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8.1 Overview of Photosynthesis 8.1 Overview of Photosynthesis

•  Photosynthesis converts solar energy
into chemical energy of carbohydrates
•  Organisms that carry on photosynthesis
are called autotrophs
–  Plants, algae, and cyanobacteria are
organisms capable of photosynthesis
•  Heterotrophs are organisms that feed on
other organisms
•  Autotrophs and heterotrophs use organic
molecules produced by photosynthesis
•  Pigments allow photosynthetic organisms to
capture solar energy
•  Most photosynthetic organisms contain the
pigment chlorophyll
•  Another common pigment group are
carotenoids

1 2

Flowering Plants as
8.1 Overview of Photosynthesis
Photosynthesizers
– Water is taken up by roots and transported
•  Photosynthesis occurs in the green parts to leaves by veins
of plants
– Carbon dioxide enters through openings in
–  Particularly leaves, contain chlorophyll and the leaves called stomata
other pigments
– Light energy is absorbed by chlorophyll and
•  Leaves contain mesophyll tissue specialized other pigments in thylakoids of chloroplasts
for photosynthesis
•  Raw materials are water and CO2

3 4

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•  Chloroplast structure
–  The chloroplast and its fluid-filled interior called cuticle

stroma are surrounded by a double membrane Leaf cross section

upper
epidermis

–  Thylakoids are a different membrane system

mesophyll
within the stroma that form flattened sacs
–  Thylakoids are stacked together to from grana
lower
CO2 epidermis

O2

–  Thylakoid space is formed by a continuous

Leaf vein
stomata

connection between individual thylakoids

5 Figure 8.2 6

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cuticle
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Leaf cross section
upper
inner membrane epidermis
outer membrane

mesophyll
stroma
stroma

lower
granum epidermis
CO2

O2
leaf vein
stomata
inner membrane
Chloroplast Chloroplast, micrograph 37,000x outer membrane

stroma
stroma

granum

thylakoid space
thylakoid membrane
Chloroplast Chloroplast, micrograph 37,000x

Grana
channel between
thylakoids
thylakoid space
© Dr. George Chapman/Visuals Unlimited
thylakoid membrane

Grana
channel between
Figure 8.2 7 Figure 8.2 thylakoids
© Dr. George Chapman/Visuals Unlimited 8

Photosynthetic Reaction Two Sets of Reactions

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solar energy •  Photosynthesis consists of two sets of reactions

CO2 + 6 H2O C6H12O6 + 6 O2
pigments –  Photo refers to capturing solar energy
–  Synthesis refers to producing a carbohydrate
•  Glucose and oxygen are the products of
photosynthesis
•  The two sets of reactions are called the:
•  The oxygen given off comes from water
–  Light Reactions (light-dependent)
•  CO2 gains hydrogen atoms and becomes –  Calvin Cycle Reactions (light-independent)
a carbohydrate •  Nicotinamide adenine dinucleotide phosphate (NADP
+) links these reactions
9 10

8.2 Plants as Solar Energy

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Converters
H 2O CO2
•  During the light reactions, different
pigments within the thylakoid membranes
solar
energy

absorb energy
ADP + P

NADP+
Calvin
•  Solar energy can be described in terms of
cycle
Light
reactions NADPH
reactions
its wavelength and energy content
ATP

thylakoid stroma
membrane

O2 CH2O

Figure 8.3 11 12

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8.2 Plants as Solar Energy

Visible Light
Converters
•  The electromagnetic •  Visible light contains various wavelengths
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spectrum extends from Increasing wavelength

•  The colors of visible light range from:

very short gamma rays to
very long radio waves –  Violet light
Increasing energy •  Shortest wavelength but high energy
•  White or visible light is Gamma Micro- Radio
rays
X rays UV Infrared
waves waves –  Red light
only a small portion of the
•  Longest wavelength but lowest energy
spectrum
visible light
–  Only about 42% of solar radiation that hits Earth’s
•  Visible light is further atmosphere reaches the surface of Earth – most is in
divided into wavelengths 380 500 600 750 the visible-light range
Wavelengths (nm)
between 380 and 750 nm –  Higher wavelengths are screened by the ozone layer
Figure 8.4 13 14

Visible Light Visible Light

•  Most photosynthetic •  Green light is reflected and only minimally
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pigments in cells are

chlorophylls a and b
absorbed
and the carotenoids Chlorophyll a –  Leaves appear green
Chlorophyll b

•  Can absorb specific carotenoids

•  Other plant pigments become noticeable
various portions of in the fall when chlorophyll breaks down
Relative Absorption

visible light and the other pigments are uncovered

•  The absorption
spectrum shown in
figure on the right 380 500 600 750
Wavelengths (nm)

Figure 8.5 15 16

Light Reactions
Noncyclic Electron Pathway
•  Light Reactions •  Noncyclic electron pathway, named because
–  Take place in thylakoid membrane electron flow is traced from water to NADP+
–  Light reactions consist of two pathways: –  Uses two photosystems (Photosystems I and II)
•  Noncyclic electron pathway –  A photosystem consists of a pigment complex
•  Cyclic electron pathway and electron acceptors within the thylakoid
–  Both pathways transform solar energy to membrane
chemical energy –  The pigment complex can be described as a
–  Both pathways produce ATP “antenna” for gathering solar energy
–  Only the noncyclic pathway produces NADPH
17 18

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Noncyclic Electron Pathway Noncyclic Electron Pathway

•  Noncyclic Electron Pathway begins with •  Photosystem II would disintegrate without
photosystem II (PSII) replacement electrons
–  Pigment complex absorbs solar energy –  Electrons provided by splitting water
–  Energy passes from one pigment to another –  Releases oxygen (O2) to atmosphere which
until it is concentrated in reaction center
benefits all organisms that use O2
•  Chlorophyll a molecule
–  Hydrogen ions (H+) stay in the thylakoid space
–  Electrons in the reaction center chlorophyll
•  Contribute to formation of hydrogen ion gradient
become so energized
•  Escape from the reaction center and move to a
nearby electron acceptor
19 20

Noncyclic Electron Pathway Noncyclic Electron Pathway

•  In PSII, an electron acceptor receives energized •  PSI comes next in noncyclic electron pathway
electrons from the reaction center
–  When the photosystem I complex absorbs solar
•  It sends those electrons down an electron energy, energized electrons leave reaction
transport chain, (series of carriers that pass center and are captured by a different electron
electrons from one to the other) acceptor
•  Energy is released to pump hydrogen ions (H+) •  Low energy PSII electrons used to replace those lost
into thylakoid space forming gradient by PSI

•  When hydrogen ions flow through ATP synthase –  Electron acceptor in photosystem I passes its
it makes ATP electrons to NADP+ and it becomes NADPH

21 22

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H 2O CO2

solar
energy

ADP + P

sun sun sun NADP+

Calvin
sun
cycle
cycle
reactions
Light
reactions NADPH

ATP

electron electron electron thylakoid electron

energy level

energy level

membrane
acceptor acceptor acceptor O2 CH2O
acceptor

e– e–
ele e– ele e–
e– ctr e– ctr
on e– NADP+ on e– NADP+
tr ans tr ans
po po
rt c H+ rt c H+
hain hain

e– e– NADPH e– e– NADPH

reaction center reaction center reaction center reaction center

pigment pigment
pigment complex pigment complex
complex Photosystem I complex Photosystem I
e– Photosystem II e– Photosystem II

H 2O CO2 CH2O H 2O CO2 CH2O

Calvin cycle Calvin cycle
1
reactions 1
reactions
2H+ 2 O2
– 2H+ 2 O2
–
Figure 8.6 23 Figure 8.6 24

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Cyclic Electron Pathway

•  Energized electrons
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•  Uses only photosystem I (PSI) and begins leave the photsytem I sun

when PSI complex absorbs solar energy reaction center and

–  Energized electrons escape from the reaction return to photosystem electron

by an electron
acceptor

energy level
center and travel down electron transport chain ATP

–  Released energy is stored in the form of a H+ transport chain e–

gradient, which causes ATP production by ATP •  ATP from cyclic e– CO2 CH2O

synthase reaction center Calvin cycle

electron transport reactions and

other enzymatic
reactions

•  Spent electrons return to PSI (cyclic) used in Calvin cycle to Pigment

complex

make carbohydrates
Photosystem I

•  Pathway only results in ATP production

25 Figure 8.7 26

The Organization of the The Organization of the

Thylakoid Membrane Thylakoid Membrane
•  The following molecular complexes are –  PS I
present in the thylakoid Membrane: •  Pigment complex and electron acceptors
–  PS II •  Adjacent to enzyme that reduces NADP+ to
•  Pigment complex and electron acceptors NADPH
•  Water is split to replace energized electrons –  ATP synthase complex
•  Oxygen (O2) is released •  Has a channel for H+ flow
–  Electron transport chain •  Flow drives ATP synthase to join ADP and P
•  Carries electrons from PS II to PS I
•  Uses energy to pump H+ from the stroma into
thylakoid space
27 28

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thylakoid membrane
thylakoid
thylakoid space
granum

photosystem II photosystem II
electron transport electron transport
H+ chain H+ chain
stroma

photosystem I photosystem I
NADP H+ NADP H+
reductase reductase
Pq Pq
e– e–
e– e
–
e– e
–
NADP+ NADPH NADP+ NADPH

e– e– e– e–
H+ H+
H+ H+
1 1
H 2O 2 H+ + 2 O2 H 2O 2 H+ + 2 O2

ATP synthase ATP synthase

H+ complex H+ complex
H+ H+

ATP ATP
thylakoid thylakoid
space H+ space H+
H+ H+
chemiosmosis chemiosmosis
P + ADP P + ADP

Stroma Stroma

Figure 8.8
29
Figure 8.8
30

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ATP Production
H 2O CO2

solar
energy

ADP + P
NADP+ Calvin
cycle
Light reactions
reactions NADPH

ATP

thylakoid
membrane
thylakoid membrane
thylakoid space
thylakoid
•  ATP Production
O2 CH2O

photosystem II
granum
–  Thylakoid space acts as a reservoir for hydrogen
H+
electron transport
chain
stroma ions (H+)
photosystem I

•  H+ from water being split within thylakoid space

NADP H+
reductase
Pq
e–
e– e– NADP+ NADPH
•  Pumped in by electron transport chain
e– e–

H 2O 2 H+ +
1
2 O2
H+
H+ –  More H+ in thylakoid space than stroma
ATP synthase
•  Electrochemical gradient
H+ complex
H+

ATP
–  H+ can only flow through ATP synthase
thylakoid

–  Energy powers making ATP by chemiosmosis

space H+
H+
chemiosmosis
P + ADP

Stroma

Figure 8.8
31 32

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8.3 Plants as Carbon Dioxide solar

energy
H 2O CO2

Fixers Light
reactions
ADP+ P
NADP+

NADPH
Calvin
cycle

Metabolites of the Calvin Cycle

ATP
RuBP ribulose-1,5-bisphosphate

•  The Calvin Cycle (named after Melvin Calvin) stroma

3 CO2
3PG

BPG
3-phosphoglycerate

1,3-bisphosphoglycerate
O2 CH2O
intermediate

–  Series of reactions that use CO2 from the 3 C6

G3P glyceraldehyde-3-phosphate

atmosphere to produce carbohydrate 3 RuBP

6 3PG
C3
C5 CO2 6

–  Humans other most other organisms take in O2 fixation ATP

and release CO2 3ADP + 3 P

Calvin cycle
CO2
reduction 6ADP + 6 P
These ATP and
NADPH
molecules were
produced by the

–  Includes regeneration
of RuBP
6 BPG
light reactions.

C3
•  Carbon dioxide fixation These ATP
molecules were
produced by the
3
ATP 5 G3P 6 NADPH
light reactions. C3

•  Carbon dioxide reduction 6 G3P

C3
6 NADP+

•  Ribulose-1,5-bisphosphate (RuBP) regeneration

net gain of one G3P x2

Other organic molecules Glucose

33 34

Fixation of Carbon Dioxide Reduction of Carbon Dioxide

•  Carbon dioxide fixation is the 1st step of •  Reduction of Carbon Dioxide
the Calvin cycle –  Each 3PG molecules undergoes reduction to
–  CO2 is attached to 5-carbon RuBP molecule G3P in two steps
•  This reaction occurs three times –  Energy and electrons needed for this reaction
•  The result is a 6-carbon molecule that splits into two are supplied by ATP and NADPH (from light
3-carbon molecules 3-phoshoglycerate (3PG) reaction)
–  RuBP Carboxylase is the enzyme that makes
this happen
•  Comparatively slow enzyme so there is a lot of it

35 36

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Reduction of Carbon Dioxide Regeneration of RuBP

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•  Regeneration of RuBP
ATP ADP + P
–  It takes three turns of the Calvin cycle to allow
one G3P to exit
3PG BPG G3P –  For every three turns of Calvin Cycle, five G3P
(3-carbon molecule) used
–  This re-forms three RuBP (5-carbon molecule)
NADPH NADP+
–  5 X 3 (carbons in G3P) = 3 X 5 (carbons in RuBP)

As 3PG becomes G3P ATP becomes

ADP + P , and NADPH becomes NADP+.
37 38

Regeneration of RuBP Importance of the Calvin Cycle

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•  G3P (glyceraldehyde-3-phosphate) can be

5 G3P 3 RuBP converted to many other molecules
–  These molecules meet the plant needs
3 ATP 3 ADP + P
•  The hydrocarbon skeleton of G3P can form:
–  Fatty acids and glycerol to make plant oil
As five molecules of G3P become three
molecules of RuBP, three molecules of ATP –  Glucose phosphate (simple sugar)
become three molecules of ADP + P . –  Fructose (+ glucose = sucrose)
–  Starch and cellulose
5 × 3 (carbons in G3P) = 3 × 5 (carbons in RuBP) –  Amino acids
39 40

8.4 Alternate Pathways for

C3 Photosynthesis
Photosynthesis
•  RuBP carboxylase binds O2 as well as CO2
•  C3 Photosynthesis
–  When bound to O2, the enzyme undergoes
–  The leaves of C3 plants have a different structure
photorespiration
and means of fixing CO2 than C4 plants
–  C3 plants such as wheat, rice, oats have
–  Wasteful reaction because it uses O2 and
mesophyll cells of leaves in parallel layers releases CO2, decreasing output of Calvin cycle

–  Bundle sheath cells around the plant veins do not –  O2 concentration in leaf rises when weather is
contain chloroplasts hot and dry, because plant keeps stomata
closed to conserve water
–  As a result, cells using Calvin cycle exposed to CO2

41 42

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C4 Photosynthesis
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Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. CO2

•  C4 plants, such as sugarcane and corn, the

mesophyll cells are arranged in concentric
rings around the bundle sheath cells
RuBP
mesophyll
cells
Calvin
cycle –  They also contain chloroplasts
3PG
(C3)
–  In the mesophyll cells, CO2 is initially fixed into a
G3P
4-carbon molecule
vein
bundle sheath
cell
stoma mesophyll cell –  The 4-carbon molecule is later broken down into
a. C3 Plant a 3-carbon molecule and CO2
a. CO2 fixation in a C3 plant, tuplip
© The McGraw-Hill Companies, Inc./Evelyn Jo Johnson, photographer –  CO2 enters the Calvin cycle

Figure 8.11 Figure 8.10 43 44

C4 Photosynthesis Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
CO2
•  C4 Pathway
mesophyll
C4
–  C4 plants tend to be found in hot, dry climates cell

–  In these climates, stomata tend to close to conserve bundle

sheath
CO2
mesophyll
water cells cell

Calvin
–  Oxygen then builds-up in the leaves cycle

–  But, RuBP carboxylase is not exposed to this O2 in C4

plants and photorespiration does not occur bundle sheath
vein
G3P
cell stoma

–  Instead, in C4 plants, the CO2 is delivered to the Calvin b. CO2 fixation in a C4 plant, corn
b. C4 Plant
cycle, which is located in bundle sheath cells that are © Corbis RF

sheltered from the leaf air spaces

45
Figure 8.11 Figure 8.10 46

C4 Photosynthesis CAM Photosynthesis

•  When the weather is moderate, C3 plants ordinarily •  CAM Pathway
have the advantage.
–  This pathway is prevalent among most succulent
•  When the weather is hot and dry, C4 plants have plants that grow in deserts, including the cacti.
the advantage, and can be expected to
–  CAM plants partition carbon fixation according to
predominate.
time.
•  In the early summer, C3 plants such as Kentucky •  During the night, CAM plants fix CO2, forming C4
bluegrass predominate in lawns in the cooler parts molecules.
of the United States, but by midsummer,
•  The C4 molecules are stored in large vacuoles.
crabgrass, a C4 plant, begins to take over.
•  During daylight, C4 molecules release CO2 to the
Calvin cycle.
47 48

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CO2
night

C4

day CO2

Calvin
cycle

G3P

c. CO2 fixation in a CAM plant, pineapple

Figure 8.10 © S. Alden/PhotoLink/Getty RF
49