CHLOROPLAST
CHLOROPLAST
Introduction
Chloroplasts are organelles involved in photosynthesis, the process of formation of
carbohydrates from carbon-di-oxide and water in the presence of sunlight by the plants.
Chloroplasts were first described as ‘chloroplyllkornern’ (chlorophyll granules) by German
botanist Hugo von Mohl in 1837, though because of their large size they were seen long back by
Nehemiah Grew who described them in 1682 as green precipitates! It was A.F.W.
Schimper who in 1883 introduced the term ‘plastids’ as a substitute for chlorophyll granule.
In the same year A. Mayer described ‘grana’ as dense dot like structures embedded in the
‘stroma’ of these plastids. The terms grana and stroma are still in use but chloroplast has
replaced the term plastids for the green organelles of leaves. T.Engelmann a German
biologist in 1881 identified the chloroplasts as the sites where photosynthesis occurs. He
observed that outside the cells of green alga Spirogyra just close to the large ribbon shaped
chloroplasts, bacteria would collect probably to utilize the oxygen being liberated during
photosynthesis in the chloroplasts (bacterial chemotaxis).
In addition to photosynthesis plastids are also involved in:
s in the
chloroplast providing the plant with the nitrogen for the synthesis of amino acids and
nucleotides.
A typical plant cell may have 20-40 chloroplasts. There are many alga which have single
chloroplast per cell that occupies most of the cell (remember cup shaped chloroplast of
Chlamydomonas).
Chloroplasts are organelles that belong to the plastid family. This group of organelles which
are present in various plant cells differ in their color and function. In young meristems small
pro-plastids develop which mature according to the requirement of the cell and also as
dictated by the nuclear genome.
For example:
Etioplasts- these develop in leaves grown in dark. These have a yellow chlorophyll
precursor and semi crystalline membranes. When exposed to light these develop to
form chloroplasts.
1) Envelope: Just like nucleus and mitochondria, chloroplasts are bound by a double
membrane envelope called outer membrane and inner membrane. Both the
membranes are made up of phospholipid bilayer. The space in between the
membranes is inter-membrane space. The outer membrane of the chloroplast
envelope contains ‘porins’ and so is freely permeable to small molecules. Whereas
the inner membrane is impermeable to ions and metabolites and they need specific
membrane transporters (integral membrane proteins) to enter the chloroplast. The
outer membrane has very less protein content about 30% and have unusually have
very little phospholipid. These have a high percentage of galactose containing
glycolipids, which have several double bonds in their fatty acids making the
membranes highly fluid. This facilitates the lateral diffusion of the protein complexes
in the membrane.
Unlike the mitochondria, the inner membrane of the chloroplast is not
involved in either electron transport or photosynthesis. Instead the
thylakoid membrane contains the electron transport system. The protons
are pumped across into the lumen of the thylakoids creating a proton
gradient that drives the ATP synthesis as protons cross back into the
stroma.
2) Thylakoids: The internal membrane system (separate from the double membrane
envelope) is organized into disc shaped/flattened membranous structures with empty
lumen called thylakoids that are frequently arranged in stacks. Each stack is called granum
(plural: grana). Adjacent grana are connected by stromal thylakoid (thylakoids in direct contact
with the stroma).The thylakoid membrane contains the light absorbing pigments, the chain of
electron carriers (ETC) and the ATP synthesizing machinery (in this sense it is analogous to the
inner membrane of mitochondria). The space inside the thyakoid is the lumen. The protons are
pumped across this membrane from the stroma to the thylakoid lumen. This results in
formation of an electrochemical gradient which generates ATP when protons are transported
back to the stroma.
3) Stroma: The space enclosed by the inner membrane of the chloroplast is called stroma.
Stromal matrix is rich in metabolic enzymes and has small double stranded circular DNA
molecules, ribosomes (70-S) and plastoglobuli (lipid granules). Dark reaction or the C-3
cycle (Calvin cycle) takes place in stroma. Enzymes for amino acid synthesis and fatty acid
synthesis are also present in stroma.
Semiautonomous nature
Some organelles enjoy some amount of autonomy or independence like in growth, division,
production of some enzymes, they are known as semiautonomous organelles. Mitochondria
and chloroplast fall in this category. They are able to do this because they have their own DNA/
genome. Both the organelles are also known to have evolved from free living prokaryotes,
according to the endosymbiotic theory. Endosymbiotic theory was proposed
by Lynn Margulis in1960s. She proposed that primitive anaerobic bacteria engulfed another
aerobic bacteria and instead of endocytosis endosymbiosis occurred, that is the association
became mutually beneficial for both the bacteria and resulted in early aerobic heterotrophic
eukaryote. Further, when this eukaryote engulfed photosynthetic bacteria another
endosymbiotic relationship evolved and resulted in an autotrophic eukaryote. This theory
also explains double membrane of these organelles, one derived from each of the organism
involved.
Since the autonomy is not complete and the chloroplasts depend on nuclear genome for
majority of their proteins and enzymes, hence, these organelles are known as
semiautonomous in nature. For example, in chloroplast the enzyme RuBisCo is made up of
multiple units of polypeptides assembled into two types of subunits called the large chain
and small chain. The large chain gene is a part of chloroplast DNA but the small chain genes
are located on the nuclear genome. The small chain subunit is thus synthesized in the
cytosol and is imported into the stroma.
Light reaction
Sunlight is captured by photosynthetic pigments the most abundant of which are the –
chlorophylls. The photosynthetic pigments are arranged into photocenters with each
photocenter containing hundreds of antenna pigment molecules which absorb the photons
and transfer energy to the reaction center chlorophyll.
The energy derived from the sunlight excites an electron (derived from the splitting of water
molecule and the generation of O2) of the chlorophyll thus converting the energy of the
sunlight to potential chemical energy. This electron then moves along an electron transport
chain (a series of electron carrier proteins located in the thylakoid membrane). Four protein
complexes located in the thylakoid membrane function as electron carriers and are also
involved in the synthesis of ATP and NADPH.
Energy initially harvested by PSII (photosystem II) is used to split a water molecule within
the thylakoid lumen. Electrons are then carried by plastoquinone to the cytochrome bf
complex where they are transferred to low energy state and protons are pumped into the
thylakoid lumen. Electrons are transferred to photosystem I by plastocyanin(PC). At PSI the
sunlight absorbed is again used to generate high-energy electrons. The photosystem I uses the
high energy electrons to reduce NADP+ to NADPH.Light is captured by the photosystems.
Photolysis of water takes place. Electron travels from PSII to PSI and NADPH is synthesised from
NADP. Proton gradient is generated across the thylakoid membrane which drives ATP formation
in the stroma. Finally, the high energy electrons that move through the electron transport chain
of light reaction and the H+ convert the NADP+ to NADPH. The movement of electron through
the photosystem I and II generates both ATP and NADPH which are then used in the Calvin cycle
to convert CO2 to carbohydrates.Another electron transport pathway that operates to produce
ATP is called the cyclic electron flow. However unlike the non-cyclic pathway no NADPH is
generated. The light energy is harvested at the photosystem I and the high energy electrons
generated are transferred to cytochrome bf complex coupled to which protons are transferred
across the thylakoid membrane establishing a proton gradient and thus synthesizing ATP .
Plastocyanin returns these electrons to photosystem I thus completing the cycle. Thus PS I
participates in both cyclic and non cyclic pathways forming ATP and NADPH depending on the
requirement of the cell.