Ray F. Evert • Susan E.

Eichhorn

Raven Biology of Plants

Eighth Edition

CHAPTER 7
Photosynthesis, Light, and Life
Lecture presentation by Dr. Mohammad Brake
Jerash University

© 2013 W. H. Freeman and Company

CHAPTER OUTLINE
Photosynthesis: A Historical Perspective
The Nature of Light
The Role of Pigments
The Reactions of Photosynthesis
The Carbon-Fixation Reactions
Photosynthesis is the route by which virtually all energy enters our biosphere.
Photosynthesis produces yearly a large amount of sugar.
Photosynthesis: A historical perspective
1) Aristotle and other Greeks thought that plants derived all of their food from
the soil.
2) Helmont (Belgian, 350 years ago), offered the first experimental evidence
that soil alone not nourish the plants?
•Grew a small willow tree in earthen ware plot adding only water
•After 5 years, the willow tree increased in weight by 74.4 kilogram, and the
soil had decreased by 57 gram.
•On the basis of the results, Helmont concluded that all the substances of the
plant were produced from the water not from the soil.
3) Toward the end of the eighteenth century, Joseph priestley reported that
the air restored by vegetation?
4) Shortly thereafter, Janingen housz confirmed priestley’s work and showed
that the air was restored only in the presence of sunlight and only the green
parts of plants.
Ingenhouzs (1796) suggested that carbon dioxide is split in photosynthesis to
yield carbon (added to water) and Oxygen which will be released as gas with
an overall reaction for photosynthesis:
CO2 + H2O + light → CH2O + O2
5) Van niel: did not accept the work of ingenhouzs.
His work was concentrated on the activities of different types of
photosynthetic bacteria.
One particular bacterium (purple sulfur bacteria) produces carbohydrate from
carbon during photosynthesis but does not release Oxygen.
The purple sulfur bacteria require hydrogen sulfide for their photosynthetic
activity, and he found that the following reaction take place during
photosynthesis in these bacteria:
CO2 + 2H2S + light → CH2O + H2O + 2S
Niel proposed the following generalized equation for photosynthesis:
CO2 + 2H2A + light → CH2O + H2O + 2A
Where H2A, represents an oxidizable substance such as:
1. Hydrogen sulfide
2. Free hydrogen
3. Water in plants and algae
Niel proposed that water, not CO2, was the source of the Oxygen in
photosynthesis
6) Robin Hill (1937): the isolated chloroplast was able to produce O2 in the
absence of CO2 when exposed to light.
This result supports the work of niel.
Hill reaction: the light-driven release of O2 in the absence of CO2,.
7) Samnel ruben and martin kamen proved that O2 released in
photosynthesis is derived from H2O by using heavy isotope of Oxygen (18O2)
CO2 + 2H2O18 + light → CH2O + H2O + 18O2
Based in the previous results the complete equation of photosynthesis in
plants and algae is:
3CO2 + 6 H2O ++ light → C3H6O3 + 3O2 + 3 H2O
It is important to remember that a carbohydrate result from photosynthesis is
three-carbon sugar not glucose (6C).
8) Blackman (1905) stated that photosynthesis has:
1. Light-dependent stage: increased in rate as temperature is increased
until 30 Co, which means that photosynthesis is controlled by enzymes.
2. Light-independent stage.
The nature of light
Newton (1642-1727) was able to separate light into a spectrum of visible
colors by passing it through a prism.
Newton showed that white light actually ranging from violet at one end and
red at the other.
James Maxwell (1831-1879) demonstrated that light is but a small part of a
vast continuous spectrum of radiation (electromagnetic spectrum).
All the radiations in this spectrum travel in waves
The wavelength: is the distance from the crest of one wave to the crest of the
next.
e.g
Gamma rays: measured in fractions of nanometer and has great energy
(high energy).
Radio waves: measured in kilometers (low energy)
Within the visible light
1- violet has the shortest wavelength with high energy.
2-red: has the longest wavelength with low energy.
Light has the properties of waves and particles:
by 1900 it had become clear that the wave model of light was not adequate,
why?
•light energy can remove electrons from the metal atoms
•each metal has a critical maximum wave length for electron removal (the
light must be of a particular wavelength or shorter for the removal of the
electrons).
•some metals have a critical wavelength in the spectrum of visible light.
What is the problem with the wave model of light?
The wave model predicts that the brighter the light (the stronger) the greater
the force with which the electrons will be removed from the metal.
The removal of electrons from metal depends only on the wavelength of the
light, not on its intensity.
1. A very weak beam of the critical wavelength is effective , while the
stronger beam of larger wavelength is not.
2. The stronger beam increase the number of electrons removed but not the
velocity at which they are removed from the metal.
3. To increase the velocity , shorter wavelength must be used

The particle model of light was proposed by Einstein in 1905

This model stated that the light composed of particles of energy called
photons (quanta of light)
The energy of a photon is inversely proportional to its wavelength (i.e)
longer wavelength, the lower energy (red). Shorter wavelength, the higher
energy (violet)
The two models of light are complementary to each other for complete
description of the phenomenon of light.
The role of pigments:
For light energy to be used by living organism, it must be first absorbed by
pigments.
Some pigments absorb all wavelength of light and so appear black and most
pigments absorb only certain wavelength and transmit the wavelengths they
don’t absorb
e.g:
Chlorophyll: absorb violet, blue and also red and reflect green.
Action spectrum: is the relative effectiveness of different wavelength of light
for a specific light-requiring process (photosynthesis).
Similarities between the absorption spectrum of a pigment and the
action spectrum of a light-requiring process provide evidence that the
pigment is responsible for that particular process
When chlorophyll molecules absorb light, electrons are temporarily boosted
to a higher energy level, called the excited state. As the electrons return to
their lower energy level, the energy released has three possible faiths:
1- The energy is converted to heat or partly to heat, but mostly released as
another less energetic photon (fluorescence).
The wavelength of the emitted light is slightly longer than the absorbed
(some energy lost as heat).
2- The energy (not the electron) may be transferred from the excited
chlorophyll molecule to neighboring chlorophyll molecule (resonance energy
transfer).
3- The light energy electron itself may be transferred to neighbor molecule
that is part of an electron transport chain.
The main photosynthetic pigments are the
chlorophyll, the carotenoids and phycobilins.
1- There are several kinds of chlorophyll: differ in
structure and light absorption.
Chlorophyll a:
-occurs in eukaryotes and cyanobacteria
- Essential for the oxygen-generating
photosynthesis carried out by organisms of these
groups.
-Constitutes 67% of the chlorophyll while
chlorophyll b is the rest (33%)
chlorophyll b:
-occurs in plants, green algae and euglenoid
-has differ absorption spectrum of chlorophyll a
-is an accessory pigment: not involved directly in
photosynthesis. Absorb light and transfer it to
chlorophyll a.
chlorophyll c:occurs is some groups of algae(brown)
d)-bacteriochlorophyll: occurs in photosynthetic bacteria like purple bacteria
(other than cayanobacteria)
chlorobium chlorophyll: occurs in green sulfur bacteria which use H2S
instead of water.
carotenoides and phycoblins: they are accessory pigments, transfer energy
to chlorophyll a like chlorophyll b and c
carotenoides: red, orange or yellow.
Soluble in lipids
Found in all chloroplast and cyanobacteria
Associated with hydrophilic protein and embedded in the thylakoid membrane.
Two types:
1. Carotenes
2. Xanthophylls
Beta carotene found in plants is the principle source of the vitamin A.
In green leaves, the chlorophyll colors mask the color of carotenoids, but in
autumn when the chlorophyll break down the color of carotenoids become
visible.
The carotenoid pigments can help collect light of different wavelengths; their
principal function is that of an antioxidant, preventing oxidative damage to the
chlorophyll molecules by light. Without the carotenoids, there would be no
photosynthesis in the presence of oxygen.
Phycobilins:
Found in cyanobacteria and red algae.
Water soluble
The reactions of photosynthesis:
Photosynthesis reactions are divided into two major processes:
1) Energy transduction reactions or light reactions
Occurs in the thylakoid membrane
ATP + NADPH will formed
Water molecules will split into:
•O2: released as O2 gas.
•H: will be used to reduce NADP to NADPH.
2) Carbon fixation reactions: conversion of CO2 into organic molecule
Occurs in the stroma
ATP and NADPH will be used
Organic molecules (simple sugars) will be formed from CO2 and H atoms from
NADPH
Two photosystems are involved in the light reactions:
Photosystem: a discrete unit of organization of chlorophyll and other pigment
molecules embedded in the thylakoids of chloroplasts and involved with the
light-dependent reactions of photosynthesis.
Each photosystem has 250-400 pigment molecules divided into:
•Antenna complex
•Reaction center
The function of antenna complex is gathering light energy and transfer it to the
reaction center.
The function of reaction center is the conservation of light energy to chemical
energy molecules.
The photosystem composed of:
•Complex of proteins
•Chlorophyll molecules
Different pigments are present in every photosystem, out of these pigments
there is only one pair of special chlorophyll a molecule which convert light
energy to chemical energy.
The pathway of energy is:
Light energy → antenna complex → reaction center → NADPH + ATP
Kinds of photosystem:
1) Photosystem I:
Discovered first
The special chlorophyll a (P700) absorb light at 700 nm
Located in the stroma thylakoids or in the margins or outer portion of the
grana thylakoids
2) Photosystem II:
Discovered later
The special chlorophyll a (P680) absorb light at 680 nm
Located in the grana thylakoids
In the light reactions, electrons flow from water to photosystem II to
photosystem I to NADP+
1) Light arrive incidently to photosystem II
2) Light energy absorbed by molecules of P680 in the reaction center
•Light energy arrived directly to P680 in the reaction center
•Light energy arrived through antenna molecules
3) When P680 molecule is excited, electron will leave the P680 molecule to
primary acceptor molecule.
4) The electrons will be transferred through electron transport chain and ATP
will form.
5) The P680 molecules will replace its electrons by electrons come from the
splitting of water molecule
2H2O → 4 e- + 4H+ + O2
Photolysis: the light-dependent oxidative splitting of water molecules.
Photolysis is catalyzed by water-splitting enzyme complex which is located in
the inside of thylakoid membrane
The proton resulted from the splitting of water will be released inside the
thylakoid membrane space and proton gradient across thylakoid membrane
will formed which is responsible for ATP synthesis in photosynthesis.
Mn, Cl, Ca: are essential atoms for photosynthesis?
The electron transport chain composed of different electron acceptor
molecules.
Photophosphorylation: the process of generating ATP from ADP+P by means
of proton-motive force generated by the thylakoid membrane of the chloroplast
during the light reactions of photosynthesis.
6) Light arrives incidently to photosystem I
7) Light energy absorbed by molecules of P700 in the reaction center
•Light energy arrived directly to P700 in the reaction center
•Light energy arrived through the antenna complex
8) When P700 molecule is excited, electrons will leave the P700 molecule to
primary electron acceptor (Ao).
9) The electrons will transfer through the electron transport chain and NADPH
will form.
10) The P700 molecules will replace its electrons by electrons arrived from
photosystem II.
2 photons must be absorbed by photosystem II and 2 by photosystem I to
produce:
NAPDH
½ O2
From one H2O
This light reaction called noncyclic reaction flow.
The total energy harvested from noncyclic electron flow is:
ATP, NADPH
From one H O molecule
Cyclic photoposphorylation generates only ATP
Photosystem I work only
The electrons turn back to photosystem I through electron transport chain
between photosystem I and photosystem II and ATP will be formed.
Occurs in primitive photosynthetic organisms (bacteria)
Comparison between non cyclic and cyclic pathways:

non cyclic cyclic

energy ATP, NADPH ATP

water Water splitting No water

O2 O2 will released O2 not involved

In plants and algae:

These organisms needs more ATP than NADPH at ratio of 3:2 for Calvin
cycle, so by the cyclic these organisms synthesis only ATP to meet the need
of Calvin cycle.
The carbon-fixation reactions:
The ATP and NADPH (the results of light reactions) will be used to fix and
reduce carbon to synthesize simple sugar in Calvin cycle.
For plants, C is available to photosynthesis in the form of CO2 and reaches
the photosynthetic cells through stomata.
For algae and cyanobacteria, CO2 is found dissolved in the surrounding
water.
In Calvin cycle, CO2 is fixed via a three-carbon pathway.
In many plants species, the reduction of carbon to simple sugar occurs in
the stroma of the chloroplast by means of series of reactions called Calvin
cycle.
The starting and ending compound in Calvin cycle is five-carbon sugar with
two phosphate groups, known as ribulose 1,5-bisphosphate (RuBP).
The Calvin cycle occurs in three stages:
The first stage: carbon fixation
CO2 enters the cycle and combined with RuBP by an enzyme called RuBP
carboxylase/oxygenase or Rubisco.
The result is an unstable six-carbon compound, immediately hydrolyzed to
generate two molecules of 3-phosphoglycerate or 3-phosphglyceric acid
(PGA).
PGA contains 3 carbon atoms and Calvin cycle is also known as the C3
pathway.
The second stage: reduction
PGA is reduced to glyceraldehyde 3-phosphate (G3P)
The third stage: regeneration of acceptor RuBP
Five of the six molecules of G3P are used to regenerate three molecules of
RuBP.
Important notes about Calvin cycle:
1. 3 CO2 and 3 RuBP are used for each Calvin cycle.
2. Each step in Calvin cycle is catalyzed by specific enzyme.
3. Every three turns of Calvin cycle produce one molecule of G3P.
4. The overall equation for the production of one molecule of G3P is
3 CO2 +9 ATP + 6 NADPH + H+→ G3P + 9 ADP + 8Pi + 6 NADP+ + 3H2O
5. G3P is transported from the chloroplast to the cytosol of the cell.
Most carbon is converted to sucrose or starch.
Much of the G3P produced by Calvin cycle is goes in two ways:
•Exported to cytosol, where, through a series of reactions, it is converted to
sucrose
•Remains in the chloroplast and converted to starch, which will be converted
to sucrose at night and will be transported to other parts of the plant.
Photorespiration occurs when Rubisco binds to O2 instead of CO2
The enzyme Rubisco is not absolutely specific for CO2 as a substrate.
Oxygen competes with CO2 at the active site.
Rubisco catalyzes the condensation of O2 with RuBP to form 1 molecule of
3 phosphglycerate (3C) and 1 of phsphoglycolate (2C).
In photorespiration, no carbon is fixed
Photorespiration is a wasteful process, yielding neiter ATP nor NADPH.
When photorespiration occurs?
1) Under today’s atmospheric conditions (21% O2, and 0.036% CO2)→
photorespiration occurs.
2) When a plant is subjected to hot, dry conditions, it must close its stomata
to conserve water. In these conditions CO2 will decrease, while O2 will
accumulate which means more photorespiration.
3) When plants are growing in close proximity to one another :
-concentration of CO2 in the air closet to the leaves may be low due to the
activity of photosynthesis
- Even when stomata are opens, CO2 is little and O2 accumulates more
photorespiration little photosynthesis.
The four- carbon pathway is a solution to photorespiration.
In some plant, the first detectable product of CO2 fixation is not the three-
carbon molecule 3-phosphoglycerate, but rather the four-carbon molecule
oxaloacetate
Plants which fixed C in four-carbon molecule (C4-pathway) are commonly
called C4 plants
C4-pathway:-
1) CO2 is fixed to phosphoenolpyruvate (PEP) in a reaction catalyzed by the
enzyme PEP carboxylase, which is found in the mesophyll cells of C4 plants,
to produce oxaloacetate.
2) Oxaloacetate is then reduced to malate or asparatate in the chloroplast of
mesophyll cells.
3) CO2 used in step one come from the air spaces.
4) Malate moves from the mesophyll cells to bundle-sheath cells surrounding
the vascular bundles of the leaf, where it is decarboxylated to yield CO2 and
pyruvate.
5) CO2 then enters Calvin cycle by reacting with RuBP to form PGA and
pyruvate reenters to the mesophyll cells, where its reacts with ATP to
generate PEP.
The substrate for the enzyme PEP carboxylase is HCO3- not CO2 as Rubisco.
PEP carboxylase is highly specific for HCO3- and not affected by the presence
of O2.
The anatomy of the leaf of C4 plants is different from the leaf of C3 plants.
In C4 plants, the vascular bundle is surrounded by a layer of bundle-sheath
and then by a layer of mesophyll cells.
Maize is an example of C4 plants.
Photosynthesis is generally more efficient in C4 plants than in C3
plants.
Fixation of CO2 in C4 plants has a larger energy cost than in C3 plants
1. In C4 plants→ 5ATP
2. In C3 plants→3ATP
Why C4 plants have evolved such an energetically expensive method of
providing CO2 to Calvin cycle?
High CO2 and low O2 concentration means more photosynthesis and less
photorespiration.
C4 plants have advantages over C3 plants because CO2 fixed by C4 plants
is pumped from the mesophyll cell into bundle-sheath cells which maintain a
high ratio of CO2 to O2.
CO2 liberated from photorespiration moves from the bundle-sheath can be
refixed by C4 pathway.
Compared with C3 plants, C4 plants are superior utilizers of available CO2
(PEP carboxylase is not inhibited by O2)
As a result the photosynthetic rates of C4 grasses, can be two to three time
the rates of C3 grasses under same environmental conditions.
In short, the gain in efficiency from the elimination of photorespiration in C4
plants more than compensates the energetic cost of the C4 pathway.
C4 plants are plants well adapted to high light intensities, high temperature
and dryness.
The optimal temperature range C4 photosynthesis is much higher than
that for C3 photosynthesis.
C4 plant can attain the same photosynthetic rate as C3 plants but with
smaller stomatal openings and less water loss.
All of the plants known to utilize C4 photosynthesis are flowering plants
(angiosperm)3 families of monocot and 16 families of dicots.
No family has been found, however, that contains only C4 species.

Plants having crassulacean acid metabolism can fix CO2 in the dark
CAM plant:-A second photosynthetic adaptation to arid conditions has
evolved in succulent (water–storing) plants.
These plants open their stomata during the night and close them during
day
Closing stomata during the day helps desert plants to conserve water, but
it also prevents CO2 from entering the leaves.
CAM pathway:
1) Night: stomata open
-CO2 is fixed at night when the stomata are open
-starch is broken down to give phosphoenal pyruvate (PEP)
-CO2 hydrated to HCO3-
-HCO3- reacts with (PEP) to form oxaloacetate
-oxaloacetate then is reduced to malate
-malate is pumped into the vacuole and stored there as malic acid.
2) Day: stomata closed:
-malic acid is recovered from the vacuole and decarboxylated,
producing CO2 and pyruvate
CO2 enters Calvin cycle
-pyruvate converted to sugar and starch by reverse glycolysis
Comparison between C3, C4, CAM in consuming water for CO2
fixation:
C3:- 400-500 gram of water for each gram of CO2 gained
C4:250-300 gram of water for each gram of CO2 gained
CAM:50-100 gram of water for each gram of CO2 gained

CAM plants is more widespread than C4 plants

CAM plants has been reported in at least 23 families of flowering plants
CAM plants has been reported in gymnosperns

Each of the carbon fixation mechanisms has its advantages and

disadvantages in nature
C4 plants generally tolerate higher temperature and drier conditions
than C3 plants
C4 plants may not compete successfully at temperature below 25 Co
because C4 plants are more sensitive to cold than C3 plants
CAM plants conserve water by closing their stomata during the day, a
practice that severely reduces their ability to take in and fix CO2
CAM plants grow slowly and compete poorly with C3 and C4 species
under conditions other than extreme aridity.
The End