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
• Photosynthesis occurs in plants, algae, certain
other protists, and some prokaryotes
• These organisms feed not only themselves but
also most of the living world
 (a) Plants

(c) Unicellular protist

10 µm

(e) Purple sulfur

1.5 µm
bacteria

(b) Multicellular alga (d) Cyanobacteria

40 µm
• 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 from H2O and CO2
• 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 O2
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 (organ) 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
• Chloroplasts are found mainly in cells of the
mesophyll, the interior tissue of the leaf
• A typical mesophyll cell has 30–40 chloroplasts

• The chlorophyll is in the membranes of

thylakoids (connected sacs in the chloroplast);
thylakoids may be stacked in columns called
grana (granum)
• Chloroplasts also contain stroma, a dense
fluid
 Leaf cross section

Vein

Mesophyll

Stomata
CO2 O2

Chloroplast
Mesophyll cell

Outer
membrane
Thylakoid
Intermembrane 5 µm
Stroma Thylakoid space
Granum
space
Inner
membrane

1 µm
Tracking Atoms Through Photosynthesis

• Photosynthesis can be summarized as the

following equation:

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 as a Redox Process

• Photosynthesis is a redox process in which

H2O is oxidized and CO2 is reduced

Reactants: 6 CO2 12 H2O

Products: C6H12O6 6 H2O 6 O2

The Two Stages of Photosynthesis
• 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 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
+ P
i
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
• 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 photons
 1m
10–5 nm 10–3 nm 1 nm 103 nm 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
• A spectrophotometer measures a pigment’s
ability to absorb various wavelengths
• This machine sends light through pigments and
measures the fraction of light transmitted at
each wavelength
TECHNIQUE

White Refracting Chlorophyll Photoelectric

light prism solution tube
Galvanometer
2 3
1 4

The high transmittance

Slit moves to Green (low absorption)
pass light light reading indicates that
of selected chlorophyll absorbs
wavelength very little green light.

The low transmittance

Blue (high absorption)
light reading indicates that
chlorophyll absorbs
most blue light.
• An absorption spectrum is a graph plotting a
pigment’s light absorption versus wavelength
• The absorption spectrum of chlorophyll a
suggests that violet-blue and red light work
best for photosynthesis
• An action spectrum profiles the relative
effectiveness of different wavelengths of
radiation in driving a process
 RESULTS

Absorption of light by
Chloro-

chloroplast pigments
phyll a Chlorophyll b

Carotenoids

(a) Absorption spectra 400 500 600 700

Wavelength of light (nm)

(measured by O2 release)
Rate of photosynthesis

(b) Action spectrum

Aerobic bacteria
Filament
of alga

(c) Engelmann’s
experiment 400 500 600 700
• The action spectrum of photosynthesis was
first demonstrated in 1883 by Theodor W.
Engelmann
• In his experiment, he exposed different
segments of a filamentous alga to different
wavelengths
• Areas receiving wavelengths favorable to
photosynthesis produced excess O2
• He used the growth of aerobic bacteria
clustered along the alga as a measure of O2
production
• 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
 CH3 in chlorophyll a
CHO in chlorophyll b

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

Hydrocarbon tail:
interacts with hydrophobic
regions of proteins inside
thylakoid membranes of
chloroplasts; H atoms not
shown
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) funnel the energy
of photons to the reaction center
• A primary electron acceptor in the reaction
center accepts an excited electron from
chlorophyll a
• Solar-powered transfer of an electron from a
chlorophyll a molecule to the primary electron
acceptor is the first step of the light reactions
 Photosystem STROMA
Photon
Light-harvesting Reaction-center Primary
complexes complex electron
acceptor
Thylakoid membrane

e–

Transfer Special pair of Pigment

of energy chlorophyll a molecules
molecules
THYLAKOID SPACE
(INTERIOR OF THYLAKOID)
• 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
• Photosystem I (PS I) is best at absorbing a
wavelength of 700 nm
Non Cyclic Electron Flow

• During the light reactions, there are two

possible routes for electron flow: cyclic and
non-cyclic
• Non cyclic is the primary pathway, involves
both photosystems and produces ATP and
NADPH using light energy
• A photon hits a pigment and its energy is
passed among pigment molecules until it
excites P680…this excited electron from P680
is transferred to the primary electron acceptor
• 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 reaction

• 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
• 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
• 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
 E
tra lect
n ro
c h s po n
Primary a i n rt
E le acceptor
Primary ctro 4 7
n tr
acceptor ans Fd
p o rt
Pq cha e–
2 in e– 8
e– e– NADP+
+ H2O Cytochrome
2H complex NADP+ + H+
+ reductase
3 NADPH
/2 O2
1
Pc
e–
e– P700
P680 5 Light

1 Light 6

ATP

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

• 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
• Some organisms such as purple sulfur bacteria
have PS I but not PS II. Cyclic electron flow is
thought to have evolved before linear electron
flow. Cyclic electron flow may protect cells from
light-induced damage
 Primary
Primary acceptor
Fd
acceptor Fd
NADP+
Pq
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
• 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 chain membrane

ATP
synthase
Matrix Stroma
Key
ADP + P i
ATP
Higher [H+] H+
Lower [H+]
• 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
Photosystem II Photosystem I
complex
4 H+ Light NADP+
Light reductase
Fd 3
NADP+ + H+

Pq NADPH

e– Pc
e– 2
H2O
THYLAKOID SPACE 1 1
/2 O 2
(high H+ concentration) +2 H+ 4 H+

To
Calvin
Cycle

Thylakoid
membrane ATP
synthase
STROMA
ADP
(low H+ concentration)
+ ATP
Pi
H+
The Calvin cycle uses ATP and NADPH to convert
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
• Carbon enters the cycle as CO2 and leaves as
a sugar named glyceraldehyde-3-phospate
(G3P) or PGAL
• For net synthesis of 1 G3P, the cycle must take
place three times, fixing 3 molecules of CO2

• 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)
5 P
G3P
6 P
Glyceraldehyde-3-phosphate Phase 2:
(G3P) Reduction

1 P Glucose and
Output G3P other organic
(a sugar) compounds
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 H2O but also limits photosynthesis

• The closing of stomata reduces access to CO2

and causes O2 to build up

• These conditions favor a seemingly wasteful

process called photorespiration
Photorespiration

• In most plants (C3 plants), initial fixation of

CO2, via rubisco, forms a three-carbon
compound

• In photorespiration, rubisco adds O2 instead

of CO2 in the Calvin cycle

• Photorespiration consumes O2 and organic fuel

and releases CO2 without producing ATP or
sugar
• 1. Rice
2. Wheat
3. Oranges
4. Grapes
5. Coffee plant
6. Tea Plants
7. Peanut
8. Lemon
9. Potatoes
• 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
• 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
• C4 plants minimize the cost of photorespiration by
incorporating CO2 into four-carbon compounds in
mesophyll cells
• This step requires the enzyme PEP 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
 C4 leaf anatomy The C4 pathway
Mesophyll
Mesophyll cell cell CO2
Photosynthetic PEP carboxylase
cells of C4 Bundle-
plant leaf sheath
cell
Oxaloacetate (4C) PEP (3C)
Vein ADP
(vascular tissue)
Malate (4C) ATP

Pyruvate (3C)
Bundle-
Stoma sheath CO2
cell
Calvin
Cycle

Sugar

Vascular
tissue
• 1. Crab Grass
2. Corn
3. Amaranth
4. Sorgham
5. Millet
6. Sugarcane
7. Nut grass
8. Barnyard grass
9. fourwinged salt bush.
CAM Plants

• Some plants, including succulents, use

crassulacean acid metabolism (CAM) to fix
carbon
• CAM plants open their stomata at night,
incorporating CO2 into organic acids

• Stomata close during the day, and CO2 is

released from organic acids and used in the
Calvin cycle
Sugarcane Pineapple
C4 CAM
CO2 CO2
Mesophyll 1 CO2 incorporated Night
cell Organic acid into four-carbon Organic acid
organic acids
(carbon fixation)

Bundle- CO2 CO2 Day

sheath
cell
2 Organic acids
Calvin release CO2 to Calvin
Cycle Cycle
Calvin cycle

Sugar Sugar

(a) Spatial separation of steps (b) Temporal separation of steps

• Thanks for listening!