Chloroplast

The chemical energy created is then used to make sugar and other organic molecules from carbon dioxide in a process called the Calvin cycle.

Despite this, chloroplasts can be found in extremely diverse organisms that are not directly related to each other—a consequence of many secondary and even tertiary endosymbiotic events.

The first definitive description of a chloroplast (Chlorophyllkörnen, "grain of chlorophyll") was given by Hugo von Mohl in 1837 as discrete bodies within the green plant cell.

[12][32] Chlorarachniophytes are a rare group of organisms that also contain chloroplasts derived from green algae,[29] though their story is more complicated than that of the euglenophytes.

Thiodictyon intracellulare (Chromatiaceae), a purple sulfur bacterium with a genome just half the size of their closest known relatives; and Chlorella sp.

They synthesize ordinary starch, which is stored in granules found in the periplastid space—outside the original double membrane, in the place that corresponds to the ancestral red alga's cytoplasm.

[32] However the diatom endosymbiont can't store its own food—its storage polysaccharide is found in granules in the dinophyte host's cytoplasm instead.

[12][65] The diatom endosymbiont's nucleus is present, but it probably can't be called a nucleomorph because it shows no sign of genome reduction, and might have even been expanded.

[77] While chloroplast genomes can almost always be assembled into a circular map, the physical DNA molecules inside cells take on a variety of linear and branching forms.

[85][86] The results of the microscopy experiments led to the idea that chloroplast DNA replicates using a double displacement loop (D-loop).

[23][119] In addition, in terms of function, the inner chloroplast membrane, which regulates metabolite passage and synthesizes some materials, has no counterpart in the mitochondrion.

[129] The protein-rich,[23] alkaline,[119] aqueous fluid within the inner chloroplast membrane and outside of the thylakoid space is called the stroma,[23] which corresponds to the cytosol of the original cyanobacterium.

[136] Plastoglobuli form when a bubble appears between the layers of the lipid bilayer of the thylakoid membrane, or bud from existing plastoglobuli—though they never detach and float off into the stroma.

[143][144] Thylakoids (sometimes spelled thylakoïds),[146] are small interconnected sacks which contain the membranes that the light reactions of photosynthesis take place on.

[129] Inside the photosystems embedded in chloroplast thylakoid membranes are various photosynthetic pigments, which absorb and transfer light energy.

Chl = chlorophyll[12][155][157] To fix carbon dioxide into sugar molecules in the process of photosynthesis, chloroplasts use an enzyme called RuBisCO.

This is a big problem, since O2 is produced by the initial light reactions of photosynthesis, causing issues down the line in the Calvin cycle which uses RuBisCO.

In mesophyll cells, chloroplasts are specialized for the light reactions, so they lack RuBisCO, and have normal grana and thylakoids,[140] which they use to make ATP and NADPH, as well as oxygen.

[140][159] Because the job of bundle sheath chloroplasts is to carry out the Calvin cycle and make sugar, they often contain large starch grains.

Lower levels of reactive oxygen species initiate systemic acquired resistance, triggering defense-molecule production in the rest of the plant.

[163] One of the main functions of the chloroplast is its role in photosynthesis, the process by which light is transformed into chemical energy, to subsequently produce food in the form of sugars.

Like ATP synthase, ferredoxin-NADP+ reductase, the enzyme that reduces NADP+, releases the NADPH it makes into the stroma, right where it is needed for the dark reactions.

Most of the G3P molecules are recycled back into RuBP using energy from more ATP, but one out of every six produced leaves the cycle—the end product of the dark reactions.

[166] Several mechanisms have evolved in different lineages that raise the carbon dioxide concentration relative to oxygen within the chloroplast, increasing the efficiency of photosynthesis.

[180] Other lipids are derived from the methyl-erythritol phosphate (MEP) pathway and consist of gibberelins, sterols, abscisic acid, phytol, and innumerable secondary metabolites.

[190] Studies of Vallisneria gigantea, an aquatic flowering plant, have shown that chloroplasts can get moving within five minutes of light exposure, though they don't initially show any net directionality.

An etioplast is a plastid that lacks chlorophyll, and has inner membrane invaginations that form a lattice of tubes in their stroma, called a prolamellar body.

[188][196] Late into the constriction phase, dynamin proteins assemble around the outer plastid-dividing ring,[196] helping provide force to squeeze the chloroplast.

There are many other documented mechanisms that prevent paternal inheritance in these flowering plants, such as different rates of chloroplast replication within the embryo.

This makes plastid transformation a valuable tool for the creation and cultivation of genetically modified plants that are biologically contained, thus posing significantly lower environmental risks.

Chloroplasts, containing thylakoids, visible in the cells of Rosulabryum capillare , a type of moss
Cladogram of chloroplast evolution
Possible cladogram of chloroplast evolution [ 29 ] [ 31 ] [ 32 ] Circles represent endosymbiotic events. For clarity, dinophyte tertiary endosymbioses and many nonphotosynthetic lineages have been omitted.
a It is now established that Chromalveolata is paraphyletic to Rhizaria . [ 32 ]
Euglena , a euglenophyte , contains secondary chloroplasts from green algae.
Chlorarachnion reptans is a chlorarachniophyte. Chlorarachniophytes replaced their original red algal endosymbiont with a green alga .
The photosynthetic pigments present in their chloroplasts make diatoms greenish-brown.
Ceratium furca , a peridinin -containing dinophyte [ 63 ]
Karenia brevis is a fucoxanthin -containing dynophyte responsible for algal blooms called " red tides ". [ 53 ]
Dinophysis acuminata has chloroplasts taken from a cryptophyte . [ 29 ]
Chloroplast DNA replication via multiple D-loop mechanisms. Adapted from Krishnan NM, Rao BJ's paper "A comparative approach to elucidate chloroplast genome replication."
Over time, base changes in the DNA sequence can arise from deamination mutations. When adenine is deaminated, it becomes hypoxanthine, which can pair with cytosine. During replication, the cytosine will pair with guanine, causing an A --> G base change.
Transmission electron microscope image of a chloroplast. Grana of thylakoids and their connecting lamellae are clearly visible.
Ribosome Plastoglobule Chloroplast DNA Chloroplast DNA Starch granule Starch granule Starch granule Starch granule Thylakoid space Thylakoid space Thylakoid Granal thylakoid Granal thylakoid Stromal thylakoid Stromal thylakoid Granum Granum Granum Granum Granum Granum Plastoglobule Stroma Inner chloroplast membrane Outer chloroplast membrane
Instead of an intermembrane space, glaucophyte algae have a peptidoglycan wall between their inner and outer chloroplast membranes.
Scanning transmission electron microscope imaging of a chloroplast
(Top) 10-nm-thick STEM tomographic slice of a lettuce chloroplast. Grana stacks are interconnected by unstacked stromal thylakoids, called "stroma lamellae". Round inclusions associated with the thylakoids are plastoglobules. Scalebar=200 nm. See. [ 145 ]
(Bottom) Large-scale 3D model generated from segmentation of tomographic reconstructions by STEM. grana=yellow; stroma lamellae=green; plastoglobules=purple; chloroplast envelope=blue. See. [ 145 ]
Granum-stroma assembly structure The prevailing model of the granum-stroma assembly is stacks of granal thylakoids wrapped by right-handed helical stromal thylakoids which are connected to large parallel sheets of stromal thylakoids and adjacent right-handed helices by left-handed helical structures. (Based on [ 145 ] ).
Proplastid Etioplast Leucoplast Chromoplast Amyloplast Elaioplast Proteinoplast Proplastid Leucoplast Etioplast Chromoplast Amyloplast Elaioplast Proteinoplast Chloroplast Chloroplast File:Plastids types flat.svg
In this light micrograph of some moss chloroplasts, some dumbbell-shaped chloroplasts can be seen dividing. Grana are also just barely visible as small granules.
In this light micrograph of some moss chloroplasts, some dumbbell-shaped chloroplasts can be seen dividing. Grana are also just barely visible as small granules.
In this light micrograph of some moss chloroplasts, some dumbbell-shaped chloroplasts can be seen dividing. Grana are also just barely visible as small granules.
In this light micrograph of some moss chloroplasts, some dumbbell-shaped chloroplasts can be seen dividing. Grana are also just barely visible as small granules.