Overview: The Process That Feeds the

Biosphere
• Photosynthesis is the process that converts
solar energy into chemical energy
• Directly or indirectly, photosynthesis nourishes
almost the entire living world
Overview: The Process That Feeds the
Biosphere
• Autotrophs sustain themselves without eating
anything derived from other organisms
• Autotrophs are the producers of the biosphere,
producing organic molecules from CO2 and other
inorganic molecules
• Almost all plants are photoautotrophs, using the
energy of sunlight to make organic molecules
Overview: The Process That Feeds the
Biosphere
• Photosynthesis occurs in plants, algae, certain
other protists, and some prokaryotes
• These organisms feed not only themselves but
also most of the living world
 (b) Multicellular
(a) Plants alga

(d) Cyanobacteria
40 m
(c) Unicellular
10 m
protists

(e) Purple sulfur

1 m
bacteria
Overview: The Process That Feeds the
Biosphere

• Heterotrophs obtain their organic material from

other organisms
• Heterotrophs are the consumers of the
biosphere
• Almost all heterotrophs, including humans,
depend on photoautotrophs for food and O 2
Overview: The Process That Feeds the
Biosphere
• The Earth’s supply of fossil fuels was formed
from the remains of organisms that died
hundreds of millions of years ago
• In a sense, fossil fuels represent stores of solar
energy from the distant past
Photosynthesis converts light energy
to the chemical energy of food
• Chloroplasts are structurally similar to and likely
evolved from photosynthetic bacteria
• The structural organization of these cells allows
for the chemical reactions of photosynthesis
 Chloroplasts: The Sites of
Photosynthesis in Plants
• Leaves are the major locations of
photosynthesis
• Their green color is from chlorophyll, the
green pigment within chloroplasts
• Chloroplasts are found mainly in cells of the
mesophyll, the interior tissue of the leaf
• Each mesophyll cell contains 30–40
chloroplasts
 Chloroplasts: The Sites of
Photosynthesis in Plants
• CO2 enters and O2 exits the leaf through
microscopic pores called stomata
• 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
interior fluid
 Leaf cross section
Chloroplasts Vein

Mesophyll

Stomata
CO2 O2

Chloroplast Mesophyll
cell

Outer
membrane
Thylakoid Intermembrane
Stroma Granum Thylakoid space 20 m
space Inner
membrane

1 m
Tracking Atoms Through Photosynthesis:
Scientific Inquiry
• Photosynthesis is a complex series of reactions
that can be summarized as the following
equation:
6 CO2 + 12 H2O + Light energy  C6H12O6 + 6 O2 + 6 H2O
 The Splitting of Water
• Chloroplasts split H2O into hydrogen and
oxygen, incorporating the electrons of hydrogen
into sugar molecules and releasing oxygen as a
by-product
Reactants: 6 CO2 12 H2O

Products: C6H12O6 6 H2O 6 O2

 Photosynthesis as a Redox Process
• Photosynthesis reverses the direction of electron
flow compared to respiration
• Photosynthesis is a redox process in which H 2O
is oxidized and CO2 is reduced
• Photosynthesis is an endergonic process; the
energy boost is provided by light
Photosynthesis as a Redox Process

becomes reduced

Energy  6 CO2  6 H2O C6 H12 O6  6 O2

becomes oxidized
 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
 The Two Stages of Photosynthesis: A
Preview
• 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
 H2O CO2

Light

NADP
ADP
+Pi
Calvin
Light Cycle
Reactions

ATP

NADPH

Chloroplast

O2 [CH2O]
(sugar)
The light reactions convert solar energy to
the chemical energy of ATP and NADPH

• Chloroplasts are solar-powered chemical

factories
• Their thylakoids transform light energy into the
chemical energy of ATP and NADPH
 The Nature of Sunlight
• 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
 1m
105 nm 103 nm 1 nm 10 nm
3
106 nm (109 nm) 103 m

Gamma Micro- Radio

X-rays UV Infrared waves waves
rays

Visible light

380 450 500 550 600 650 700 750 nm

Shorter wavelength Longer wavelength
Higher energy Lower energy
 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
 Light
Reflected
light

Chloroplast

Absorbed Granum
light

Transmitted
light
 Photosynthetic Pigments: The Light
Receptors

• Chlorophyll a is the main photosynthetic pigment

• Accessory pigments, such as chlorophyll b,
broaden the spectrum used for photosynthesis
• Accessory pigments called carotenoids absorb
excessive light that would damage chlorophyll
 Excitation of Chlorophyll by Light
• 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
• If illuminated, an isolated solution of chlorophyll
will fluoresce, giving off light and heat
 Excited
e state
Energy of electron

Heat

Photon
(fluorescence)
Photon
Ground
Chlorophyll state
molecule

(a) Excitation of isolated chlorophyll molecule (b) Fluorescence

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
 Photosystem STROMA
Photon
Light- Reaction- Primary
harvesting center electron
complexes complex acceptor
Thylakoid membrane

e

Transfer Special pair of Pigment

of energy chlorophyll a molecules
molecules
THYLAKOID SPACE
(INTERIOR OF THYLAKOID)
(a) How a photosystem harvests light
• A primary electron acceptor in the reaction
center accepts excited electrons 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
• 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
• 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

• 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+)
 Linear Electron Flow

Primary
acceptor
2
e

P680

1 Light

Pigment
molecules

Photosystem II
(PS II)
 Linear Electron Flow
• 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
 Linear Electron Flow

Primary
acceptor
2
e
2 H H2O
+
3
1
/ 2 O2
e
e
P680

1 Light

Pigment
molecules

Photosystem II
(PS II)
 Linear Electron Flow

• 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

E le
Primary ctro 4
acceptor n tr
ans
por
Pq t ch
2 ain
e
2H  H2O Cytochrome
+ complex
3
1
/ 2 O2
e Pc
e
P680 5

1 Light

ATP

Pigment
molecules

Photosystem II
(PS II)
 Linear Electron Flow

• 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
Primary
E le acceptor
Primary ctro 4
acceptor n tr
ans
por e
Pq t ch
2 ain
e
2H  H2O Cytochrome
+ complex
3
1
/ 2 O2
e Pc
e P700
P680 5 Light

1 Light 6

ATP

Pigment
molecules
Photosystem I
(PS I)
Photosystem II
(PS II)
 Linear Electron Flow

• 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
• This process also removes an H+ from the
stroma
 Linear Electron Flow
E
tra lect
c h ns r on
a i po
Primary n rt
E le acceptor
Primary ctro 4 7
acceptor n tr
ans Fd
por e
Pq t ch
2 ain e 8
e  e NADP
2H  H2O Cytochrome
NADP + H
+ complex
3 reductase
1
/ 2 O2 NADPH
e Pc
e P700
P680 5 Light

1 Light 6

ATP

Pigment
molecules
Photosystem I
(PS I)
Photosystem II
(PS II)
 e

e e
Mill
makes
ATP NADPH
e 
e
e

n
Photo
e
ATP
n
Photo

Photosystem II Photosystem I
 Cyclic Electron Flow
• Cyclic electron flow uses only photosystem I
and produces ATP, but not NADPH
• No oxygen is released
• Cyclic electron flow generates surplus ATP,
satisfying the higher demand in the Calvin cycle
 Primary
Primary acceptor
Fd
acceptor Fd
Pq NADP
NADP + H
reductase
Cytochrome NADPH
complex

Pc

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
 A Comparison of Chemiosmosis in
Chloroplasts and Mitochondria
• 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
 Mitochondrion Chloroplast

MITOCHONDRION CHLOROPLAST
STRUCTURE STRUCTURE
H Diffusion
Intermembrane Thylakoid
space space
Electron
Inner Thylakoid
transport
membrane membrane
chain

ATP
synthase
Matrix Stroma
ADP  P i ATP
Key Higher [H ]

H 

Lower [H ]
 A Comparison of Chemiosmosis in
Chloroplasts and Mitochondria

• 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
STROMA
(low H concentration) Cytochrome NADP
complex Photosystem I reductase
Photosystem II
Light
Light 4 H+ 3
NADP + H
Fd

Pq
NADPH
Pc
2
H2O
1 /2 O 2
1

THYLAKOID SPACE +2 H+ 4 H+
(high H concentration)
To
Calvin
Cycle

Thylakoid
membrane ATP
synthase
ADP
+ ATP
STROMA
Pi
(low H concentration) H+
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
The Calvin cycle uses the chemical energy
of ATP and NADPH to reduce CO2 to sugar
• 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 CO 2
• The Calvin cycle has three phases
– Carbon fixation (catalyzed by rubisco)
– Reduction
– Regeneration of the CO2 acceptor (RuBP)
 Input
3 (Entering one
at a time)
CO2

Phase 1: Carbon fixation

Rubisco
3 P P
Short-lived
intermediate
3P P 6 P
Ribulose bisphosphate 3-Phosphoglycerate
(RuBP) 6 ATP
6 ADP

3 ADP Calvin
Cycle
3 6 P P
ATP
1,3-Bisphosphoglycerate
6 NADPH
Phase 3:
Regeneration of 6 NADP
the CO2 acceptor 6 Pi
(RuBP) P
5
G3P
6 P
Glyceraldehyde 3-phosphate Phase 2:
(G3P) Reduction

1 P
G3P Glucose and
(a sugar) other organic
Output compounds
The Importance of Photosynthesis: A Review
• 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
 H2O CO2

Light
NADP
ADP
+ Pi
Light RuBP
Reactions: 3-Phosphoglycerate
Photosystem II Calvin
Electron transport chain Cycle
Photosystem I
Electron transport chain
ATP
G3P
Starch
NADPH (storage)

Chloroplast

O2 Sucrose (export)