BIOL 110: General Biology I

Chapter 10
Photosynthesis

Henry H. Ruiz, PhD

Biological & Chemical Sciences
NYIT, Old Westbury
 Photosynthesis
• Life on Earth is solar powered

• Photosynthesis is the process that converts solar

energy into chemical energy

• Directly or indirectly, photosynthesis nourishes

almost the entire living world

• Organisms obtain organic compounds by one of two

major modes: autotrophic or heterotrophic nutrition
 Photosynthesis
• 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
• Photosynthesis occurs in plants, algae, certain other
protists, and some prokaryotes
• These organisms feed not only themselves but also
most of the living world
 Photosynthesis

(b) Multicellular
(a) Plants alga

(d) Cyanobacteria
40 µm
(c) Unicellular
10 µm
protists

(e) Purple sulfur

1 µm
bacteria
 Photosynthesis

• Heterotrophs obtain their organic material from

other organisms

• Heterotrophs are the consumers of the biosphere

– Feed on other organisms

• Almost all heterotrophs, including humans, depend

on photoautotrophs for food and O2
 Photosynthesis

• Chloroplasts are structurally similar to and likely

evolved from photosynthetic bacteria

• The structural organization of these cells allows

for the chemical reactions of photosynthesis

• The endosymbiont theory suggests that original

chloroplast was a photosynthetic prokaryote that
lived inside a eukaryotic cell
 Chloroplasts: Sites of Photosynthesis

• Leaves - major locations of photosynthesis

– Green color is from chlorophyll (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

• CO2 enters & O2 exits leaf through pores – stomata

• Chlorophyll is in the membranes of thylakoids
(connected sacs in chloroplast); thylakoids may be
stacked in columns - grana
• Chloroplasts also contain stroma -dense interior fluid
 Leaf cross section
Chloroplasts Vein

Mesophyll

Stomata
CO2 O2

Chloroplast Mesophyll
cell

Outer
membrane
Thylakoid Intermembrane
Stroma Granum Thylakoid space 20 µm
space Inner
membrane

1 µm
 Photosynthesis
• 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

• Chloroplasts split H2O into hydrogen & oxygen,

incorporating electrons of hydrogen into sugar
molecules & releasing oxygen as a by-product

Reactants: 6 CO2 12 H2O

Products: C6H12O6 6 H2O 6 O2

 Photosynthesis: Redox Process
• Photosynthesis reverses direction of electron flow
compared to respiration
• Photosynthesis is a redox process in which H2O is
oxidized & CO2 is reduced
• Photosynthesis is an endergonic process; the energy
boost is provided by light

becomes reduced

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

becomes oxidized
 Photosynthesis: Two stages

• Light reactions (the photo part) & 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 & NADPH
– begins with carbon fixation, incorporating CO2 into organic
molecules
 H2O

Light

NADP+
ADP
+ Pi

Light
Reactions

Chloroplast
 H2O

Light

NADP+
ADP
+ Pi

Light
Reactions

ATP

NADPH

Chloroplast

O2
 H2O CO2

Light

NADP+
ADP
+ Pi
Calvin
Light Cycle
Reactions

ATP

NADPH

Chloroplast

O2
 H2O CO2

Light

NADP+
ADP
+ Pi
Calvin
Light Cycle
Reactions

ATP

NADPH

Chloroplast Light reactions

convert solar
energy to the [CH2O]
O2
chemical energy of (sugar)
ATP and NADPH
 Nature of Sunlight
• Light - form of electromagnetic energy, also called
electromagnetic radiation
– Like other electromagnetic energy, light travels in rhythmic
waves
• Wavelength = distance between crests of waves
– Wavelength determines the type of electromagnetic energy
• Electromagnetic spectrum - 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
 Electromagnetic Spectrum
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: Light Receptors
• Pigments are Light
Reflected
substances that light
absorb visible light
Chloroplast
• Different pigments
absorb different
wavelengths
• Wavelengths that
are not absorbed
are reflected or
transmitted
• Leaves appear green Absorbed Granum
light
because chlorophyll
reflects and
transmits green Transmitted
light
light
 (a) Absorption spectra
• Absorption spectrum - graph

Absorption of light by
Chloro-

chloroplast pigments
plotting a pigment’s light absorption phyll a Chlorophyll b
versus wavelength
• Absorption spectrum of Carotenoids

chlorophyll a: violet-blue & red

light work best for photosynthesis 400 500 600 700
• Action spectrum profiles the Wavelength of light (nm)
relative effectiveness of different

(measured by O2 release)
(b) Action spectrum

Rate of photosynthesis
wavelengths of radiation in driving
a process
• (c) Engelmann’s experiment (1883)
exposed different segments of
filamentous alga to different
wavelengths 400 500 600 700

• Areas receiving wavelengths

Aerobic bacteria
favorable to photosynthesis
Filament
produced excess O2 of alga
• He used the growth of aerobic
bacteria clustered along the alga as
a measure of O2 production 400 500
(c) Engelmann’s experiment
600 700
 CH3 in chlorophyll a
Chlorophylls CH3
CHO in chlorophyll b

• Chlorophyll a is the Porphyrin ring

main photosynthetic
pigment
• Accessory
pigments, such as
chlorophyll b,
broaden the
spectrum used for
photosynthesis
• Accessory pigments Hydrocarbon tail
called carotenoids (H atoms not shown)
absorb excessive
light that would
damage chlorophyll
 Chlorophylls
• When a pigment
absorbs light, it goes
from a ground state to
Excited an excited state,
e− state which is unstable
• When excited
Energy of electron

Heat
electrons fall back to
the ground state,
photons are given off,
an afterglow called
Photon fluorescence
Photon (fluorescence) • If illuminated, an
Ground
Chlorophyll
molecule
state isolated solution of
chlorophyll will
fluoresce, giving off
(a) Excitation of isolated chlorophyll molecule
light and heat
 Photosystem: Photon
Photosystem STROMA
Light- Reaction-
Reaction-Center harvesting
complexes
center
complex
Primary
electron

Complex
acceptor

• A photosystem
consists of a

Thylakoid membrane
reaction-center
complex (a type of e−
protein complex)
surrounded by light-
harvesting
complexes
• The light-
harvesting
Transfer Special pair of Pigment
complexes (pigment chlorophyll a molecules
of energy
molecules bound to molecules
proteins) transfer THYLAKOID SPACE
the energy of (INTERIOR OF THYLAKOID)
photons to the (a) How a photosystem harvests light
reaction center
 Photosystem: Reaction-Center Complex
Chlorophyll STROMA
Thylakoid membrane

Protein
subunits THYLAKOID
SPACE

(b) Structure of photosystem II

 Photosystems
• A primary electron acceptor in the reaction center accepts
excited electrons & 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
• Two types of photosystems in thylakoid membrane
• Photosystem II (PS II) functions first (the numbers reflect
order of discovery) & is best at absorbing a wavelength of
680 nm
• Reaction-center chlorophyll a of PS II - P680
• Photosystem I (PS I) is best at absorbing a wavelength of
700 nm
• Reaction-center chlorophyll a of PS I - 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

• During the light reactions, electrons flow through

the photosystems and other molecular components
built into the thylakoid membrane
 Linear Electron Flow

1. A photon hits a pigment

Primary
acceptor and its energy is passed
e−
2
among pigment molecules
until it excites P680

P680

1 Light 2. An excited electron from

P680 is transferred to
the primary electron
Pigment
molecules acceptor (we now call it
Photosystem II
(PS II)
P680+)
 Linear Electron Flow
3. P680+ is a very strong
Primary
acceptor
oxidizing agent
2 H+
H2O e−
2
4. H2O is split by enzymes,
1/
2
+
O2 3 and the electrons are
e−
e− transferred from the
P680
hydrogen atoms to
P680+, thus reducing it
1 Light

to P680
Pigment
molecules
5. O2 is released as a by-
Photosystem II product of this reaction
(PS II)
 Linear Electron Flow 6. Each electron “falls”
down an electron
transport chain from
Primary
acceptor
4
the primary electron
acceptor of PS II to
Pq
2
H2O e−
Cytochrome
PS I
2 H+
+ complex
1/ O2 3
2

7. Energy released by the

e− Pc
e−
P680 5
fall drives the creation
1 Light
of a proton gradient
ATP
across the thylakoid
Pigment
membrane
molecules
8. Diffusion of H+
Photosystem II
(PS II) (protons) across the
membrane drives ATP
synthesis
 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 electron transport chain
Primary
Primary 4 acceptor
acceptor
Pq e−
2
H2O e−
2 H+ Cytochrome
+ complex
1/ O2 3
2
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
Primary
Primary 4 acceptor 7
acceptor Fd
Pq e−
2 e− 8
e− e− NADP+
H2O Cytochrome
2 H+ 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)
 Linear Electron Flow
e−
Mechanical
Analogy! e− e−
Mill
makes
ATP NADPH
e−
e−
e−

e−
ATP

Photosystem II Photosystem I
 Cyclic Electron Flow
• Cyclic electron flow uses only photosystem I & produces ATP, but not
NADPH (i.e. purple sulfur bacteria)
• 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
Chemiosmosis: Chloroplasts vs Mitochondria

• Chloroplasts & 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 & mitochondria but also shows similarities
– Mitochondria – protons pumped to intermembrane space & drive
ATP synthesis as they diffuse back into mitochondrial matrix
– Chloroplasts, protons are pumped into thylakoid space & drive
ATP synthesis as they diffuse back into 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+ ]
 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 1/
2
O2
THYLAKOID SPACE +2 H+ 4 H+
(high H+ concentration)
To
Calvin
Cycle

Thylakoid
membrane ATP
synthase
ADP
+ ATP
STROMA
Pi
(low H+ concentration) H+

• ATP & NADPH are produced on side facing stroma, where the
Calvin cycle takes place
• In summary, light reactions generate ATP & increase the potential
energy of electrons by moving them from H2O to NADPH
 Calvin cycle
• The Calvin cycle, like citric acid cycle, regenerates its starting
material after molecules enter & leave cycle
• The cycle builds sugar from smaller molecules by using ATP &
reducing power of electrons carried by NADPH

• Carbon enters cycle as CO2 & leaves as a sugar glyceraldehyde 3-

phosphate (G3P)
• For net synthesis of 1 G3P, cycle must take place 3X, fixing 3
molecules of CO2
• The Calvin cycle has 3 phases
– Carbon fixation (catalyzed by rubisco)
– Reduction
– Regeneration of the CO2 acceptor (RuBP)
 Input
3 (Entering one
CO2 at a time)

Phase 1: Carbon fixation

Rubisco
3 P P
Short-lived
intermediate
3 P P 6 P
Ribulose bisphosphate 3-Phosphoglycerate
(RuBP)
 Input
3 (Entering one
CO2 at a time)

Phase 1: Carbon fixation

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

Calvin
Cycle
6 P P
1,3-Bisphosphoglycerate
6 NADPH

6 NADP+
6 Pi

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

1 P
G3P Glucose and
(a sugar) other organic
Output compounds
 Input
3 (Entering one
CO2 at a time)

Phase 1: Carbon fixation

Rubisco
3 P P
Short-lived
intermediate
3 P 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
G3P Glucose and
(a sugar) other organic
Output compounds
 3 CO2

Carbon fixation

3 × 5C 6 × 3C

Calvin
Cycle
Regeneration of
CO2 acceptor
5 × 3C

Reduction

1 G3P (3C)
 Alternative mechanisms of carbon
fixation

• Dehydration - problem for plants, sometimes requiring

trade-offs with other metabolic processes, especially
photosynthesis

• Hot, dry days - plants close stomata, which conserves

H2O but also limits photosynthesis
– Closing of stomata reduces access to CO2 & causes O2 to
build up
– These conditions favor an apparently wasteful process called
photorespiration
 Photorespiration
• In most plants (C3 plants)
• initial fixation of CO2, via rubisco, forms a three-carbon
compound (3-phosphoglycerate)

• In photorespiration
• rubisco adds O2 instead of CO2 in the Calvin cycle, producing a
two-carbon compound
• Photorespiration consumes O2 & organic fuel & releases CO2
without producing ATP or sugar
• 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 4-carbon compounds in mesophyll
cells

• This step requires 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 4-carbon compounds are exported to bundle-

sheath cells, where they release CO2 that is then used
in the Calvin cycle
 C4 Plants
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
 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, & CO2 is released from

organic acids & used in Calvin cycle
Sugarcane Pineapple
C4 CAM CO2
CO2
1 CO2 incorporated
Mesophyll Organic acid (carbon fixation) Organic acid Night
cell

CO2 CO2
Bundle- 2 CO2 released Day
sheath Calvin Calvin
Cycle to the Calvin Cycle
cell
cycle

Sugar Sugar
(a) Spatial separation of steps (b) Temporal separation of steps
 Importance of Photosynthesis
• 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)