Photosynthesis

➢ The process by which green plants and other organisms with photosynthetic
pigments can produce their own food by assimilating/getting solar energy
from the sun.
➢ The most important process on earth because all life forms are very
dependent on photosynthesis.
➢ Without plant/producers we can’t survive. We are very dependent on plant
because they are the producers of food.
➢ Exhibited/performed by autotrophic organisms/autotrophs
➢ Auto = “self”, trophos = “feeding”
➢ Photosynthesis are performed by self-feeding organisms (i.e. plants,
cyanobacteria, different species of algae like seaweeds).
➢ We are heterotrophic organisms – fed by autotrophs.
➢ Hetero = “different”

7.1 PHOTOSYNTHETIC ORGANISMS

• Photosynthesis provides food for the biosphere, oxygen for cellular

respiration, and various significant products.
• In flowering plants, photosynthesis takes place within membrane-bounded
chloroplasts, organelles that contain membranes thylakoids surrounded by
a fluid called stroma.

7.2 THE PROCESS OF PHOTOSYNTHESIS

• Photosynthesis has two sets of reactions. During the light reactions, solar
energy is captured by the pigments in thylakoid membranes, and during the
Calvin cycle reactions, carbon dioxide is reduced by enzymes to a
carbohydrate in the stroma.

7.3 PLANTS AS SOLAR ENERGY CONVERTERS

• Plants use solar energy in the visible light range when they carry on
photosynthesis.
• The light reactions, which occur in thylakoid membranes, produce ATP and
NADH.
7.4 CALVIN CYCLE REACTIONS

• The Calvin cycle reactions, which occur in the stroma, use ATP and NADH
from the light reactions to reduce carbon dioxide to a carbohydrate.

7.5 OTHER TYPES OF PHOTOSYNTHESIS

• Plants use C3 or C4 or CAM photosynthesis, which are distinguishable by the

manner in Co2 which is fixed.

The Nature of Light

Light exhibits the particle-wave duality. Sunlight contains particles called:

• Photons – move in a wave-like behavior.

LIGHT AS A WAVE
Wavelength – the distance
between two light waves.

• Measured in
nanometers (nm); 1
nm = 10-9m
• Arranged from shortest
to longest in an electromagnetic spectrum.

Sunlight consists of:

• 52% Infrared
• 44% Visible light
• 4% Ultraviolet
light
➢ Light is a narrow band
of electromagnetic
spectrum of radiation
emitted by sun.
➢ Plant pigments are
able to observe wavelengths of the visible light (400-700 nm).
➢ The higher the wavelengths, the lower the energy (inversely proportional).
➢ Photosynthetic pigments maximally absorb red and blue lights. The green light is
not absorbed, it is reflected hence it’s appearance as color green in the human
eyes.
ABSORPTION AND ACTION SPECTRA
Absorption Spectra - Tells
the wavelengths that is
being absorbed by the
different chlorophyll
molecules.

Action Spectra – tells the

rate of photosynthesis.

7.1 Photosynthetic organisms

Photosynthesis converts solar energy into the chemical energy of carbohydrate.
Photosynthetic organisms, including land plants, algae, and cyanobacteria, are called
autotrophs because they produce their own food. Photosynthesis produces an
enormous amount of carbohydrate. So much that, if it were instantly converted to
coal and the coal were loaded into standard railroad cars (each car holding about 50
tons), the photosynthesizers of the biosphere would fill more than 100 cars per
second with coal. No wonder photosynthetic organism are able to sustain
themselves and all other living things on Earth. With few exceptions, it is possible to
trace any food chain back to plants and algae. In other words, producers, which have
the ability to synthesize carbohydrates, feed not only on themselves but also
consumers, which must take in preformed organic molecules. collectively,
consumers are called heterotrophs. Both autotrophs and heterotrophs use organic
molecules produced by photosynthesis as a source of building blocks for growth and
repair and as a source of chemical energy for cellular work.

Photosynthesizers also produce copious amounts of oxygen as a by-product. Oxygen,

which is required by organisms when they carry on cellular respiration, rises high into
the atmosphere, where it forms an ozone shield that filters out ultraviolet radiation
and makes terrestrial life possible.

Our analogy about photosynthetic products becoming coal is apt because the bodies
of many ancient plants did become the coal we burn today, usually to produce
electricity. Coal formation happened several hundred million years ago, and that is
why coal is called fossil fuel. Today’s trees are also commonly used as fuel. Then too,
the fermentation of plant materials produces ethanol, which can be used directly to
fuel automobiles or as a gasoline incentive.
The products of photosynthesis are critical to human-kind in a number of other ways.
They serve as a source of building materials, fabrics, paper, and pharmaceuticals.
And while we are thanking green plants for their services, let’s not forget the simple
beauty of a magnolia blossom or the majesty of the Earth’s forest.

FLOWERING PLANTS AS PHOTOSYNTHESIZERS

Photosynthesis takes place in the green portions of plants. The leaves of a flowering
plant contain mesophyll tissue in which cells are specialized for photosynthesis. The
raw materials for photosynthesis are water and carbon dioxide. The roots of a plant
absorb water, which then moves in vascular tissue (xylem) up the stem to a leaf by
way of the leaf veins. Carbon dioxide in the air enters a leaf through small openings
called stomata (sing., stoma). After entering a leaf, carbon dioxide and water diffuse
into chloroplasts [Gk, chloros, green, and plastos, formed, molded], the organelles
that carry on photosynthesis.

A double-membrane surrounds a chloroplast, and its semifluid interior called stroma

[Gk, thylakos, sack, and eides, like, resembling], which in some places are stacked to
form grana (sing., granum), so called because they looked like piles of seeds to early
microscopists. The space of each thylakoid is within a chloroplast, thereby forming
an inner compartment within chloroplasts called the thylakoid space.
The thylakoid membrane contains chlorophyll and other pigments that are capable
of absorbing solar energy, the type of energy that drives photosynthesis. The stroma
contains an enzyme-rich solution where carbon dioxide is first attached to an organic
compound and is then reduced to a carbohydrate.

• Leaves - major photosynthetic organ

• In herbacious plants, any green part of the body can be utilized in
photosynthesis (i.e. green stems, green branches) .
• Leaves are flat, broad, and very, very thin so that it can maximally absorb
wavelgnths from the sun.
• The leaves are composed of an upper and lower epidermis. The lower
epidermis is said to be provided with a lot of stomates.
• Stomates – openings; very important forthe passage and exchange of gases.
Serves as an entry for carbon dioxide and release of oxygen.
• Mesophyll – major photosynthetic are of the leaf. Made up of parenchyma
cells specifically chlorenchyma.
• Chlorenchyma - parenchyma cells with chloroplast.
In Dicots, the mesophyll is divded into 2 layers:

1. Palisade mesophyll – a compact arrangement of the chlorenchyma.

2. Spongey mesophyll – composed of intracellular spaces utilized for the
exchange of gases.

The chloroplast is a tripled-layer organelle.

• The third layer is the thylakoid membrane.

• Thylakoid membrane -> organized = stacks also called the granum.
• Thylakoids are arranged in stacks to increase the surface area so that more
chemical energy can be produced from solar energy.

Therefore, it is proper to associate the absorption of solar energy with the thylakoid
membranes making up the grana and to associate the reduction of carbon dioxide to
a carbohydrate with the stroma of a chloroplast.

Human beings, and indeed nearly all organisms, release carbon dioxide into the air.
This is some of the same carbon dioxide that enters a leaf through the stoma and is
converted to carbohydrate. Carbohydrate, in the form of glucose, is the chief source
of chemical energy for most organisms.

7.2 The process of photosynthesis

• In the presence of light and chlorophyll, carbon dioxide and water are converted
to form sugar and there is a release of oxygen as a by-product.
• Photosynthetic reaction. In the presence of light and chlorophyll (solar energy),
carbon dioxide is reduced to become carbohydrate, and water is oxidized (split)
to become oxygen. The hydrogen atoms especially the electrons are
incorporatedto the forming carbohydrate.
REDUCTION-OXIDATION (REDOX) REACTIONS

• When two hydrogen atoms are added to

pyruvic acid, it becomes reduced to lactic
acid.
• Reduction – addition of electrons, making a
compound more negative.
• Reduced – when a compound accepts
electrons and become more negative.

• NADP+ - Nicotinamide Adenine Dinucleotide Phosphate – A coenzyme

• Coenzyme – electron-carriers from one
molecule to another. Help enzymes to
function very well. Enzymes are proteins
and coenzymes is an organic non-
protein.
• The electron neutralizes the positive sign in NADP+ to be NADPH
• In a reaction, reduction is always paired with oxidation and vice versa.
• Redox reaction is always coupled, it something lost an electron, then something
must have gained one.

• Pyruvic acid is oxidized to Acetyl group,

it lost an electron and a hydrogen atom.
• The NAD+ reduced, it gained an electron
and a hydrogen atom.
•

This overall equation can be used to represent the process of photosynthesis:

In this equation, (CH2O) represents carbohydrate. Of the equation were multiplied by

six, the carbohydrate would be C6H12O6, or glucose.
The overall equation shows that photosynthesis involves oxidation-reduction (redox)
and the movement of electrons from one molecule to another. Recall that oxidation
is the loss of electrons, and reductions is the gain of electrons. In living things, the
electrons are very often accompanied by hydrogen ions so that oxidation is the loss if
hydrogen atoms (H++e-), and reduction is the gain of hydrogen atoms. This simplified
rewrite of the above equation makes it clear that carbon dioxide has been reduced,
and water has been oxidized:

It takes hydrogen atoms and also energy to reduce carbon dioxide. From our study of
energy and enzymes, you expect that solar energy will not be used directly during
photosynthesis, and instead, it will be converted to ATP molecules. ATP is the energy
currency of cells and, when cells need something, they spend ATP. In this case, solar
energy will be used to generate the ATP needed to reduce carbon dioxide to a
carbohydrate. Of course, we always want to keep in mind that this carbohydrate
represents the food produced by land plants, algae, and cyanobacteria that feeds the
biosphere.

Electrons needed to reduce carbon dioxide will be carried by a coenzyme. NADP+ is

the coenzyme of oxidation-reduction (redox coenzyme) active during
photosynthesis. When NADP+ is reduced, it has accepted two electrons and one
hydrogen atom, and when it is oxidizes, it gives up its electrons:

What molecule supplies the

electrons that reduce NADPH
during photosynthesis? Put a sprig
of Elodea in a beaker, and supply it
with light, and you will observe a
bubbling (Fig. 7.3). The bubbling
occurs because the plant is
releasing oxygen as it
photosynthesizes. A very famous
experiment performed by C. B. van
Niel of Stanford University found
that the oxygen given off by
photosynthesizers comes from water. This was the first step toward discovering that
water splits during photosynthesis. When water splits, oxygen is released and the
hydrogen atoms H +e) are taken up by NADPH. Later, NADH reduces carbon dioxide
to a carbohydrate.

Van Niel performed two separate experiments. When an isotope of oxygen, O, was a
part of water, the O, given off by the plant contain "O. When "O was a part of carbon
dioxide supplied to a plant, the O, given off by a plant did not contain the 10. Why
not?

TWO SETS OF REACTIONS

Many investigators have contributed to our understanding of the overall equation of

photosynthesis and to our current realization that photosynthesis consists of two
separate sets of reactions. F. E. Blackman was the first to suggest, in 1905, that
enzymes must be involved in the reduction of carbon dioxide to a carbohydrate and
that, therefore, the process must consist of two separate sets of reactions. We will
call the two sets of reactions the light reactions and the Calvin cycle reactions.

Light Reactions
The light reactions are so named because they only occur when solar energy is
available (during daylight hours). The overall equation for photosynthesis gives no
hint that the green pigment chlorophyll, present in thylakoid membranes, is largely
responsible for absorbing the solar energy that drives photosynthesis. During the

light reac- tions, solar energy energizes electrons that move down an electron
transport chain (see Figure 6.12). As the electrons move down the chain, energy
released and captured for the production of ATP molecules. Energized electrons are
also taken up by NADP*, which becomes NADPH. This equation can be used to
summarize the light reactions be- cause, during the light reactions, solar energy is
converted to chemical energy

Calvin Cycle Reactions

The Calvin cycle reactions are named for Melvin Calvin, who received a Nobel Prize
for discovering the enzymatic reactions that reduce carbon dioxide to a
carbohydrate in the stroma of chloroplast (Fig. 7.5). The enzymes that are able to
speed the reduction of carbon dioxide during both day and night are located in the
semifluid substance of the stroma.
During the Calvin cycle reactions, CO, is taken up and then reduced to a
carbohydrate that can later be converted to glucose. This equation can be used to
summarize the Ca vin cycle reactions because, during these reactions, the ATP and
NADPH formed during the light reactions are used to reduce carbon dioxide:

Summary
Figure 7.4 can be used to summarize our discussion so far. This figure shows that
during the light reactions, (1) solar energy is absorbed, (2) water is split so that
oxygen is released, and (3) ATP and NADPH are produced.
 During the Calvin cycle reactions, (1) CO2 is absorbed and (2) reduced to a
carbohydrate (CH2O) by utilizing ATP and NADPH from the light reactions (see
bottom set of red arrows). The top set of red arrows takes ADP + and
NADP back to light reactions, where they become ATP and NADPH once
more so that carbohydrate production can continue.

7.3 Plants as solar energy converters

Solar energy can be described in terms of its wavelength and its energy content.
Figure 7.6a lists the different types of radiant energy from the shortest wavelength,
gamma rays, to the longest, radio waves. Most of the radiation reaching the Earth is
within the visible-light range. Higher- energy wavelengths are screened out by the
ozone layer in the atmosphere, and lower-energy wavelengths are screened out by
water vapor and carbon dioxide before they reach the Earth's surface. The
conclusion is, then, that organic molecules and processes within organisms, such as
vision and photosynthesis, are chemically adapted to the radiation that is most
prevalent in the environment visible light (fig. 7.6a).

Pigment molecules absorb wavelengths of light. Most pigments absorb only some
wavelengths; they reflect or transmit the other wavelengths. The pigments found in
chloroplasts are capable of absorbing various portions of visible light. This is called
their absorption spectrum. Photosynthetic organisms differ by the type of
chlorophyll they contain. In plants, chlorophyll a and chlorophyll b play prominent
roles in photosynthesis. Carotenoids play an accessory role. Both chlorophylls a and
b absorb violet, blue, and red light better than the light of other colors. Because
green light is transmitted and reflected by chlorophyll, Plant leaves appear 8reen to
us. 1he carotenoids, which are shades or yellow and orange, are able to absorb light
in the violet-blue-green range. These pigments become noticeable in the tall when
chlorophyll breaks down.

How do you determine the absorption spectrum or pigments? To identify the

absorption spectrum of a particular pigment, a purified sample is exposed to
different wavelengths of light inside an instrument called a spectrophotometer. A
spectrophotometer measures the amount of light that passes through the sample,
and from this it is possible to calculate how much was absorbed. The amount of light
absorbed at each wavelength is plotted on a graph, and the result is a record of the
pigment's absorption spectrum (Fig. 7.6b).

THE NATURE OF A CHLOROPHYLL MOLECULE

• A chlorophyll molecule is capable of absorbing a photon of light that contains
energy in the form of quantum.
• When a photon of light is absorbed by a chlorophyll molecule, that quantum of
energy is utilized to excite an electron.
• The photo-excited electron will
not stabilize and can easily
become unstable or could go back
to the unexcited state, and the
energy that originally came from
the quantum can be wasted in the
form of heat or fluorescence
(light).
• In photosynthesis this should not
be the case. In order to solve this
problem in plants, the chlorophyll
molecules are arranged to form a
photosystem.
The photosystem have 2 important components:

1. Antenna complex – antenna chlorophyll molecules absorb incoming photons

and funnel the excitation energy to the reaction center.
o Chlorophyll molecules – funnel/transfer the energy into a reaction
center.
o Antenna – receives and transmits wavelengths/signals to the
television/radio.
2. Reaction center – photochemical oxidation-reduction occurs which stabilize
electrons. Includes 2 special chlorophyll a molecules and other carriers; the 2
chlorophyll a maximal absorption is 700 nm (PS 1) and 680 nm for PS II.
o Very important in a redox reaction.
o The redox-reaction stabilizes the electron until such time that the
photo-excited electron is accepted by a primary electron acceptor so
that it can go along the process.
• The solar energy is not wasted anymore because solar energy is channelled until
such time that the energy is said to be stabilized and used for photosynthesis.

Light Reactions

The light reactions utilize two photosystems, called photo- system1(PS) and
photosystem ll (PS). The photosystems are named for the order in which they were
discovered, not for the order in which they occur in the thylakoid membrane or
participate in the photosynthetic process. A photosystem consists of a pigment
complex (molecules of chlorophyll a, chlorophyll b, and the carotenoids) and
electron acceptor molecules within the thylakoid membrane. The pigment complex
serves as an "antenna" for gathering solar energy.
PHOTOSYSTEMS

The 2 photosystems have different reaction centers.

PS I

• Reaction center of P700 (wacelengths) – maximally absorb 700 nm of light

PS II

• Reaction center of P680 (wavelengths) – maximally absorb 680 nm of light

 Overview of Photosynthesis

• Has 2 stages: (1) light reactions, and (2) dark reaction/calvin cycle.
• In the presence of light and chlorophyll, in the thylakoid membrane
• Thylakoid membrane – made up of chlorophyll molecules which can absorb
photons of light. These photons contain energy in the form of quantum.
Contains pigments necessary for absorbing solar energy.
• When light energy is harvested, that solar energy is now converted to become 2
forms of chemical energy: ATP and NADPH.
• Utilizes water as a raw material. Water is now oxidized to become oxygen.
Water is the source of electrons.
• After producing ATP and NADPH, these two forms of energy are now utilized in
the Calvin Cycle.
• The calvin cycle happens in the stroma of the chloroplast.
• The calvin cycle is the production of carbohydrates so that the raw material,
carbon dioxide, is now reduced to form carbohydrate.
• After utilizing the solar/chemical energy, in making the carbohydrate, there is a
regeneration of ADP+ and NADP+ - acceptors, these go back to the light reactions
so that the cycle can repeat.
• Solar energy is harvested by the chlorophyll molecules, and that solar energy is
converted to 2 forms of chemical energy: (1) ATP, and (2) NADPH. These
energies will be utilized by the Calvin cycle in reducing carbon to become
carbohydrate.
STAGE 1: LIGHT-DEPENDENT REACTIONS

Electron Pathway 1: Non-Cyclic (Linear) Photophosphorylation

➢ Photophosphorylation – the production of ATP utilizing the energy from the sun.

➢ The electrons are transferred from: Water -> PS II -> PS I, the electrons are
trapped in the NADPH. There is a formation of two chemical energies: ATP and
NADPH.
➢ Products: ATP + NADPH
➢ Happens in the thylakoids

PHOTOSYSTEM I

• When a photon of light strikes a chlorophyll molecule, the energy is transferred

to a series of antenna complex chlorophyll molecules until the energy reaches
the reaction center and the energy is now used to photoexcite the electrons.
• The photoexcited electrons are accepted by a primary electron acceptor.
• The purpose of stabilizing the electrons so that they can be transferred to a
series of reaction in order to produce ATP.
• From the primary electron acceptor, the electrons travel along an electron
transport chain or ETC

Thylakoid’s ETC vs. Mitochondrion’s ETC

ETC – Produces ATP

Mitochondrion

• Called an ETC because it is made up of a series of electron transporters.

• The reduced NAD transfers its electrons to the first system/first electron
acceptor -> NADH is oxidized to become NAD and the first electron acceptor
becomes reduced.
• When the first electron acceptor passes the electrons to the second electron
acceptor, the second electron acceptor becomes reduced and the first electron
acceptor becomes oxidized, and so on, and so forth.
Series of electron transporters until the electrons reaches the Ferredoxin

chemiosmosis

• In cellular respiration, the electron acceptors/electron transporters are

organized to form complexes.
• The energy carried by the electrons’ purpose is utilized by different proteins to
pump hydrogen ions. (Mitochondrion: from the matrix to the intermembrane
space).
• Hydrogen ions are accumulated so that when these hydrogen ions diffuse back
into the interior of the mitochondrion, the kinetic energy provided by the
hydrogen ions is now utilized to power the production of ATP molecules. ->
protein = ATP Synthase = also the protein in the thylakoid membrane.

Thylakoid

• The energy from the electron is also utilized to accumulate hydrogen ions going
into the thylakoid space.
• When the hydrogen ions diffuse, going to the stroma, energy is now utilized to
form ATP -> Chemiosmosis
• Chemiosmosis – production of ATP utilizing energy from chemical ions.
• In phosphorylation produces ATP utilizing energy from the sun, while in
chemiosmosis, energy is already converted.

• Electrons will pass through ETC until they are received by Ferredoxin (Fd), the
electrons are now accepted by NADP+ (very important coenzyme).
• NADP+ is not reduced to become NADPH by the enzyme NADP+ reductase.
• NADP+ reductase gets the electrons from Ferredoxin and transfers the electron
to the 2 NADP+, and the NADP is reduced to become NADPH
• NADP+ - Final electron acceptor; reduced to become NADPH
• Problem: when the electrons are trapped by NADPH, the PS I will be rid of
electrons, there will be electron holes because the electrons are captured by the
NADP+ to become NADPH. Since all the electrons are captured, where would the
PS I get its electrons from? -> PS II then Water

PHOTOSYSTEM II

• PS II – Same thing happens. When a photon of light strikes a chlorophyll

molecule of the antenna complex, the energy is captured/funnelled until it
reaches the reaction center where the electrons are photoexcited and stabilize
until the electron is accepted by the primary electron acceptor.
• Same however, the Plastoquinone (Pq) comes first.
• Primary acceptor -> Pq -> Cytochrome Complex (ETC) -> Plastocyanin (Pc)
• As the electrons pass the ETC, there is a production of ATP; the electrons go to
plastocyanin and then goes back to the reaction center of PS I.
• Now, both PS I and PS II have no electrons, the PS II will get electrons from
water, which is why it is a raw material during the light reactions in
photosynthesis.
• Water is oxidized to become oxygen because the hydrogen ions split to provide
electrons.
• Water molecules = hydrogen ions + electrons + oxygen > HILL REACTION
• Water molecules is split into hydrogen ions and electrons and also oxygen, so
that oxygen is released.
• Hill Reaction – in honor of Sir Robert Hill who studied the process of the rate of
photosynthesis of an isolated chloroplast molecules.
Electron Pathway 2: Cyclic Photophosphorylation

• Performed by PS I
• If there is a very high ratio of NADPH to NADP+, cyclic photophosphorylation
happens.
• All the NADP has received electrons and became NADPH, meaning no more
NADP can receive electrons.
• From the Ferredoxin, the electrons would have to go back to the Cytochrome
Complex.
• As the electrons are passed through a series of electron transporters again,
there will be a production of ATP.
• Electrons go back to Plastocyanin, to the reaction center and so on, and so forth.
• The cycle repeats.
• Product: ATP

During the light reactions, electrons usually follow a noncyclic pathway that begins
with photosystem II (Fig.7.7). The pigment complex absorbs solar energy, which is
then passed from one pigment to the other until it is concentrated in a particular
pair of chlorophyll a molecules, called the reaction center. Electrons (e -) in the
reaction center become so energized that they escape from the reaction center and
move to nearby electron acceptor molecules.
PS II would disintegrate without replacement electrons, and these are removed from
water, which splits, re- leasing oxygen to the atmosphere. Notice that with the loss
of electrons, water has been oxidized, and that indeed, the oxygen released during
photosynthesis does come from water. Many organisms, including plants and even
our- selves, use this oxygen within their mitochondria. The hydrogen ions (H +) stay in
the thylakoid space and contribute to the formation of a hydrogen ion gradient.

An electron acceptor sends energized electrons, received from the reaction center,
down an electron trans- port chain (ETC), a series of carriers that pass electrons from
one to the other (see Fig. 6.13). As the electrons pass from one carrier to the next,
energy is captured and stored in the form of a hydrogen ion (H) gradient. When
these hydrogen ions flow down their electrochemical gradient through ATP synthase
complexes, ATP production occurs (see page 124). Notice that this ATP will be used
by the Calvin cycle reactions in the stroma to reduce carbon dioxide to a
carbohydrate.

When the PS I pigment complex absorbs solar energy, energized electrons leave its
reaction center and are captured by electron acceptors. (Low-energy electrons from
the electron transport chain adjacent to PS II replace those lost by PS I.) The electron
acceptors in PS I pass their electrons to NADPT molecules. Each one accepts two
electrons and an H to become a reduced form of the molecule, that is, NADPH. This
NADPH will be used by the Calvin cycle reactions in the stroma to reduce carbon
dioxide to a carbohydrate.
The Organization of the Thylakoid Membrane

As we have discussed, the following molecular complexes are in the thylakoid

membrane (Fig. 7.8):

PS II, which consists of a pigment complex and electron-acceptor molecules,

receives electrons from water asS water spnts, releasing Oxy8en.

The electron transport chain (ETC), consisting of Pq (plastoquinone) and cytochrome

complexes, carries electrons from PS to PSI. Pq also pumps H + from the stroma into
the thylakoid space.

PS I, which also consists of a pigment complex and electron-acceptor molecules, is

adjacent to NADP reductase, which reduces NADP+ to NADPH.

The ATP synthase complex has a channel and a protruding ATP synthase, an enzyme
that joins ADP +
ATP Production

The thylakoid space acts as a reservoir for hydrogen ions (H+). First, each time water
is oxidized, two H+ remain in the thylakoid space. Second, as the electrons move
from carrier to carrier along the electron transport chain, the electrons give up
energy, which is used to pump H+ from the stroma into the thylakoid space.
Therefore, there are more H+ the thylakoid space than in the stroma. The flow of H+
(often referred to as protons in this context) from high to low concentration provides
kinetic energy that allows an ATP synthase complex enzyme to enzymatically
produce ATP from ADP + . This method of producing ATP is called chemiosmosis
because ATP production is tied to the establishment of an H gradient (see Fig.
6.13).

Tropical rain forest destruction and global warming

Algore, former presidential candidate, won the 2007 Nobel Peace Prize tor raising
public awareness concerning global warming. The Nobel Committee said that "global
warming could induce large-scale migrations and lead to greater competition for the
Earth's resources. As such, it may increase the danger of violent conflicts and wars,
within and between countries." Global warming refers to an expected rise in the
average global temperature during the twenty-first century due to the introduction
of certain gases into the atmosphere. For at least a thousand years prior to l850,
atmospheric carbon dioxide (CO2) levels remained fairly constant at 0.028%. Since
the 1850s, when industrialization began, the amount of CO 2, in the atmosphere has
increased to 0.038% (Fig. 7Ao).

Role of Carbon Dioxide

In much the same way as the panes of a green- house, CO2 and other gases in our
atmosphere trap radiant heat from the sun. Therefore, these gases are called
greenhouse gases. Without any greenhouse gases, the Earth's temperature would be
about 33°C cooler than it is now. Likewise, increasing the concentration of
greenhouse gases is predicted to cause global warming

Certainly, the burning of fossil fuels adds CO2 to the atmosphere. But another factor
that contributes to an increase in atmospheric CO, is tropical rain forest destruction.
Role of Tropical Rain Forests

Between 10 and 30 mllion hectares of rain forests are lost every year to ranching,
loggng. mining. and otherwise developing areas of the forest for human needs. The
clearing of forests often involves burning them (Fig 7Ab). Each year, deforestation in
tropical rain forests accounts for 20-30% of all Co, in the atmosphere. The
consequence of burning forests is greater trouble tor global warming because
burning a forest adds CO2 to the atmosphere and, at the same time removes trees
that would ordinarily absorb CO2

The Argument for Preserving Forests

The process of photosynthesis and also the oceans act as a sink for CO2. Despite
their reduction in size from an original 14% to 6% of land surface today, tropical rain
forests make a substantial contribution to global CO2, removal. Taking into account
all ecosystems, marine and terrestrial, photosynthesis produces organic matter that
is 300 to 600 times the mass o people currently on Earth this year. Tropical rain
forests contribute greatly to the uptake of CO2 and the productivity of
photosynthesis because they are the most efficient of all terrestrial ecosystems.

Tropical rain forests occur near the equator. They can exist wherever temperatures
are above 26 C and rainfall is heavy (from 100-200 cm) and regular: Huge trees with
buttressed trunks and broad, undivided, dark-green leaves predominate. Nearly all
land plants in a tropical rain forest are woody, and woody vines are also abundant.

It might be hypothesized that an increased amount of CO 2 in the atmosphere will

cause photosynthesis to increase in the remaining portion of the forest. To study this
possibility. investigators measured at atmospheric Co levels, daily temperature
levels, and tree girth in La Selva, Costa Rica, for 6 years. The data collected
demonstrated relatively lower forest productivity at higher temperatures. These
findings suggest that, as temperatures rise, tropical rain forests may add to ongoing
atmospheric accumulation and accelerated global warming rather than the reverse.
All the more reason to slow global warming and preserve forests.

Some countries have programs to combat the problem of deforestation. In the mid-
1970s, Costa Rica established a system of national parks and reserves to protect 12%
of the country's land area from degradation. The current Costa Rican government
wants to expand the goal by increasing protected areas to 25% in the near future.
Similar efforts in other countries may help slow the ever-increasing threat of global
warming.
7.4 Calvin Cycle Reactions
The Calvin cycle reactions occur after the light reactions. The Calvin cycle is a series
of reactions that produce carbohydrate before returning to the starting point once
more (Fig. 7.9). The cycle is named tor Melvin Calvin, who, with colleagues, used the
radioactive isotope C as a tracer to discover the reactions making up the cycle.

This series of reactions uses carbon dioxide from the atmosphere to produce
carbohydrate. How does carbon di- oxide get into the atmosphere? We and most
other organ systems take in oxygen from the atmosphere and release carbon dioxide
to the atmosphere. The Calvin cycle includes (1) carbon dioxide fixation, (2) carbon
dioxide reduction, and (3) regeneration of RuBP (ribulose-1,5-bisphosphate).

STAGE 1: DARK REACTIONS/CALVIN-BENSON CYCLE/CARBON FIXATION CYCLE

• Light-independent reactions
• Happens in the stroma of the chloroplast
• Melvin Calvin and Andrew Benson -> discovered the pathway
• The raw material during here is carbon dioxide, a source of carbon.
• Carbon is reduced to become sugar.
• The carbon from carbon dioxide is fixed/added to RuBP (Ribulose Bisphosphate)
• C3 Cycle – the true product in photosynthesis is a 3-carbon sugar ->
Glyceraldehyde 3-phosphate (G3P) or phosphoglyceraldehyde (PGAL)
• Exhibited/performed by 80% of plants on earth -> C3 Plants
• Photosynthetic Carbon Reduction (PCR) Cycle – carbon is reduced to become
carbohydrate.

• Correspond to the synthesis part in photosynthesis

• 3 Phases: (1) Carbon Fixation, (2) Reduction, and (3) Regeneration of the CO2
Acceptor (RuBP)

Fixation of carbon dioxide

Carbon dioxide (CO2) fixation is the first step of the Calvin cycle. During this reaction,
carbon dioxide from the atmosphere is attached to RuBP, a 5-carbon molecule. The
result is one 6-carbon molecule, which splits into two 3-carbon molecules

The enzyme that speeds this reaction, called RuBP carboxylase, is a protein that
makes up about 20-50% of the protein content in chloroplasts. The reason for its
abundance may be that it 1s unusually slow (it processes only a few molecules of
substrate per second compared to thousands per second for a typical enzyme), and
so there has to be a lot of it to keep the Calvin cycle going.

• Carbon form CO2 is now fixed/added/incorporated to the starting material ->

RuBP by an enzyme Rubisco (Ribulose-1, 5-bisphosphate carboxylase/oxygenase
• Rubisco – the most abundant protein on earth. The carboxylase part is active in
incorporating carbon from CO2 into RuBP, hypothetical 6-carbon sugar is
produced and this immediately splits into 2 molecules of phosphoglycerate
containing 3 carbons each.
Reduction of Carbon Dioxide

The first 3-carbon molecule in the Calvin cycle is called 3PG (3-phosphoglycerate).
Each of two 3PG molecules undergoes reduction to G3P in two steps:

This is the sequence of reactions that uses some ATP and NADPH from the light
reactions. This sequence signifies the reduction of carbon dioxide to a carbohydrate
because R--CO2 has become R--CH2O. Energy and electrons are needed for this
reduction reaction, and these are supplied by ATP and NADPH.

• Reduction of carbon to become carbohydrate.

• The phosphoglycerate is now phosphorylated, it is now energized (given a
phosphate group).
• NADPH gives its electrons.
• 1, 3-bisphosphoglycerate is reduced
• NADP collects electrons from PS I, PS II, and water, and the electrons are now
transferred to the forming carbohydrate.
• 1, 3-bisphosphoglycerate is reduced to become G3P
• And NADPH is now oxidized to become NADP
• The ADP and NADP go back to the light reactions to form ATP and NADPH
• G3P/PGAL – True product of photosynthesis (C3 Cycle)

Regeneration of RuBP
 Notice that the Calvin cycle reactions in Figure 7.9 are multiplied by three because it
takes three turns of the Calvin cycle to allow one G3P to exit. Why? Because, for
every three turns of the Calvin cycle, five molecules of G3P are used to re-form three
molecules of RuBP and the cycle continues. Notice that 5 x 3 (carbons in G3P) = 3 x5
(carbons in RuBP):

This reaction also uses some of

the ATP produced by the light
reactions.

• Other G3P undergo a series

of reduction in the 3rd
phase (regeneration of CO2
acceptor – RuBP) until there is a production of the carbon acceptor so that the
carbon cycle can repeat.
The Importance of the Calvin Cycle

G3P (glyceraldehyde-3-
phosphate) is the product of the
Calvin cycle that can be
converted to other molecules a
plant needs. Notice that ghucose
phosphate is among the organic
molecules that result from G3P
metabolism (Fig. 7.10). This is of
interest to us because glucose is
the molecule that plants and
animals most often metabolize to
produce the ATP molecules they
re- quire for their energy needs.

Glucose phosphate can be

combined with fructose (and the
phosphate removed) to form
sucrose, the molecule that plants
use to transport carbohydrates
from one part of the plant to the
other.

Glucose phosphate is also the

starting point for the synthesis of starch and cellulose. Starch is the storage form of
glucose. Some starch is stored in chloroplasts, but most starch is stored in
amyloplasts in roots. Cellulose is a structural component of plant cell walls and
becomes fiber in our diet because we are unable to digest it.

A plant can use the hydrocarbon skeleton of G3P to form fatty acids and glycerol,
which are combined in plant oils. We are all familiar with corn oil, sunflower oil, or
olive oil used in cooking. Also, when nitrogen is added to the hydrocarbon skeleton
derived from G3P, amino acids are formed.

Photorespiration
• Occurs on hot (30-45 C),
dry, bright days
• For plants not to be
dehydrated, they close
their stomata in order
to prevent too much
transpiration –
evaporation of water
from the surface of the plants.
• When stomates are open, lots of water are loss are evaporate.
• When the stomates are closed, carbon dioxide cannot enter anymore and
oxygen will increase (since it is a by-product of photosynthesis).
• When there is more oxygen compared to carbon dioxide, oxygen will be fixed by
Rubisco and CO2 will be released.
• Instead of carbon dioxide being used in the calvin cycle, carbon dioxide is
released.
• RuBP + O2 instead of CO2

Competes with photosynthesis

• When the level of carbon

dioxide is very low compared
to the level of oxygen, the
enzyme ribulose-1, 5-
bisphosphate
carboxylase/oxygenase fixes
oxygen rather than carbon
dioixde.
• Carbon dioxide is released
instead of being utilized to
produce sugar, but ATP and
NADPH are still being used up
in order to regenerate RuBP.
• No sugar is produced, because CO2 is released and wasted, but ATP and NADPH
are still being used up to regenerate RuBP.
• Super less productive

Photorespiration and the C2 glycolate/PCO Cycle

• If there is an increase in the amount of oxygen, compared to carbon dioxide,
Rubisco fixes oxygen to RuBP to become 2-phosphoglyolate which is converted
to glycolate, and glycolate contains only 2 carbon -> C2 glycolate cycle.
• From the chloroplast, glycolate enters the peroxisome (reduces H2O2/hydrogen
peroxide).
• Glycolate oxidizes to become Glyoxylate -> Glycine
• Glycine enters the mitochondrion
• In the mitochondrion, glycine is now oxidized, and NAD is reduced, and carbon
dioxide is released -> photorespiration
• From the mitochondrion, Serine returns to the peroxisome, becomes Glycerate,
enters the chloroplast, regeneration of carbon dioxide acceptor (RuBP), ATP is
utilized until there is regeneration of RUBP
• Experienced by 80% of plants in the world
• The remaining 20%, evolved in order to compensate for this condition.
7.5 Other types of photosynthesis
The majority of land plants such as azaleas, maples, and tulips carry on
photosynthesis as described and are called C3 plants (Fig. 7.11a). C3 plants use the
enzyme RuBP carboxylase to fix CO2 to RuBP in mesophyll cells. The first detected
molecule following fixation is the 3- carbon molecule 3PG:

As shown in figure 7.2, leaves have small openings called stomata through which
water can leave and carbon dioxide (CO2) can enter If the weather is hot and dry, the
stomata close, conserving water. (Water loss might cause the plant to wilt and die.)
Now the concentration of CO2 decreases in leaves, while O2, a by-product of
photosynthesis, increases. When O2, rises in C plants, RuBP carboxylase combines it
with RuBP instead of CO2. The result is one molecule of 3PG and the eventual release
of CO2. This is called photorespiration because in the presence of light (photo),
oxygen is taken up and CO2 is released (respiration).

An adaptation called C4 photosynthesis enables some plants to avoid

photorespiration.

Because of photorespiration, plants have special adaptations to limit the effect of

photorespiration:

1. C4 plants
2. CAM Plants

C4 Photosynthesis

Objective: Concentrate the carbon dioxide in the bundle sheath cell in the form of
malic acid

C4 PLANTS

• Generally of tropical or subtropical origins

• 15% of plants (grasses, corn, sugarcane)
• Partially opened during the day and during the night
• Not exclusive, they utilize both C3 and C4 pathways
• Spatial partitioning – change of location of photosynthesis from the mesophyll
to the bundle sheath cell
• The calvin cycle happens in the bundle sheath cell
• The chlorenchyma are said to form a ring-like arrangement around the bundle
sheath cell – Kranz anatomy
Phosphoenol pyruvate

• The principal effect of the C4 cycle remains to concentrate CO2 in the bundle-
sheath cells where the enzymes of the PCR cycle are located
• C4 crops species can assimilate CO2 at rates 2 to 3 times that of C3 species
• Mesophyll cells surround the bundle sheath cell
• Carbon dioxide from the air is now incorporated into another compound – PEP
or Phosphoenol pyruvate
• Another enzyme – PEP carboxylase – incorporates carbon from carbon dioxide
to PEP to become oxaloacetic acid – which contains 4 carbons, hence the name,
C4 cycle
• Oxaloacetic acid is now reduced to become malic acid, NADPH is oxidized to
NADP
• Malic acid from the mesophyll goes to the bundle sheath cell
• Within the bundle sheath cell within the chloroplast, malic acid is now oxidized
again to become pyruvic acid and CO2 is freed
• CO2 is now utilized in the C3 Cycle
• Pyruvic acid is returned to the mesophyll cell which is phosphorylated to
regenerate PEP so that the cycle can continue.
 In a C3 plant, the mesophyll cells
contain well-formed chloroplasts and
are arranged in parallel layers. In a C
leaf, the bundle sheath cells, as well as
the mesophyll cells, contain
chloroplasts. Further, the mesophyll
cells are arranged concentrically
around the bundle sheath cells:

C4 plants use the enzyme PEP

carboxylase (PEPCase) to fix CO2 to PEP (phosphoenolpyruvate, a C molecule). The
result is oxaloacetate, a C molecule:

In a C4 plant, CO2 is taken up in mesophyll cells, and then malate, a reduced form of
oxaloacetate, is pumped into the bundle sheath cells (Fig. 7.11b). Here, and only
here, does CO2 enter the Calvin cycle. It takes energy to pump molecules, and you
would think that the C4 pathway would be disadvantageous. Yet in hot, dry climates,
the net photosynthetic rate of C4, plants such as sugarcane, corn, and Bermuda grass
is about two to three times that of C3 plants such as wheat, rice, and oats. Why do C4
plants enjoy such an advantage? The answer is that they can avoid photorespiration,
discussed previously. Photorespiration is wasteful because it is not part of the Calvin
cycle. Photorespiration does not occur in C4 leaves because PEPCase, unlike RuBP
carboxylase, does not combine with O2 Even when stomata are closed, CO2 is
delivered to the Calvin cycle in the bundle sheath cells.

When the weather is moderate, C3 plants ordinarily have the advantage, but when
the weather becomes hot and dry, C4 plants have the advantage, and we can expect
them to predominate. In the early summer, C3 plants such as Kentucky bluegrass and
creeping bent grass predominate in lawns in the cooler parts of the United States,
but by mid- summer, crabgrass, a C4 plant, begins to take over.
 CAM Photosynthesis

CAM PLANTS

• Hot, dry environments

• 5% of plants = succulent plants, cacti and bromeliads (e.g., pineapple)
• Crassulacean-acid metabolism
o Studied most extensively in the family Crassulaceae
o Found in 23 families of flowering plants (including Cactaceae and
Euphorbiaceae), 1 family of ferns (Polypodiaceae)
• Whereas C4 Plants represent partitioning in space, CAM is partitioning by the
use of time (temporal partitioning)
o Stomates closed during day to perform C3 cycle
o Stomates open during the night to accumulate carbon dioxide
• Light reaction – occurs during the day
• Calvin cycle – occurs when CO2 is present
• Succulents – a part of their stem and leaf are very watery in nature
• Do not need much watering because they are able to store water in their body.
• Are not CAM exclusive, they also perform the C3 cycle

CAM PATHWAY

Night time

• During night time, the stomates are open to accumulate carbon dioxide from
the atmosphere
• The carbon dioxide is converted to carbonic acid
• Carbonic acid is fixed to PEP by PEP carboxylase to become oxaloacetate
• Oxaloacetate is now reduced to become malate, and is now stored in the
vacuole during the night time.

Day time

• Stomates are now closed, and the malic acid will be released from the vacuole
going to the chloroplast
• Malic acid -> Malate will be converted to pyruvate, and carbon dioxide will be
utilized in the calvin cycle in the C3 pathway -> the enzymes necessary for the
PCR cycle are also dependent on sunlight

Night time

• Starch is converted to PEP so that the cycle can repeat

CAM stands for crassulacean-acid

metabolism; the Crassu-laceae is
a family of flowering succulent
(water-containing) plants that live
in warm, dry regions of the world.
CAM was first discovered in these
plants, but now it is known to be
prevalent among other groups of
plants.

Whereas a C4 plant represents

partitioning in space carbon
dioxide fixation occurs in
mesophyll cells and the Calvin
cycle occurs in bundle sheath
cells-CAM is partitioning by the use of time. During the night, CAM plants use
PEPCase to fix some CO2, forming C4 molecules, which are stored in large vacuoles in
mesophyll cells. During the day, C4 molecules (malate) release CO2 to the Calvin cycle
when NADPH and ATP are available from the light reactions (Fig. 7.12). The primary
advantage for this partitioning again has to do with the conservation of water. CAM
plants open their stomata only at night, and therefore only at that time does
atmospheric CO2 enter the plant. During the day, the stomata close; this conserves
water, but CO2 cannot enter the plant.

Photosynthesis in a CAM plant is minimal because a limited amount of CO 2 is fixed at

night, but it does allow CAM plants to live under stressful conditions.
Photosynthesis and Adaptation to the Environment

The different types of photosynthesis give us an opportunity to consider that

organisms are metabolically adapted to their environment. Each method of
photosynthesis has its advantages and disadvantages, depending on the climate.

C4 plants most likely evolved in, and are adapted to, areas of high light intensities,
high temperatures, and limited rainfall. C4 plants, however, are more sensitive to
cold, and C3 plants do better than C4 plants below 25°C. CAM plants, on the other
hand, compete well with either type of plant when the environment is extremely
arid. Surprisingly, CAM is quite widespread and has evolved in 23 families plants,
including some lilies and orchids! And it is found among nonflowering plants,
including some ferns bearing trees.
 Connecting the Concepts

"Have You Thanked a Green Plant Today? Is a bumper sticker that you may have
puzzled over until now. Plants, you now know, capture solar energy and store it in
carbon-based organic nutrients that are passed to other organisms when they feed
on plants and/or on other organisms. In this context, plants are called autotrophs
because they make their own organic food. Heterotrophs are organisms that take in
preformed organic food.

The next chapter considers cellular respiration, the process that produces ATP
molecules. We have to keep in mind that cells cannot create energy, and therefore
when we say that the cell produces ATP we mean that it converts the energy within
glucose molecules to that found in ATP molecules, with a loss of heat, of course. Why
do cells carry out this wasteful process? Because ATP is the energy currency of cells
and is able to contribute energy to many different cellular processes and reactions.

In the carbon cycle, living and dead organisms contain organic carbon and serve as a
reservoir of carbon. Some 300 million years ago, a host of plants died and did not
decompose. These plants were compressed to form the coal that we mine and burn
today. (Oil has a similar origin, but it most likely formed in marine sedimentary rocks
that included animal remains.)

The amount of carbon dioxide in the atmosphere is increasing steadily, in part

because we humans burn fossil fuels to run our modern industrial society. This
buildup of car bon dioxide will contribute to global warming. Autotrophs such as
plants take in carbon dioxide when they Photosynthesize. Carbon dioxide is returned
to the atmosphere when autotrophs and heterotrophs carry on cellular respiration.
In this way, the very same carbon atoms cycle from the atmosphere to autotrophs,
then to heterotrophs, and then back to autotrophs again. Energy does not cycle, and
therefore all life is dependent on the ability of plants to capture solar energy and
produce carbohydrate molecules.
 SUMMARY

7.1 PHOTOSYNTHETIC ORGANISMS

Photosynthesis produces carbohydrates and releases oxygen, both of which are used
by the majority of living things. Cyanobacteria, algae, and land plants carry on
photosynthesis. In plants, photosynthesis takes place in chloroplasts. A chloroplast is
bounded by a double membrane and contains two main components: the semifluid
stroma and the membranous grana made up of thylakoids.

7.2 THE PROCESS OF PHOTOSYNTHESIS

The overall equation for photosynthesis shows that it is a redox reaction. Carbon
dioxide is reduced, and water is oxidized. During photosynthesis, the light reactions
take place in the thylakoid membranes, and the Calvin cycle reactions take place in
the stroma.

7.3 PLANTS AS SOLAR ENERGY CONVERTERS

Photosynthesis uses solar energy in the visible-light range. Specifically, chlorophylls a

and b absorb violet, blue, and red wavelengths best. This causes chlorophyll to
appear green to us. The carotenoids absorb light in the violet-blue-green range and
are yellow to orange pigments.

The noncyclic electron pathway of the light reactions begins when solar energy
enters PS II. In PS II, energized electrons are picked up by electron acceptors. The
oxidation (splitting) of water replaces these electrons in the reaction-center
chlorophyll a molecules. Oxygen is released to the atmosphere, and hydrogen ions
(H*) remain in the thylakoid space. An electron acceptor molecule passes electrons
to PSI by way of an electron transport chain. When solar energy is absorbed by PS I,
energized electrons leave and are ultimately received by NADP+, which also
combines with H+ from the stroma to become NADPH.

Chemiosmosis requires an organized membrane. The thylakoid membrane is highly

organized: PS l is associated with an enzyme that oxidizes (splits) water, the
cytochrome complexes transport electrons and pump H +; PS l is associated with an
enzyme that reduces NADP+, and ATP synthase produces ATP.

The energy made available by the passage of electrons down the electron transport
chain allows carriers to pump H+ into the thylakoid space. The buildup of H +
establishes an electrochemical gradient. When H + flows down this gradient through
the channel present in ATP synthase complexes, ATP is synthesized from ADP and
by ATP synthase. This method of producing ATP is called chemiosmosis.

7.4 CALVIN CYCLE REACTIONS

The energy yield of the light reactions is stored in ATP and NADPH. These molecules
are used by the Calvin cycle reactions to reduce CO 2 to carbohydrate, namely G3P
which is then converted to all the organic molecules a plant needs.

During the first stage of the Calvin cycle, the enzyme RuBP carboxylase fixes CO 2 to
RuBP producing a 6-carbon molecule that immediately breaks down to two C3
molecules. During the second stage, CO2 (incorporated into an organic molecule) is
reduced to carbohydrate (CH20O). This step requires the NADPH and some of the
ATP from the light reactions. For every three turns of the Calvin cycle, the net gain is
one G3P molecule; the other five G3P molecules are used to re-form three molecules
of RuBP This step also requires ATP for energy. It takes two G3P molecules to make
one glucose molecule.

7.5 OTHER TYPES OF PHOTOSYNTHESIS

In C4 plants, as opposed to the C3 plants just described, the enzyme PEPCase fixes
carbon dioxide to PEP to form a 4-carbon molecule, oxaloacetate, within mesophyll
cells. A reduced form of this molecule is pumped into bundle sheath cells, where CO2
is released to the Calvin cycle. PEPCase has an advantage over RuBP carboxylase
because RuBP carboxylase, but not PEPCase, combines O 2 with RuBP instead of CO2
when the stomata close and the concentration of O2 rises. C4 plants avoid this
complication by a partitioning of pathways in space. Carbon dioxide fixation occurs
utilizing PEPCase in mesophyll cells, and the Calvin cycle occurs in bundle sheath
cells.
In CAM plants, the stomata are open only at night, conserving water. PEPCase fixes
CO2 to PEP only at night, and the next day, CO2 is released and enters the Calvin cycle
within the same cells. This represents a partitioning of pathways in time: Carbon
dioxide fixation occurs at night, and the Calvin cycle occurs during the day. CAM was
discovered in desert plants, but since then it has been discovered in many different
types of plants.