Campbell: Biology
Twelfth Edition
Chapter 10
Photosynthesis
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Figure 10.1 Each leaf of this tree is harvesting energy from sunlight
and using it to convert C O2 and H2O into chemical energy stored in
sugar and other organic molecules. The tree uses these sugars for
energy and to build its own trunk, branches, and leaves.
Remarkably, enough sugars are left over to feed other organisms
(like moth larvae, shown below) that cannot carry out this
extraordinary transformation
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Figure 10.1 How do photosynthetic cells
use light to change carbon dioxide and
water into organic molecules and oxygen?
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Chemical reactions either release or store
energy
Endergonic reactions use energy.
These reactions need energy input energy to
create bonds and form more complex molecules
ANABOLIC REACTIONS: FORM or BUILD
complex molecules from simpler ones
Photosynthesis is one big anabolic pathway
involves many steps,
Uses energy from the sun to convert carbon dioxide
and water into sugar (organic molecules/food)
The Process that Feeds the Biosphere
Plants and other photosynthetic organisms contain
organelles called chloroplasts
Photosynthesis is the process that converts solar
energy into chemical energy within chloroplasts
Photosynthesis occurs in plants, algae, certain other
unicellular eukaryotes, and some prokaryotes
Directly or indirectly, photosynthesis nourishes
almost the entire living world
The Process That Feeds the Biosphere,
Continued
Autotrophs
are “self-feeders” that is they make their own food
Do not usually consume organic molecules from other
organisms.
are the producers of the biosphere, producing
organic molecules from
Two types:
Photoautptophs: use energy of light to produce food
Chemoautotrophs: use inorganic chemicals as their
energy source
Figure 10.2 Photoautotrophs
The Process That Feeds the Biosphere,
Continued-2
Heterotrophs
obtain organic material from other organisms
consumers of the biosphere
Some eat other living organisms; others, called
decomposers, consume dead organic material or
feces
Almost all heterotrophs, including humans, depend
on photoautotrophs for food and
Concept 10.1: Photosynthesis converts light
energy to the chemical energy of food
Photosynthesis in plants
• takes place in chloroplasts,
• mostly within leaves,
• converts carbon dioxide and water into
organic molecules, and
• releases oxygen.
BioFlix® Animation: Photosynthesis
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Chloroplasts: The Sites of Photosynthesis
in Plants
Leaves are the major locations of photosynthesis in
plants
Their green color is from chlorophyll, the green
pigment that absorbs light
Chloroplasts are found mainly in cells of the
mesophyll, the interior tissue of the leaf
Each mesophyll cell contains 30-40 chloroplasts
Gases enter and exit the leaf through pores called
stomata
Chloroplasts: The Sites of Photosynthesis
in Plants, Continued
A chloroplast has an envelope of two membranes
surrounding a dense fluid called the stroma
Thylakoids are connected sacs in the chloroplast
that compose a third membrane system
Thylakoids may be stacked in columns called grana
Chlorophyll, the pigment that gives leaves their
green color, resides in the thylakoid membranes
Figure 10.3 Zooming in on the
location of photosynthesis in a plant
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Video: Chloroplasts in Motion
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Tracking Atoms Through
Photosynthesis: Scientific Inquiry
• Photosynthesis is a complex series of reactions that can
be summarized as the following equation:
6 CO 2 + 12 H 2 O + Light energy C6 H12O 6 +6 O 2 +6 H 2O
• The overall chemical change during photosynthesis is the
reverse of cellular respiration
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Figure 10.4 Tracking atoms through
photosynthesis
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Figure 10.UN01 Photosynthesis as a
redox process
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Photosynthesis as a Redox Process
Photosynthesis reverses the direction of electron
flow compared to respiration
Photosynthesis is a redox process in which is
oxidized and is reduced
Photosynthesis is an endergonic process; the
energy boost is provided by light
Figure 7.UN01
Becomes reduced
Light
energy
6 CO2 6 H2O C6H12O6 6 O2
Carbon dioxide Water Oxygen gas
Photosynthesis Glucose
Becomes oxidized
The Splitting of Water
Chloroplasts split into hydrogen and oxygen,
incorporating the electrons of hydrogen into sugar
molecules and releasing oxygen as a by-product
Photosynthesis: A Preview
Two stages: the Light reactions (the photo part) and
Calvin cycle (the synthesis part)
[Link] reactions: occurs in the thylakoid
membranes in these reactions
• H2O is split, providing electrons and giving off O2
as a by-product,
• ATP is generated from ADP and a phosphate
group by photophosphorylation and
• light energy is absorbed by the chlorophyll
molecules to drive the transfer of electrons and
H+ from water to the electron acceptor NADP+
reducing it to NADPH.
• ATP and NADPH produced by the light reactions
provides the electrons and power to the Calvin
[Link] cycle: occurs in the stroma of the
chloroplast.
• The Calvin cycle is a cyclic series of reactions
that assembles sugar molecules using CO2 and
the energy-rich products of the light reactions.
• CO2 is incorporated into organic compounds in
a process called carbon fixation.
• After carbon fixation, enzymes of the cycle
make sugars by further reducing the carbon
compounds.
• The Calvin cycle is often called the dark
reactions or light-independent reactions,
because none of the steps requires light
directly.
Figure 10.5 An overview of
photosynthesis: cooperation of the
light reactions and the Calvin cycle
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The Nature of Sunlight
Light is electromagnetic energy, also called
electromagnetic radiation
Electromagnetic energy travels in rhythmic waves
Wavelength is the distance between crests
of electromagnetic waves
Wavelength determines the type of electromagnetic
energy
The Nature of Sunlight, Continued
The electromagnetic spectrum is the entire range
of electromagnetic energy, or radiation
Visible light consists of wavelengths (380 nm to
750 nm) that produce colors we can see
Visible light also includes the wavelengths that drive
photosynthesis
Light also behaves as though it consists of discrete
particles, called photons
Figure 10.6 The electromagnetic spectrum
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Photosynthetic Pigments: The Light Receptors
Pigments 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
Figure 10.7 Why leaves are green:
interaction of light with chloroplasts
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Animation: Light Energy and Pigments
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Photosystems capture solar energy
Pigments in chloroplasts absorb
Figure 10.11 Excitation of Isolated
photons (capturing solar power), which Chlorophyll by Light
increases the potential energy of
the pigment’s electrons and
sends the electrons into an
unstable state.
These unstable electrons
drop back down to their
“ground state,” and as they
do,
release their excess
energy as heat.
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, i.e. they
function like lightning rods.
• The light 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
in the transformation of light energy to
chemical energy in the light reactions.
Figure 10.12a The Structure and Function of a
Photosystem (Part 1: Mechanism)
a) How a photosystem harvests light
Figure 7.7B
Excited state
Photosystem
Light
Light-harvesting Reaction-center
complexes complex
Primary electron
acceptor
Thylakoid membrane
Pigment
Transfer Pair of molecules
of energy chlorophyll a molecules
• Two types of photosystems (photosystem I and
photosystem II) cooperate in the light reactions.
• Each type of photosystem has a characteristic
reaction center.
– Photosystem II or PSII, which functions first, is
called P680 because its pigment absorbs light with a
wavelength of 680 nm.
– Photosystem I or PSI, which functions second, is
called P700 because it absorbs light with a
wavelength of 700 nm.
Two photosystems connected by an electron
transport chain generate ATP and NADPH
In the light reactions, light energy is transformed into the
chemical energy of ATP and NADPH.
To accomplish this, electrons are
removed from water,
passed from photosystem II to photosystem I, and
accepted by NADP+, reducing it to NADPH.
Between the two photosystems, the electrons
move down an electron transport chain and
provide energy for the synthesis of ATP.
Figure 7.8A
Electron transport chain NADP H NADPH
Light
Light Provides energy for
synthesis of ATP Photosystem I 6
Photosystem II
Stroma by chemiosmosis
1
Primary
Primary
acceptor
Thylakoid membrane
acceptor
2
4
5
P680 P700
Thylakoid
3
space
H2O 1
2 O2 2 H
1. What is the source of electrons for PSII? PS?
2. Who is the final electron acceptor in the ETC
of the thylakoid membrane?
Figure 10.14 A mechanical analogy for linear
electron flow during the light reactions
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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
Linear Electron Flow, Continued
There are eight steps in linear electron flow:
1. A photon hits a pigment in a light-harvesting
complex of PS II, and its energy is passed among
pigment molecules until it excites P680
2. An excited electron from P680 is transferred to the
primary electron acceptor (we now call it )
Linear Electron Flow, Continued-1
3. is split by enzymes, and the electrons are
transferred from the hydrogen atoms to , thus
reducing it to P680
is the strongest known biological oxidizing agent
Th are released into the thylakoid space
e is released as a by-product of this reaction
Linear Electron Flow, Continued-2
4. 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
5. Potential energy stored in the proton gradient drives
production of ATP by chemiosmosis
Linear Electron Flow, Continued-3
6. In PS I (like PS II), transferred light energy excites
P700, which loses an electron to the primary electron
acceptor
(P700 that is missing an electron) accepts an
electron passed down from PS II via the electron
transport chain
Linear Electron Flow, Continued-4
7. Each electron “falls” down an electron transport
chain from the primary electron acceptor of PS I to
the protein ferredoxin (Fd)
8. reductase catalyzes the transfer of electrons
to , reducing it to NADPH
The electrons of NADPH are available for the reactions
of the Calvin cycle
This process also removes an from the stroma
Figure 10.13 How linear electron flow during the
light reactions generates AT P and NADP H
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Figure 10.Un02 Schematic of light
reactions
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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
A Comparison of Chemiosmosis in
Chloroplasts and Mitochondria, Continued
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
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 using oxidative phosphorylation
Chloroplasts transform light energy into the chemical
energy of ATP using photophosphorylation
Spatial organization of chemiosmosis differs
between chloroplasts and mitochondria but also
shows similarities
Chemiosmosis powers ATP synthesis in
the light reactions
In photophosphorylation, using the initial energy
input from light,
the electron transport chain pumps H+ from the
stroma (FLUID) into the thylakoid SPACE and
the resulting concentration gradient drives H + back
through ATP synthase, producing ATP.
Figure 10.16 Comparison of chemiosmosis in
mitochondria and chloroplasts
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Figure 7.UN03
Mitochondrion Chloroplast
Intermembrane H c.
space
Membrane
Matrix d.
a.
b. e.
A Comparison of Chemiosmosis in
Chloroplasts and Mitochondria, Continued-1
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 to NADPH
Figure 10.17 The light reactions:
organization of the thylakoid membrane
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Chloroplast
To Calvin
Cycle H+
Light ADP P ATP
Light
Stroma
(low H+ NADPH
H+ NADP+ H+
concentration)
H+ H+
Thylakoid
membrane
H+ H+ H+
H+
H2O H +
1 H+
O + 2 H+ H+ H+
Thylakoid space 2 2 H+ H+ H+
(high H+ Electron H+
H+
concentration) Photosystem II transport chain Photosystem I ATP synthase
Animation: The Light Reactions
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BioFlix® Animation: The Light Reactions
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Figure 10.Un04 Calvin cycle schematic
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Animation: The Calvin Cycle
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BioFlix® Animation: The Calvin Cycle
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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 Calvin cycle is anabolic; it builds sugar from
smaller molecules by using ATP and the reducing
power of electrons carried by NADPH
The Calvin Cycle uses the Chemical Energy of ATP
and NADPH to Reduce CO2 to Sugar, Continued
Carbon enters the cycle as and leaves as a
sugar named glyceraldehyde 3-phospate (G3P)
For net synthesis of one G3P, the cycle must take
place three times, fixing three molecules of
The Calvin cycle has three phases:
1. Carbon fixation (catalyzed by rubisco)
2. Reduction
3. Regeneration of the acceptor (RuBP)
Figure 10.18 The Calvin cycle
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Figure 10.UN03
3 CO2
Carbon fixation
RuBP 3-PGA
Calvin
Cycle
Regeneration of
RuBP
Reduction
1 G3P (3C)
Input for 3 CO2 6 CO2
Making Input for
1 G3P
ATP Making ATP
NADPH 1 glucose NADPH
Calvin Calvin
Cycle Cycle
Output: G3P
C6H12O6
(a 3C molecule)
(a 6C molecule)
The energy entering chloroplasts as sunlight gets stored as chemical energy in organic compounds
Concept 10.5: Life depends on
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 chloroplasts and other structures such as roots, tubers, seeds, and fruits
photosynthesis
The Importance of Photosynthesis: A
Review
Figure 10.21 A review of photosynthesis
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Figure 10.22b A Review of Photosynthesis (Part 2)
Figure 10.UN05 Summary of Key Concepts:
Linear Electron Flow
Figure 10.22c A Review of Photosynthesis
(Part 3)
LIGHT REACTIONS CALVIN CYCLE REACTIONS
• Are carried out by molecules • Take place in the stroma
in the thylakoid membranes • Use ATP and NADPH to convert
• Convert light energy to the chemical to the sugar G3P
energy of ATP and NADPH • Return ADP, inorganic phosphate,
• Split and release and to the light reactions
to the atmosphere
Figure 10.22 Make Connections: The
working cell
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Figure 10.22a Make Connections: The Working
Cell (Part 1)
Figure 10.22b Make Connections: The Working
Cell (Part 2)
Figure 10.22c Make Connections: The Working
Cell (Part 3)
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 but also limits photosynthesis
The closing of stomata reduces access to and
causes to build up
These conditions favor an apparently wasteful
process called photorespiration
Photorespiration: An Evolutionary Relic?
In most plants ( plants), initial fixation of , via
rubisco, forms a three-carbon compound
(3-phosphoglycerate)
In photorespiration, rubisco adds instead of
in the Calvin cycle, producing a two-carbon
compound
Photorespiration consumes and organic fuel and
releases without producing ATP or sugar
Photorespiration: An Evolutionary Relic?,
Continued
Photorespiration may be an evolutionary relic
because rubisco first evolved at a time when the
atmosphere had far less and more
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
C4 Plants
plants minimize the cost of photorespiration by
incorporating into four-carbon compounds
There are two distinct types of cells in the leaves of
plants:
Bundle-sheath cells are arranged in tightly packed
sheaths around the veins of the leaf
Mesophyll cells are loosely packed between the
bundle sheath and the leaf surface
C4 Plants, Continued
Sugar production in plants occurs in a three-step
process:
1. The production of the four-carbon precursors is
catalyzed by the enzyme PEP carboxylase in the
mesophyll cells
PEP carboxylase has a higher affinity for than
rubisco does; it can fix even when
concentrations are low
2. These four-carbon compounds are exported to
bundle-sheath cells
3. Within the bundle-sheath cells, they release
Figure 10.19 C4 leaf anatomy and the C4 pathway
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C4 Plants, Continued-2
Since the Industrial Revolution in the 1800s,
levels have risen greatly
Increasing levels of may affect and plants
differently, perhaps changing the relative abundance
of these species
The effects of such changes are unpredictable and a
cause for concern
C4 Plants, Continued-3
Suitable agricultural land is decreasing due to the
effects of climate change, while the world demand
for food continues to increase
photosynthesis uses less water and resources
than photosynthesis
Scientists have genetically modified rice, a plant,
to carry out photosynthesis
They estimate 30-50% increase in yield compared to
rice
CAM Plants
Some plants, including succulents, use
crassulacean acid metabolism (CAM) to
fix carbon
CAM plants open their stomata at night,
incorporating into organic acids that are stored
in the vacuoles
Stomata close during the day, and is released
from organic acids and used in the Calvin cycle
CAM Plants, Continued
The CAM pathway is similar to the pathway in
that they both incorporate into organic
intermediates before it enters the Calvin cycle
The pathway structurally separates the initial
steps of carbon fixation from the Calvin cycle
In the CAM pathway, these steps occur in the same
cell, but are separated in time
Figure 10.20 C4 and CA M
photosynthesis compared
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Animation: Photosynthesis in Dry
Climates
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