Chloroplast

Plastids are cell organelles which occur only in plant cells. They multiply by division and in most cases are inherited maternally. This means that all the plastids in a plant usually have descended from the proplastids in the egg cell. During cell differentiation, the proplastids can differentiate into green chloroplasts, colored chromoplasts, and colorless leucoplasts.

Plastids possess their own circular chromosome as well as enzymes for gene duplication, gene expression, and protein synthesis.

The plastid genome (plastome) has properties similar to that of the prokaryotic genome, as for instance in the cyanobacteria, but encodes only a minor part of the plastid proteins The majority of these proteins are encoded in the nucleus and subsequently are transported into the plastids.

The proteins encoded by the plastome comprise part of the proteins of photosynthetic electron transport and of ATP synthesis. Proplastids are very small organelles (diameter 11.5 mm). They are undifferentiated plastids found in the meristematic cells of the shoot and the root. They, like all other plastids, are enclosed by two membranes forming an envelope. Chloroplasts (Fig. 1) are formed by differentiation of the proplastids

A mature mesophyll cell contains about 50 chloroplasts. By definition chloroplasts contain chlorophyll. However, they are not always green. Chloroplasts are lens-shaped and can adjust their position within the cell to receive an optimal amount of light. In higher plants their length is 3 to 10 mm. The two envelope membranes enclose the stroma. The stroma contains a system of membranes arranged as flattened sacks (Fig. 1), which were given the name thylakoids (in Greek, sac-like) by

During differentiation of the chloroplasts, the inner envelope membrane invaginates to form thylakoids, which are subsequently sealed off. In this way a large membrane area is provided as the site for the photosynthesis apparatus . The thylakoids are connected to each other by tubelike structures, forming a continuous compartment of the thylakoid space. Many of the thylakoid membranes are squeezed very closely together; they are said to be stacked. These stacks can be seen by light microscopy as small particles within the chloroplasts and have been named grana.

Differentiation of a proplastid to a chloroplast.

The inner envelope membrane is a permeability barrier for metabolites and nucleotides, which can pass through only with the aid of specific translocators. In contrast, the outer envelope membrane is permeable to metabolites and nucleotides (but not to macromolecules such as proteins or nucleic acids). This permeability is due to the presence of specific membrane proteins called porins, which form pores permeable to substances with a molecular mass below 10,000 Dalton.

About 10 to 100 identical plastid genomes are localized in a special region of the stroma known as the nucleoide. The ribosomes present in the chloroplasts are either free in the stroma or bound to the surface of the thylakoid membranes. In leaves grown in the dark (etiolated plants), the plastids have a yellowish color and are termed etioplasts. These etioplasts contain some, but not all, of the chloroplast proteins.

They are devoid of chlorophyll but contain instead membrane precursors, termed prolaminar bodies, which probably consist of lipids. The etioplasts are regarded as an intermediate stage of chloroplast development. Leucoplasts are a group of plastids that include many differentiated colorless organelles with very different functions (e.g., the amyloplasts), which act as a store for starch in non-green tissues such as root, tubers, or seeds. Leucoplasts are also the site of lipid biosynthesis in non-green tissues.

Lipid synthesis in plants is generally located in plastids. The reduction of nitrite to ammonia, a partial step of nitrate assimilation, is also always located in plastids. In those cases in which nitrate assimilation takes place in the roots, leucoplasts are the site of nitrite reduction. Chromoplasts are plastids that, due to their high carotenoid content, are colored red, orange, or yellow. They are the same size as chloroplasts but have no known metabolic function.

Their main function may be to house the pigments of some flowers and fruit (e.g., the red color of tomatoes).

Mitochondria- structure
Mitochondria are the site of cellular respiration where substrates are oxidized for generating ATP. Mitochondria, like plastids, multiply by division and are maternally inherited. They also have their own genome (consisting in plants of a large circular DNA strand and often several small circular DNA strands) and their own machinery for gene duplication, gene expression, and protein

The mitochondrial genome encodes only a small number of the mitochondrial proteins most of them are encoded in the nucleus. Mitochondria are of endosymbiontic origin. the mitochondrial outer membrane contains porins that render this membrane permeable to molecules below a mass of 4,000 to 6,000 Dalton, such as metabolites and free nucleotides. The mitochondrial inner membrane contains the proteins of the respiratory chain. In order to enlarge the surface area of the inner membrane, it is invaginated in folds

Chloroplasts-functionphotosynthesis
In photosynthesis photon energy splits water into oxygen and hydrogen, the latter bound as NADPH. This process, termed the light reaction, takes place in the photosynthetic reaction centers embedded in membranes. It involves the transport of electrons, which is coupled to the synthesis of ATP. NADPH and ATP are consumed in a socalled dark reaction to synthesize carbohydrates from CO2

Chlorophyll is the main photosynthetic pigment.In photosynthesis of a green plant, light is collected primarily by chlorophylls pigments that absorb light at a wavelength below 480 nm and between 550 and 700 nm. When white sunlight falls on a chlorophyll layer, the green light with a wavelength between 480 and 550 nm is not absorbed, but is reflected. This is why plant chlorophylls and whole leaves are green. The basic structure is a ring made of four pyrroles, a tetrapyrrole, which is also named porphyrin.

Mg++ is present in the center of the ring as the central atom. Mg++ is covalently bound with two N-atoms and coordinately bound to the other two atoms of the tetrapyrrole ring. A cyclopentanone is attached to ring c. At ring d a propionic acid group forms an ester with the alcohol phytol. Phytol consists of a long branched hydrocarbon chain with one C-C double bond. It is derived from an isoprenoid, formed from four isoprene units This long hydrophobic hydrocarbon tail renders the chlorophyll highly soluble in lipids and therefore promotes its presence in the membrane phase.

Chlorophyll always occurs bound to proteins. In ring b, chl-b contains a formyl residue instead of the methyl residue in chla. This small difference has a large influence on light absorption. In plants, the ratio chl-a to chl-b is about three to one. Only chl-a is a constituent of the photosynthetic reaction centers and therefore it can be regarded as the central photosynthesis pigment. In a wide range of the visible spectrum, however, chl-a does not absorb light. This nonabsorbing region is named the green window.

Light absorption excites the chlorophyll molecule

1. The most important path for conversion of the energy released when the first singlet state returns to the ground state is its utilization for chemical work. The chlorophyll molecule transfers the excited electron from the first singlet state to an electron acceptor and a positively charged chlorophyll radical chl+ remains. This is possible since the excited electron is bound less strongly to the chromophore molecule than in the ground state.

2. The excited chlorophyll can return to the ground state by releasing excitation energy in the form of light; this light emittance is named fluorescence. Due to vibrations and rotations, part of the excitation energy is usually lost beforehand in the form of heat, with the result that the fluorescence light has less energy (longer in wavelength) than the energy of the excitation light, which was required for attaining the first singlet state .

3. It is also possible that the return from the first singlet to the ground state proceeds in a stepwise fashion via the various levels of vibration and rotation energy, by which the energy difference is completely converted to heat. 4. By releasing part of the excitation energy in the form of heat, the chlorophyll molecule can attain an excited state of lower energy, called the first triplet state. This triplet state cannot be reached directly from the ground state by excitation.

In the triplet state the spin of the excited electrons has been reversed. As the probability of a spin reversal is low, the triplet state does not occur frequently, but in the case of a very high excitation, part of the chlorophyll can reach this state. By emitting so-called phosphorescent light, the molecule can return from the triplet state to the ground state. 5.The return to the ground state can be coupled with the excitation of a neighboring hromophore molecule. This transfer is important for the function of the antennae

efficient photosynthesis is possible only when the energy of photons of various wavelengths is captured over a certain surface by a so called antenna The antennae of plants consist of a large number of protein-bound chlorophyll molecules that absorb photons and transfer their energy to the reaction center. Only a few thousandths of the chlorophyll molecules in the leaf are constituents of the actual reaction centers; the remainder are contained in the antennae.

The results of Emerson and Arnold led to the conclusion that when the quantum requirement is evaluated at eight photons per molecule of O2 formed, and, as was recognized later, two reaction centers require four photons each upon the formation of O2, about 300 chlorophyll molecules are associated with one reaction center. These are constituents of the antennae. The antennae of plants consist of an inner part and an outer part . The outer part, formed by the light harvesting complexes (LHCs), collects the light.

The inner part of the antenna, consisting of the core complexes, is an integral constituent of the reaction centers; it also collects light and conducts the excitons collected in the outer part of the antenna into the photosynthetic reaction centers. The LHCs are formed by polypeptides, which bind chl-a, chl-b, xanthophylls, and carotenes. These proteins, termed LHC polypeptides, are encoded in the nucleus. A plant contains many different LHC polypeptides.

Plants contain two reaction centers, which are arranged in sequence: a reaction center of photosystem II (PS II), which has an absorption maximum at 680 nm, and that of photosystem I (PS I) with an absorption maximum at 700 nm.