transmits green
–  Wavelengths that are not absorbed are reflected or transmitted
–  Leaves appear green because chlorophyll reflects and transmits green
light
Light
Reflected
light
Chloroplast
Thylakoid
Absorbed
light
Transmitted
light
Visible radiation Cont d
• 
Chloroplasts contain several different pigments and all absorb light of different
wavelengths
–  The main photosynthetic pigment : Chlorophyll a absorbs blue violet and red
light and reflects green
–  The accessory pigments : Chlorophyll b absorbs blue and orange and
reflects yellow-green
–  The accessory pigments : The carotenoids absorb mainly blue-green light
and reflect yellow and orange
•  Photoprotection: absorbs and dissipate excessive light energy that may
damage chlorophyll or produce ROS
•  Phytochemicals from the plant : Antioxidants
25
Photosystems capture solar power
•  Pigments in chloroplasts are responsible for absorbing
photons (capturing solar power), causing release of electrons
–  The electrons jump to a higher energy level—the excited state—
where electrons are unstable
–  The electrons drop back down to their ground state, and, as they
do, release their excess energy
•  The energy released could be lost as heat or light, but rather
it is conserved as it is passed from one molecule to another
molecule
–  All of the components to accomplish this are organized in thylakoid
membranes in clusters called photosystems
–  Photosystems are light-harvesting complexes surrounding a reaction
center complex
–  Protein with pigment molecules (chlorophyll a, b and carotenoids)
Excitation of isolated chlorophyll by light.
Photosystems capture solar power
•  The energy is passed from molecule to molecule within the
photosystem
–  Finally it reaches the reaction center where a primary electron
acceptor accepts these electrons and consequently becomes reduced
–  This solar-powered transfer of an electron from the reaction center
pigment to the primary electron acceptor is the first step of the light
reactions
The structure and function of a photosystem
29
Photosystems I and II
•  Two types of photosystems (photosystem I and photosystem II)
–  Photosystem II, which functions first,
–  The reaction-center chlorophyll a of PS II is called P680 because its pigment
absorbs light with a wavelength of 680 nm
–  Photosystem I, which functions next,
–  The reaction-center chlorophyll a of PS I is called P700 because it absorbs
light with a wavelength of 700 nm
•  During the light reactions, light energy is transformed into the chemical
energy of ATP and NADPH
–  To accomplish this, electrons removed from water pass from photosystem II to
photosystem I and are accepted by NADP+
–  The bridge between photosystems II and I is an electron transport chain that
provides energy for the synthesis of ATP
•  NADPH, ATP, and O2 are the products of the light reactions
e–
ATP
e–
e–
e–
NADPH
e–
Mill
makes
ATP
n
Photo
e–
Photon
e–
Photosystem II
Photosystem I
Electron transport chain
Provides energy for
synthesis of
by chemiosmosis
Photon
ATP
Photosystem II
Stroma
NADP+ + H+
Photon
Photosystem I
1
Primary
acceptor
Primary
acceptor
2
e–
e–
Thylakoid
membrane
4
P700
P680
Thylakoid
space
3
H 2O
1
2
5
O2 + 2 H+
6
NADPH
Chemiosmosis for ATP synthesis
•  Interestingly, chemiosmosis is the mechanism that not only is
involved in oxidative phosphorylation in mitochondria but also
generates ATP in chloroplasts
–  ATP is generated because the electron transport chain
produces a concentration gradient of hydrogen ions across
a membrane
•  ATP synthase couples the flow of H+ to the phosphorylation
of ADP
–  The chemiosmosis production of ATP in photosynthesis is
called photo-phosphorylation
Chemiosmosis for ATP synthesis
Chloroplast
Stroma (low H+
concentration)
Light
H+
Light
H+
NADP+ + H+
ADP + P
NADPH
H+
ATP
H+
Thylakoid
membrane
H 2O
1
2
O2 + 2 H+
H+
Photosystem II Electron
transport
chain
Thylakoid space
(high H+ concentration)
H+
H+
H+
H+
H+
Photosystem I
H+
H+
H+
H+
ATP synthase
Chloroplast
Mitochondrion
CHLOROPLAST
STRUCTURE
MITOCHONDRION
STRUCTURE
H+
Intermembrane
space
Inner
membrane
Matrix
Diffusion
Electron
transport
chain
Thylakoid
membrane
ATP
synthase
Stroma
ADP + P i
Key
[H+ ]
Higher
Lower [H+ ]
Thylakoid
space
H+
ATP
35
STROMA
(low H+ concentration)
Photosystem II
Light
4 H+
Cytochrome
complex
Photosystem I
Light
NADP+
reductase
3
Fd
Pq
H 2O
NADPH
Pc
2
1
THYLAKOID SPACE
(high H+ concentration)
1/
2
NADP+ + H+
O2
+2 H+
4 H+
To
Calvin
Cycle
Thylakoid
membrane
STROMA
(low H+ concentration)
ATP
synthase
ADP
+
Pi
ATP
H+
36
Calvin cycle
•  The Calvin cycle makes sugar within a chloroplast
–  To produce sugar, the necessary ingredients are atmospheric CO2,
ATP, and NADPH, which were generated in the light reactions
–  Using these three ingredients, an energy-rich, three-carbon sugar
called glyceraldehyde-3-phosphate (G3P) is produced
•  A plant cell may then use G3P to make glucose and other organic
molecules
•  The starting material for the Calvin cycle is a five-carbon
sugar named ribulose bisphosphate (RuBP)
–  The next step is a carbon (CO2) fixation step aided by an enzyme
called rubisco (ribulose bisphosphate carboxylase/oxygenase)
–  This is repeated over and over, one carbon at a time
–  For One G3P, nine ATP and six NADPH are consumed
Overview of Calvin cycle
Input
CO2
ATP
NADPH
CALVIN
CYCLE
Output:
G3P
39
40
41
Photosynthesis uses light energy, CO2, and H2O
•  The chloroplast, which integrates the two stages of
photosynthesis, makes sugar from CO2
–  All but a few microscopic organisms depend on the food-making
machinery of photosynthesis
–  Plants make more food than they actually need and stockpile it as
starch in roots, tubers, and fruits
•  Some plants have evolved a means of carbon fixation that
saves water during photosynthesis
–  One group can shut its stomata when the weather is hot and dry to
conserve water but is able to make sugar by photosynthesis
–  These are called the C4 plants because they first fix carbon dioxide
into a four-carbon compound
H 2O
CO2
Chloroplast
Light
NADP+
ADP
+ P
Photosystem II
Thylakoid
membranes
RuBP
CALVIN
CYCLE 3-PGA
(in stroma)
Electron
transport
chains
Photosystem I
ATP
NADPH
Stroma
G3P
O2
Sugars
LIGHT REACTIONS
CALVIN CYCLE
Cellular
respiration
Cellulose
Starch
Other organic
compounds
Photorespiration
•  In hot climates, most plants (C3 plants) stomata close to
reduce water loss so oxygen builds up
–  Rubisco adds oxygen instead of carbon dioxide to RuBP and produces
a two-carbon compound, a process called photorespiration
–  Unlike photosynthesis, photorespiration produces no sugar, and unlike
respiration, it produces no ATP
•  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
Photorespiration
45
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
(phosphoenolpyruvate) 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
46
C4 pathway
The C4 pathway
C4 leaf anatomy
Mesophyll
cell
PEP carboxylase
Mesophyll cell
Photosynthetic
cells of C4
Bundleplant leaf
sheath
cell
Oxaloacetate (4C)
Vein
(vascular tissue)
PEP (3C)
ADP
Malate (4C)
Stoma
Bundlesheath
cell
CO2
ATP
Pyruvate (3C)
CO2
Calvin
Cycle
Sugar
Vascular
tissue
47
48
CAM (crassulacean acid metabolism)
•  Another adaptation to hot and dry environments has
evolved in the CAM (crassulacean acid
metabolism), such as pineapples and cacti
–  CAM plants conserve water by opening their stomata and
admitting CO2 only at night
–  Stomata close during the day, and CO2 is released from organic acids
and used in the Calvin cycle
–  When CO2 enters, it is fixed into a four-carbon compound,
like in C4 plants, and in this way CO2 is banked
–  It is released into the Calvin cycle during the day
49
Sugarcane
Pineapple
C4
Mesophyll Organic acid
cell
CAM
CO2
1 CO2 incorporated
(carbon fixation) Organic acid
Calvin
Cycle
Night
CO2
CO2
Bundlesheath
cell
CO2
2 CO2 released
to the Calvin
cycle
Sugar
(a) Spatial separation of steps
Calvin
Cycle
Day
Sugar
(b) Temporal separation of steps
51
Global warming
• 
The greenhouse effect results from solar energy warming our planet
–  Gases in the atmosphere (often called greenhouse gases), including CO2,
reflect heat back to Earth, keeping the planet warm and supporting life
–  However, as we increase the level of greenhouse gases, Earth s temperature
rises above normal, initiating problems
• 
Increasing concentrations of greenhouse gases lead to global warming, a slow
but steady rise in Earth s surface temperature
–  The extraordinary rise in CO2 is mostly due to the combustion of carbon-based
fossil fuels
–  The consequences of continued rise will be melting of polar ice, changing
weather patterns, and spread of tropical disease
• 
Perhaps photosynthesis can mitigate the increase in atmospheric CO2
–  However, there is increasing widespread deforestation, which aggravates the
global warming problem
Some heat
energy escapes
into space
Sunlight
Atmosphere
Radiant heat
trapped by CO2
and other gases
Southern tip of
South America
Antarctica
Notice for Mid-term exam
• 
• 
• 
• 
Type I (50%) : From Item pool (5점짜리 10문제)
–  5 items (with right answer) from each student
•  Due on 16th April (11:59 PM)
–  Upload to cyber campus
–  10 % of mid-term grade
–  File name should be “ID number-name.pdf” (no other format)
»  Pdf format only!!!!!!
•  Creative and comprehensive items will be accepted and presented.
–  With extra point.
–  Short answered question (Free format but no multiple choice) – 문제당 5점..
•  Wrong question and answer will not be accepted
–  Item pool will be found in cyber-campus on no later than 18th of April
Type II (30%) : Simple short answered question (5점짜리 6문제)
–  For minimum grade
Type III (20%) : Creative and comprehensive write-out answered question (10점짜리 2문제)
–  For assessment of the students
–  Prepare the answers for Why or How in the classes
–  May take 10 – 20 min each question
–  May be important for ‘A grade
–  Consists of multiple sub-questions (ex. III-1: 5점, III-2, 5점)
Mid-term Exam on 22th (Fri) April
–  Where: K302
55
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