Week 2: Photosynthesis (Form and Function of Plants)

Photosynthesis is conversion of light energy into chemical energy in the form of sugar and
other organic molecules by plants.

What is photosynthesis (refer to diagram 1)?

● Photosynthesis occurs as two reactions which are closely integrated in the
chloroplasts of green tissues.
● These reactions are known as the light reaction (light-dependent) and the dark
reaction (light independent).
● The light reaction takes place in the thylakoid membrane of the chloroplasts and
produces oxygen, ATP and NADPH.
● The dark reaction takes place in the stroma of the chloroplast and utilises the ATP
and NADPH from the light reaction to convert/fix carbon dioxide into sugars.

Photosynthesis captures light energy as reduced carbon:

Light-dependent reactions, the first step is the capture of light energy as ATP and reducing
power, NADPH → (ATP and NADPH) → light-independent reactions, the second step is
the transfer of energy and reducing power from ATP and NADPH to carbon dioxide, to
produce high-energy and reduced sugars.

Carbon fixation: The process where photosynthetic organisms (such as plants) turn inorganic
carbon into organic compounds (carbohydrates), reactions take place in the chloroplast
stroma.

Where does photosynthesis occur and how are form and function integrated (refer to diagram 2
and 3)?
● Chloroplasts are formed from three membranes that enclose three compartments.
● Outer membrane covers the entire surface of the organelle with the inner membrane
just inside the outer membrane.
● Intermembrane compartment between these two membranes.
● Fluid within the compartment formed by the inner membrane is called stroma.
● Third membrane system consists of thylakoid membranes which form flattened,
closed sacs called thylakoids.
 ● Space enclosed by thylakoid is called thylakoid lumen.
● Thylakoids are arranged into stacks called grana.
● Grana are interconnected by flattened, tubular membranes called stromal lamellae.
● Thylakoid membranes and stromal lamellae house the molecules of the
light-dependent reactions of photosynthesis.
● Light-independent reactions are concentrated in the stroma.
● Carbon dioxide required for photosynthesis diffuses into cells containing
chloroplasts after entering through stomata.
● Oxygen and water produce diffuses from cells and exit through the stomata.

The most important structure for photosynthesis is the chloroplast which is a specialised
organelle that carries the whole biochemical machinery that drives photosynthesis. The structure
of the leaf facilitates the functional reactions that take place. On the upper surface of the leaf,
that is directly facing sunlight, is this epidermal layer which acts as a protective covering for
the leaf. The palisade mesophyll (follows the epidermal layer) which contains almost rectangular
cells that have many different chloroplasts. The spongy mesophyll layer follows which has many air
spaces which facilitates the movement of gases such as carbon dioxide and oxygen as well as the
movement of water from the inside of the leaf to the outside. On the lower surface in most
plants, are stomata – specialised epidermal cells that allow for the movement of gases between
the atmosphere and the mesophyll region where carbon dioxide is required for carbon fixation and
oxygen is evolved inside the mesophyll region and then exits via the stoma.

What is the nature of light that makes it suitable for photosynthesis?

Light behaves as a wave or a stream of particles. Visible light is part of the
electromagnetic spectrum. Plants use the visible light of the electromagnetic spectrum
(400-700 nm) for photosynthesis. The wavelengths can range from nanometres (gamma)
to kilometres (radio).

During photosynthesis, light is absorbed by chlorophyll (green pigment) and carotid (orange-yellow
pigment). Chlorophyll B acts as an accessory pigment that also absorbs light:
● Light is absorbed by molecules of green pigments called chlorophylls and
yellow-orange pigments called carotenoids.
● Carotenoids are accessory pigments that absorb light energy at a different
wavelength than those absorbed by chlorophylls.
 ● These pigments (and others) are located in the thylakoid membranes of
chloroplasts.
● Held in specific position and orientation by proteins embedded in thylakoid
membranes.

Chlorophyll A is the main light absorbing component – it is where the main reactions that
facilitate the conversion of light to chemical energy actually take place. Chlorophyll B is an
accessory pigment. All the pigments of photosynthesis are all capable of absorbing light. The
carotenoids also have an accessory function that absorbs different parts of the wavelengths
that the chlorophyll molecules (A and B) do not absorb. These carotenoid pigments have other
functions, a UV quenching function – this is a protective mechanism that protects the plant's
fragile biochemical machinery that drives photosynthesis from damage. The pigments are
associated with the thylakoid membranes which hold a range of different proteins which react in
the different reactions of photosynthesis. Chlorophyll A and B are very similar structurally – they
have a light absorbing head and a hydrophilic tail which associates itself with the thylakoid
membrane. The hydrophilic light-absorbing head is situated in a way that it is associated with
the stromal components and it is well exposed so that it is able to absorb light. Each chlorophyll
molecule has a magnesium component which is central to the light absorbing head. Electrons are
absorbed by these ring structures that are associated with the light-absorbing head. The tail is
there to insert the light-absorbing head into the thylakoid membranes. The carotenoids have a
light-absorbing region which runs throughout the whole molecule.

Chlorophyll molecules are bound to specific membrane proteins – arranged efficiently to

capture light energy – refer to diagram 4.

Chlorophyll A functions as the reaction centre of photosynthesis. Chlorophyll B is an accessory

pigment which is very important for the light reactions of photosynthesis. They are the main
components in absorbing light. They are structured with a porphyrin head which is not tightly
integrated into the thylakoid membrane but this head forms complexes that are important for
light absorption. The reaction centres have chlorophyll A.

Structure of Light Reaction Components – Refer to Diagram 5

Which wavelengths are absorbed by chlorophylls and carotenoids?
Chlorophyll A and B both absorb the red and blue wavelengths, but each has a different
peak.

What is the effect of these wavelengths on photosynthesis?

Maximum photosynthesis at red and blue wavelengths and also a bit at green and yellow
light. This is because the green and yellow light are continuously reflected (bounced)
between the chloroplasts and with each reflection a bit of light is absorbed.

Chlorophylls A and B and carotenoids all absorb different wavelengths when they are not inside a
living system. When both chlorophylls and carotenoids are used together, they use maximum use of
white light. Irrespective of the type of light that is being absorbed, that type of light has the
same amount of energy to drive photosynthesis. Blue, red, green or yellow light are all used once
the reaction centre of the photosystems are excited, that reaction is exactly the same,
irrespective of the type of light.
The sun emits light at a range of different wavelengths, but much of the very short
wavelength light is absorbed by Earth’s atmosphere. The earth’s atmosphere blocks much
of the short wavelength light.

Absorption spectra of photosynthetic pigments:

● All chlorophyll based photosynthesis systems use chlorophyll A. Different antenna
systems use different subsets of accessory pigments which expand the range of
light absorbed.
● Chlorophyll B is found in land plants, green algae and cyanobacteria.
● Carotenoids are found in all chlorophyll-based photosynthesis systems.
● Phycoerythrin is found in cyanobacteria and non-green algae, and phycocyanin in
cyanobacteria.

Absorbance spectrum of photosynthesis in a green plant – action spectrum:

● The photosynthetic action spectrum shows the rate of photosynthesis that occurs
when a single wavelength of light shines on a plant.
● A different action spectrum can be measured in other organisms as a function of
their accessory pigments.
● Reaction centre chlorophylls absorb maximally at 680 and 700 nm. Longer
wavelength light does not have sufficient energy to drive photosynthesis in plants.

Organisation of Pigments in the Photosystems – Refer to Diagram 6

● Light absorbing pigments are organised with proteins and other molecules into
large complexes called photosystems.
● Photosystems are sites at which light energy is absorbed and converted into
chemical energy.
● Two photosystems carry out different parts of the light-dependent reactions.
● These photosystems are located in the thylakoid membranes of the chloroplasts.

There are two different photosystems. Photosystem I is associated with NADPH synthesis
and photosystem II is linked to a group of enzymes that carry out initial reactions of
splitting water into electrons, protons and oxygen. These photosystems are connected by
reactions that occur on the photosystems themselves, and close to the photosystems and
along the thylakoid membranes, and even the stroma (refer to diagram 7).
Each photosystem has an antenna complex and a reaction centre. The reaction centres
contain chlorophyll A. Chlorophyll B and the carotenoids are situated in the antenna
complexes. Light energy is important because it contains photons which are transferred
within the antenna complexes. There is no transfer of electrons that takes place within the
antenna complexes. The only time that an electron transfer action takes place, happens
within the reaction centres that contain chlorophyll A.

Antenna complex consists of several chlorophyll and carotenoid molecules anchored by

membrane proteins and held in a specific orientation for optimal light interception. The
reaction centre contains special subsets of chlorophyll A molecules complexed with
proteins (refer to diagram 8).

The photosystems have a very complex set of proteins and other molecules that are bound to
these photosystems and their main functions are to carry the reactions that are
electron-reductive and oxidative reactions of photosynthesis. Photosystems contain a pair of
chlorophyll A molecules that are part of the reaction centres – this is where the first
determining step of photosynthesis reactions actually takes place (where electron transfer is
initiated). Antenna complexes are really only there to direct light towards the specialised pair of
chlorophyll A molecules. The passage of electrons takes place within the photosystems from
different molecules that are part of these photosystems.
The chlorophyll A molecules form the reaction centres of both photosystems I and II. The
photosystems interact with each other via electron transport and the electrons move from
photosystem II all the way up to photosystem I and the electrons are deposited on NADP
reductase to form NADPH. The photosystems are embedded in the thylakoid membrane, but parts
of these photosystems also interact with the lumen and stroma, therefore they are considered
transmembrane photosystems. The pigments that are involved in antenna complexes do not sit
on any of these structures.

Capturing of light energy by the photosystems for ATP and NADPH production (refer to diagram
9):
● Light energy in the form of photons is absorbed by pigment molecules of the
antenna complex.
● Absorbed light energy is transferred to P680 or P700 molecules in the reaction
centre.
● Absorbed light is converted to chemical energy when an excited electron from the
special chlorophyll A molecule is transferred to a primary acceptor, also in the
reaction centre.
● High energy electrons pass out of the reaction centre and out of the photosystems
to the electron transfer system.

The first reaction that results in chemical energy or organic molecules is controlled at the
reaction centres and this happens in a unidirectional fashion or the z-scheme this means the
reaction starts off at one point and finishes off at another point, but it never reconnects to
the point where it originally started from. This light energy is captured by the photosystems and
leads to the production of ATP and NADPH. Both photosystems have the ability to absorb light
energy and they do this through photons. The antenna complexes direct this light energy in the
form of photons towards the reaction centres. That light is then transferred to the reactions
centres and excitations within the reaction centres, in the chlorophyll molecule, results in an
electron transfer reaction and the chlorophyll A molecule is therefore, the primary sector of
excited electrons and that is the very first time that physical energy is converted to chemical
energy and this reaction occurs within the chlorophyll molecule which makes the chlorophyll molecule
a primary acceptor. Once the chlorophyll A molecule has accepted these electrons, the electrons
are then passed on out of both photosystems and transferred to electron carriers in this
unidirectional fashion (the z scheme). Only when the energy from the photon reaches the
reaction centre, is the energy from the antenna complex sufficient to excite the electron in
the chlorophyll A complex to such an extent that the electron leaves its orbital. Chlorophyll A is
now an electron short. This loss of electrons must be replaced. This electron is passed onto the
electron acceptor.

Only when the energy from the photon reaches the reaction centre, is the energy from the
antenna complex sufficient to excite the electron in the chlorophyll A complex to such an
extent that the electron leaves its orbital. Chlorophyll A is now an electron short. This
lost electron must be replaced. We will focus on photosystem II as the start of the electron
transfer even though an electron is lost nearly simultaneously from P700 and P680.

Refer to diagram 10:

1. Electrons from the water-splitting reaction system are transferred to P680
chlorophyll A in the reaction centre of photosystem II. Electrons are raised to an
excited state using energy passed to the reaction centre from the antenna complex.
2. Electrons flow through a short chain of carriers within photosystem II and then
transferred to plastoquinone, a mobile carrier.
3. Plastoquinones pass electrons to the cytochrome complex. As it accepts and
releases electrons, protons (H+) are pumped from the stroma into the thylakoid
lumen. These protons drive ATP synthesis.
4. Plastocyanin transfers electrons from the cytochrome complex to photosystem I.
 5. Electrons from plastocyanin replace electrons lost from photosystem I chlorophyll
A. This electron is excited to higher energy levels by energy transfer from the
antenna complex.
6. After passage through the carrier within photosystem I, electrons are transferred to
iron-sulphur protein, ferredoxin a mobile carrier.
7. Ferredoxin transfers high energy electrons to NADP+, the final acceptor of the
non-cyclic pathway. NADP+ reduced to NADPH by NADP+ reductase.
8. Proton (H+) pumping by plastoquinones and cytochrome complex creates
concentration gradient of proton (H+) with high concentration in lumen and low
concentration in stroma.
9. Proton (H+) gradient supplies energy that drives ATP synthesis via ATP synthase.
10. Due to the gradient, proton (H+) flows across the inner membrane into the matrix
through a channel in ATP synthase.
11. Flow of protons (H+) activates ATP synthase, making the headpiece and stalk
rotate.
12. Changes in shape and position as it turns catalyses the synthesis of ATP from ADP
and Pi (via proton motive force) provides energy for ATP synthesis.

This whole process of electron flow from photosystem II to photosystem I is known as

non-cyclic electron transport. Result of non-cyclic electron transfer between photosystem II
and photosystem I is the production of oxygen, ATP and NADPH.

Chemiosmosis is the process of using a proton (H+) gradient to power ATP synthesis.
Synthesis of ATP coupled to transfer of electrons energised by photons of light is called
photophosphorylation.

Sometimes photosystem I works independently of photosystem II in a process called cyclic

electron flow.

● Electrons pass through the cytochrome complex and plastocyanin to P700

chlorophyll A in the reaction centre photosystem I where they're excited.
● Electrons then flow from photosystem I to ferredoxin, but rather than being used
for NADP+ reduction, they flow back to P700 via cytochrome complex.
● Electrons pass to plastocyanin and onto photosystem I where they receive energy
from light.
 ● Each time electrons flow around the cycle, more proton (H+) is pumped across
thylakoid membranes, driving ATP synthesis.
● Net result of cyclic electron flow is conversion of light energy into chemical
energy of ATP without production of NADPH or oxygen.
● Cyclic electron flow is adapted to prevent excessive NADPH build-up.
● Excess NADPH can lead to reactive oxygen species forming which can damage
cells.

Summary of capturing of light energy by the photosystems for ATP and NADPH production:
This whole process of electron flow from photosystem II to photosystem I is known as
non-cyclic electron transport. The result of this between photosystem II and photosystem I is
the production of oxygen, ATP and NADPH. Light hits the P680 reaction centre exciting
the electrons and that initiates their movement with photosystem II. Because the P680
reaction centre has lost electrons, it then grabs electrons from water through
water-splitting, those electrons (four of them) move along a set of electron carriers
through redox potential reactions and are then deposited in plastoquinones (a mobile
electron carrier that is able to shuttle between photosystem II and the cytochrome
complex. As this is happening, this allows for the streaming of protons from the stroma
into the lumen. These protons along with the protons being made by hydrogen splitting
increases the protons (H+) inside the lumen, causing the lumen to become more acidified.
In the meanwhile, electrons are moved from the cytochrome complex to plastocyanin –
which is a more water-soluble electron carrier that is able to shuttle between the
cytochrome complex and photosystem I. Plastocyanin takes the electrons and passes them
over to the P700. At the same time, excitation energy from the light energises the P700
and electrons are passed on over to ferredoxin which is positioned closely to NADP+
reductase. Ferredoxin then passes the electrons to this particular enzyme (NADP+
reductase) which is able to use this energy and generate NADPH from NADP+. The ATP
is generated through the mobility of the H+s that pass-through ATP synthase and this
drives the motor of ATP synthase which results in conversion of ADP into ATP through a
phosphorylation reaction.