Photosynthesis

Photosynthesis is the ultimate source of cell biological energy.

It is the capture of solar energy (by the photosynthetic organisms) and its
conversion into the energy of biomass (complex macromolecules)

Basically

Photosynthetic plants trap solar energy in the form of ATP and NADPH, which
they then use as an energy source(s) to make carbohydrates from carbon
dioxide and water. They simultaneously release oxygen into the atmosphere.

General photosynthetic equation -

6H2O + 6H2O + 6CO2 ------> C2H12O6 + 6O2 + 6H2O

As we have seen aerobic heterotrophs use the oxygen so formed to degrade the
energy rich organic products of photosynthesis to CO2 and H2O in order it
generate ATP for their own use. The CO2 formed by respiration in heterotroph
returns to the atmosphere to be used again by the photosynthetic organisms.

THUS

Solar energy provides the driving force for the continuous cycling of atmospheric
carbon dioxide and oxygen through the biosphere.
The Visible Light System and Photosynthesis

Visible light is part of the electromagnetic spectrum. This light interacts with
pigments such as chlorophyll, which are molecules that absorb the energy of light
and provide the power for photosynthesis.

Light is very difficult to describe; it is a particle and a wave? The photon is a

packet of light that has wavelength associated with it, so a photon can be
thought of as both a particle and a wave.

The energy contained in a single quantum (or photon) is directly proportional to

its wavelength. The constant of proportionality is named Planck's constant h.

Where "λ " (lambda) is wavelength

"µ " (mu) is equal to frequency (1Hz= 1 cycle per second)

"c" is the speed of light = 3.0X1010 cm per second.

Therefore, energy of a proton E = µ h

And µ = c/ λ

Therefore E = h c/ λ

Therefore, if "h" and "c" are constants (or nearly so) then energy varies as the
inverse of (wavelength).

NB the longer the wavelength the less energy.

Absorption Spectra (and the colours of things)

Different photons have different wavelengths and these show as different

colours e.g. blue light has short wavelengths and red light has longer
wavelengths.
When an object absorbs a certain wavelength of light then that wavelength can
no longer bounce back into your eye therefore, you do not see it.

Conversely: If objects reflect certain wavelength of light (reflect certain colours)

then those colours bounce off its surface (are not absorbed) therefore they can
enter your eyes and you can see them.

Therefore reflected light can be seen; absorbed light can not be seen. If you
shine different wavelengths of light on a substance (say a molecule (e.g.
chlorophyll a) then some of those wavelengths will be absorbed and some will be
reflected. A plot of these absorptions is called can absorption spectra.
NB: Chlorophyll is green.

Absorption of Light Excites Molecules

The ability of a chemical compound to absorb light depends upon the

arrangement of electrons around the atomic nuclei in its structure. When a
molecule (or atom) absorbs a photon an electron is boosted to a higher energy
level. This happens on all or none basis. To boost the electron, a photon must
have a certain minimum energy (hence the name quantum).

The molecule that has absorbed the photon is in an excited state, which is
usually very unstable.
When the excited electron falls back to its ground state it gives up the energy that
was absorbed. Some is given off as heat some as light. But the total energy is
the same therefore, the light photon that is remitted is a lower energy, therefore a
longer wavelength.

Photosynthetic Pigments and The Structure of Chlorophyll

The photosynthetic pigments in chloroplasts are located in the thylakoid

membranes. The primary light absorbing pigments in higher plants are the
chlorophylls. These are Mg2+ complexes of molecules resembling the
protoporphyrin ring system of haemoglobin.
Photosynthetic cells of higher plants always contain types of chlorophyll. One is
always chlorophyll a the other often chlorophyll b. In addition to the chlorophylls
the thylakoid membranes always contain secondary light absorbing pigments.
Together they are called accessory pigments. These include various
carotenoids, which may be yellow, red or purple. The most important
carotenoids are B-carotene and xanthophyll.

These accessory pigments absorb light in wavelengths of light other than those
of the chlorophylls therefore, increase the range of light that can be utilised.

Chloroplast Structure:

Recall:

A double membrane system or envelope that controls molecular traffic inwards

and outwards from them surrounds each chloroplast. Within the chloroplasts
there is another series of membranes the Thylakoids which contain the
photosynthetic pigments. Certain regions of the thylakoids are stacked to form
grana (granum sing.) membranes called lamellae connect grana. These
membranes are also called appressed membranes = grana and non-
appressed membranes = lamellae. A liquid material called stroma surrounds
these membranes. Inside each thylakoid is a fluid filled space called the
thylakoid compartment.
The Photosynthetic Process

Some of the processes involved in photosynthesis require light while others do

not. Therefore, there are two basic parts to photosynthesis, the light reactions
(which are light dependent) and the light independent reactions (or dark
reactions).

One can think of the light dependent reactions as a way to increase the free
energy of the system and the light independent reactions as a way to convert
that new free energy into the bonds of glucose.

Photosystems

The light-absorbing pigments of thylakoid membranes and their associated

electron carriers are arranged in functional sets or clusters. These clusters are
called photosystems. For example: in spinach chloroplasts these photosystems
contain about 200 chlorophyll molecules and about 50 carotinoids, arranged in
what are called light harvesting antenna. These photosystems can absorb light
over the entire visible spectrum but especially well between 400 to 500 nm and
600 to 700 nm.

Photochemical Reaction Centre of the Light Harvesting Antenna

When a chlorophyll molecule in the thylakoid membrane is excited by light, the
energy level of an electron in its structure is boosted by an amount equivalent to
the energy of the absorbed light and the chlorophyll becomes excited. The
packet of excitation energy (The Exciton) now migrated rapidly through the light
harvesting pigment molecules to the reaction centre of the photosystem where
it causes an electron to acquire the large amount of energy.

The photochemical reaction centre of photosystems II is called P680. The

photochemical reaction centre of photosystems I is called P700.

Photosystems I and II

The thylakoid membranes of plant chloroplasts have two different kinds of

photosystems each with its own set of light harvesting chlorophyll and carotenoid
molecules and the photochemical reaction centre.

Photosystem I - is maximally excited by light at longer wavelengths. (P700)

Photosystems II - is maximally excited by shorter wavelengths. (Less the 680).

(P680)

NB: Generalisation
All oxygen-evolving photosynthetic cells. (E.g. those of higher plants and
cyanobacteria) contain both photosystems I and II, whereas all other species of
photosynthetic bacteria, which do not evolve oxygen only contain photosystem
I.

The Light Reaction

These increase the free energy made available to the system. This free energy
can be used in three ways:

1) To build the chemiosmotic or proton gradient.

2) Generate ATP.

3) Reduce NADP+ to NADPH.

There are two ways to generate ATP

1) Non cyclic photophosphoraltion.

2) Cyclic photophosphoralation.
These two systems differ in the route taken by the "light activated" electrons and
in some of the products formed.

Non Cyclic Photophosphoralation

These events begin with photons being absorbed in photosystem II and its
energy is shunted into P680 and the chlorophyll a of this photoactivation centre
passes the energy rich electron on into the electron transport system. Once in
the electron transport system of the thylakoid membrane the electron is passed
from electron carrier to electron carrier eventually entering photosystem I. In the
"tumbling" down of the electron transport chain the electron gradually loses
energy. Some of that energy will be used to "pump protons" across the thylakoid
membrane into the "lumen" (thylakoid compartment).

NB: Each light activated electron allows the "proton pump" to pump one proton
across the membrane. These partly spent electrons then pass into the next
photosystem I P700 where they receive another "boost" to their highest energy
level yet. But these electrons do not pump protons they use their energy to
reduce NADP+ to NADPH.
Chemiosmotic Phosphoralation

The cyclic and non-cyclic events of the electron carriers and photosystems of the
thylakoid membrane serve to pump protons into the lumen of the thylakoids.

Just as in the mitochondria outer compartment the protons are allowed to flow
back into the stroma (which has a high OH- (hydroxide) content (i.e. very basic).
The controlled flow of H+ down this concentration gradient occurs through the so-
called CFi particles. Which turn out to be ATP synthase complexes.
These systems therefore contribute ATP and high-energy electrons NADPH
(reducing power) to the next series of reaction in photosynthesis. (i.e. the dark
reaction the Calvin cycle).