Chapter 10

Photosynthesis

● Overview: The Process That Feeds the

Biosphere
● Photosynthesis
○ Is the process that converts solar energy into
chemical energy
● Plants and other autotrophs
○ Are the producers of the biosphere

● Plants are photoautotrophs

○ They use the energy of sunlight to make
organic molecules from water and carbon
dioxide
 Figure
10.1

● Photosynthesis
○ Occurs in plants, algae, certain other protists,
and some prokaryotes
These organisms use light energy to drive the
synthesis of organic molecules from carbon dioxide
and (in most cases) water. They feed not only
themselves, but the entire living world. (a) On
land, plants are the predominant producers of
food. In aquatic environments, photosynthetic
organisms include (b) multicellular algae, such
as this kelp; (c) some unicellular protists, such
as Euglena; (d) the prokaryotes called
cyanobacteria; and (e) other photosynthetic
prokaryotes, such as these purple sulfur
bacteria, which produce sulfur (spherical (a) Plants
globules) (c, d, e: LMs).

(c) Unicellular 10
protist m

(e) Pruple 1.5

sulfur m
bacteria

Figure (b) Multicellular (d)

40
algae Cyanobacteria
10.2 m

● Heterotrophs
○ Obtain their organic material from other
organisms
○ Are the consumers of the biosphere
 ● Concept 10.1: Photosynthesis converts light
energy to the chemical energy of food

Chloroplasts: The Sites of Photosynthesis in

Plants
● The leaves of plants
○ Are the major sites of photosynthesis

Leaf cross
section
Vein

Mesophyll
 CO O
Stomat
a 2 2

Figure
10.3

● Chloroplasts
○ Are the organelles in which photosynthesis occurs
○ Contain thylakoids and grana

Mesophyl
l

Chloroplas
t

5
µm

Outer
membran
e
Thylakoi Thylakoi Intermembran
Strom Granu d d e
a m space Inner space
membran
e

1
µm

Tracking Atoms Through Photosynthesis:

Scientific Inquiry
● Photosynthesis is summarized as
 6 CO2 + 12 H2O + Light energy  C6H12O6 + 6 O2 + 6 H2
O

The Splitting of Water

● Chloroplasts split water into
○ Hydrogen and oxygen, incorporating the
electrons of hydrogen into sugar molecules

Reactants 6 12 H2O
: CO2

Products C6H12O6 6 6
: H2 O O2

Figure
10.4

Photosynthesis as a Redox Process

● Photosynthesis is a redox process
○ Water is oxidized, carbon dioxide is reduced
The Two Stages of Photosynthesis: A Preview
● Photosynthesis consists of two processes
○ The light reactions
○ The Calvin cycle

● The light reactions

○ Occur in the grana
 ○ Split water, release oxygen, produce ATP, and
form NADPH

● The Calvin cycle

○ Occurs in the stroma
○ Forms sugar from carbon dioxide, using ATP
for energy and NADPH for reducing power
● An overview of photosynthesis

H2 CO
O 2

Ligh
t
NADP 
ADP
+P
LIGHT CALVIN
REACTIONS CYCLE

ATP

NADP
H

Chloroplas
t
[CH2O]
O
Figure (sugar)
2
10.5

● Concept 10.2: The light reactions convert

solar energy to the chemical energy of ATP and
NADPH
The Nature of Sunlight
● Light
○ Is a form of electromagnetic energy, which
travels in waves

● Wavelength
○ Is the distance between the crests of waves
○ Determines the type of electromagnetic
energy
 ● The electromagnetic spectrum
○ Is the entire range of electromagnetic energy, or
radiation

1m
10–5 10–3 1 103 106 106 103
nm nm nm nm nm nm m
Gamm Micro- Radio
a X- UV Infrare wave waves
rays rays d s

Visible
light

38 45 50 55 60 65 70 750
0 0 0 0 0 0 0 nm
Shorter Longer
wavelength wavelength
Higher Lower
Figure energy energy
10.6

● The visible light spectrum

○ Includes the colors of light we can see
○ Includes the wavelengths that drive
photosynthesis
Photosynthetic Pigments: The Light
Receptors
● Pigments
○ Are substances that absorb visible light

● Reflect light, which include the colors we see

Ligh
t
Reflecte
d
Light
Chloroplas
t
 Absorbed Granu
light m

Transmitte
d
Figure light
10.7

● The spectrophotometer
○ Is a machine that sends light through
pigments and measures the fraction of light
transmitted at each wavelength

● An absorption spectrum
○ Is a graph plotting light absorption versus wavelength

Whit Refractin Chlorophyl Photoelectri

e g l c
light prism solution tube Galvanometer
2 3

1 0 10
4 0

Slit moves Gree The high transmittance

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

0 10
0

The low transmittance

Blu
(high absorption) reading
e
Figure chlorophyll absorbs most blue light.
light
10.8

● The absorption spectra of chloroplast

pigments
○ Provide clues to the relative effectiveness of
different wavelengths for driving photosynthesis

● The absorption spectra of three types of

pigments in chloroplasts
Three different experiments helped reveal which wavelengths of light are photosynthetically important. The
EXPERIMENT
results are shown below.

RESULTS
Chlorophyll
a
Chlorophyll
 b

Absorption of light Carotenoid

by s
chloroplast pigments

Wavelength of light (nm)

(a) Absorption spectra. The three curves show the wavelengths of light best absorbed by
�three types of chloroplast pigments.
Figure
10.9

● The action spectrum of a pigment

○ Profiles the relative effectiveness of different
wavelengths of radiation in driving photosynthesis

Rate of photosynthesis
(measured by O2
release)

(b Action spectrum. This graph plots the rate of photosynthesis versus wavelength.
) �The resulting action spectrum resembles the absorption spectrum for chlorophyll
�a but does not match exactly (see part a). This is partly due to the absorption of
light �by accessory pigments such as chlorophyll b and carotenoids.

● The action spectrum for photosynthesis

○ Was first demonstrated by Theodor W. Engelmann

Aerobic
bacteria
 Filamen
t
of alga

40 50 60 70
0
(c 0Engelmann‘s experiment. In 1883, 0 0
Theodor W. Engelmann illuminated a filamentous alga with light that had
) been passed through a prism, exposing different segments of the alga to different wavelengths. He used aerobic
bacteria, which concentrate near an oxygen source, to determine which segments of the alga were releasing the
most O2 and thus photosynthesizing most.
Bacteria congregated in greatest numbers around the parts of the alga illuminated with violet-blue or red light.
Notice the close match of the bacterial distribution to the action spectrum in part b.

Light in the violet-blue and red portions of the spectrum are most effective in driving photosynthesis.
CONCLUSION

● Chlorophyll a
○ Is the main photosynthetic pigment
● Chlorophyll b
○ Is an accessory pigment CH
3
CH
in chlorophyll
a
in chlorophyll
CH O b
2 CH
C H
H 3
C C C
H3 C C C C CH CH
Porphyrin ring:
C
Light-absorbing
C N N C 2 3
“head” of
H C M C H molecule
C N g N C note magnesium
H3
C C
C C C CH atom at center
H C C C 3
H
CH H C C
2 O
CH O
2
C O O

O CH
3
CH
2

Hydrocarbon tail:
interacts with
hydrophobic
regions of proteins inside
thylakoid membranes of
chloroplasts: H atoms not
shown
Figure
10.10

● Other accessory pigments

 ○ Absorb different wavelengths of light and pass
the energy to chlorophyll a

Excitation of Chlorophyll by Light

● When a pigment absorbs light
○ It goes from a ground state to an excited state,
which is unstable

Excite
e d
–
state

Hea
t
Energy of
election

Photon
(fluorescence
)
Groun
Photo Chlorophyl d
n l state
molecule
Figure 10.11 A

● If an isolated solution of chlorophyll is

 illuminated
○ It will fluoresce, giving off light and heat

Figure 10.11 B

A Photosystem: A Reaction Center

Associated with Light-Harvesting Complexes
● A photosystem
○ Is composed of a reaction center surrounded by a
number of light-harvesting complexes

Thylakoi
d

Photo Photosyste
n m
STROM
Light- Reaction Primary election
A
harvesting center acceptor
complexes

e
–
Thylakoid
membrane

Transfer Special Pigment

of chlorophyll molecule
energy a s
molecules THYLAKOID SPACE
(INTERIOR OF
Figure THYLAKOID)
10.12

● The light-harvesting complexes

○ Consist of pigment molecules bound to
particular proteins
○ Funnel the energy of photons of light to the
reaction center
● When a reaction-center chlorophyll molecule
absorbs energy
○ One of its electrons gets bumped up to a
primary electron acceptor

● The thylakoid membrane

○ Is populated by two types of photosystems, I
and II
Noncyclic Electron Flow
● Noncyclic electron flow
○ Is the primary pathway of energy
transformation in the light reactions

● Produces NADPH, ATP, and oxygen

H2 CO
Ligh O 2
t
NADP
+
AD
P
LIGHT CALVIN
REACTION CYCLE
S AT
P
NADP
H

Electron
O [CH2O] Transport
2 (sugar) chain
Primary
accepto 7
Primary r
4
accepto
r F
P Electron transport chain
2 d
q e 8
e e
H2 NADP
Cytochrome –
2 O +
complex NADP+
H++ reductas + 2 H+
3
O e NADP
P H
2 e C
– +
e 5 P70
Ligh H+
– 0
t1 P68 Ligh
0 t
6
 ATP

Photosystem-
Photosystem
Figure II
I
(PS I)
10.13 (PS II)

● A mechanical analogy for the light reactions

e
ATP –

e e
– –

NADP
e H
– e
e –
–

Mill
make
Photo
s
n
ATP

e
–

Photo
n Photosystem Photosystem
Figure II I
10.14

Cyclic Electron Flow

● Under certain conditions
○ Photoexcited electrons take an alternative
path
 ● In cyclic electron flow
○ Only photosystem I is used
○ Only ATP is produced

Primary
Primary accepto
F
accepto r
d F
r d NADP
P
q NADP+ +

Cytochrom reductas
e NADP
e H
complex
P
c

ATP Photosystem
Photosystem
Figure II
I
10.15

A Comparison of Chemiosmosis in
Chloroplasts and Mitochondria
● Chloroplasts and mitochondria
○ Generate ATP by the same basic mechanism:
chemiosmosis
○ But use different sources of energy to
accomplish this
● The spatial organization of chemiosmosis
○ Differs in chloroplasts and mitochondria

Ke
y
Higher
Lower
[H+]
[H +]
Mitochondrio Chloroplas
n t

MITOCHONDRION CHLOROPLAS
STRUCTURE T
H Diffusio STRUCTURE
Intermembranc + n Thylakoi
e Electron d
space transpor space
Membranc
e t
ATP
chain
Synthas Strom
Matri e a
ADP P
x ATP
+ H
Figure +
10.16

● In both organelles
○ Redox reactions of electron transport chains
generate a H+ gradient across a membrane
● ATP synthase
○ Uses this proton-motive force to make ATP
 ● The light reactions and chemiosmosis: the
organization of the thylakoid membrane
H2 CO
O 2
LIGHT
NADP
+
AD
P CALVIN
LIGHT
CYCLE
REACTOR
ATP
NADP
H

STROMA O [CH2O]
(Low H+ Cytochrom
2 (sugar)
Photosystem e Photosystem
concentration)
II complexLight I NADP+
2 reductas 3
H+ F e NADP+ +
d 2H+
NADP +
P H H+
q P
2 c
H2 1
⁄ O
THYLAKOID SPACE O 1 2
2 +2
2
(High H+ H+ H+
concentration) To
Calvi
n
cycle

ATP
Thylakoid synthas
STROMA membran e AD
(Low H+ e P ATP
concentration) P
H
Figure +

10.17

● Concept 10.3: The Calvin cycle uses ATP

and NADPH to convert CO2 to sugar
● The Calvin cycle
 ○ Is similar to the citric acid cycle
○ Occurs in the stroma

● The Calvin cycle has three phases

○ Carbon fixation
○ Reduction
○ Regeneration of the CO2 acceptor

● The Calvin cycle

 H2
Ligh O
CO
2
Inpu
t NADP 3 t (Entering
+
AD CO one
LIGHT�R P CALVIN at a time)
�CYC
EACTION
ATP LE 2
NADP
Phase 1: Carbon fixation
H

Rubisc
O [CH2O]
2 (sugar)
o 3P P

Short-
lived�inter P
3P P 6
Ribulose mediate
3-
bisphosphate�(RuBP Phosphoglycerate 6 ATP
) 6 ADP

3 ADP CALVIN
CYCLE 6P P
3 ATP
1,3-
Bisphoglycerate 6
Phase NADPH
3:�Regeneratio 6
+
n of�the CO2 6 NADPH
P

acceptor�(RuB 5 P
P) (G3P 6 P
) Glyceraldehyde-3- Phase
phosphate 2:�Redu
(G3P) ction

1 P
G3P�(a Glucose
Figure sugar)� and�other
10.18 Output organic�co
mpounds

● Concept 10.4: Alternative mechanisms of

carbon fixation have evolved in hot, arid
climates
 ● On hot, dry days, plants close their stomata
○ Conserving water but limiting access to CO2
○ Causing oxygen to build up

Photorespiration: An Evolutionary Relic?

● In photorespiration
○ O2 substitutes for CO2 in the active site of the
enzyme rubisco
○ The photosynthetic rate is reduced
C4 Plants
● C4 plants minimize the cost of
photorespiration
○ By incorporating CO2 into four carbon
compounds in mesophyll cells

● These four carbon compounds

○ Are exported to bundle sheath cells, where
they release CO2 used in the Calvin cycle
 ● C4 leaf anatomy and the C4 pathway
Mesophyll cell Mesophyl
Photosynthetic l
cell PEP CO
CO
cells of C4 Bundle carboxylase 22
plant -
leaf sheath
cell

Oxaloacetate (4 PEP (3
Vein C) C) AD
(vascular P
tissue) Malate (4 ATP
C)

C4 leaf
anatomy Bundle Pyruate (3
- C)
Sheath CO
cell 2
Stom
a
CALVIN
CYCLE

Suga
r

Vascular
tissue

Figure
10.19

CAM Plants
● CAM plants
○ Open their stomata at night, incorporating CO2
into organic acids
● During the day, the stomata close
○ And the CO2 is released from the organic
acids for use in the Calvin cycle

● The CAM pathway is similar to the C4

pathway

Sugarcan Pineappl
e e
C CA
4
CO M CO
Mesophyll Cell
2 2
Night
Organic 1 CO2 Organic
 Bundle- acid incorporated acid
sheath into four-carbon Day
cell organic acids
(a Spatial separation (carbon fixation) (b Temporal separation
) of steps. In C4 CALVIN 2 Organic acids CALVIN ) of steps. In CAM
plants, carbon fixation �CYC release CO2 �CYC plants, carbon fixation
and the Calvin cycle LE LE and the Calvin cycle
to
occur in different Calvin cycle occur in the same
Suga Suga
Figure types of cells.
r
cells�at different times.
r
10.20

The Importance of Photosynthesis: A Review

● A review of photosynthesis
Light Calvin
reaction
H2 cycle
CO
O 2
Ligh
t NADP
+
AD
+P
P
1 RuB 3-
Photosystem II P Phosphoglycerate
Electron transport
chain
Photosystem I ATP G3
P Starch
NADP
(storage
H )
Amino
acids
Chloroplas Fatty acids
t O Sucrose
(export)
2 Calvin cycle reactions:
Light reactions:
• Take place in the stroma
• Are carried out by molecules in the
• Use ATP and NADPH to convert
thylakoid membranes
CO2 to the sugar G3P
• Convert light energy to the
chemical • Return ADP, inorganic phosphate,
energy of ATP and NADPH and �NADP+ to the light reactions
Figure • Split H2O and release O2 to the
10.21 atmosphere

● Organic compounds produced by

photosynthesis
○ Provide the energy and building material for
ecosystems