Plastid

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Plastid
Plant cells with visible
chloroplasts
Scientific classification Edit this classification
Domain: Bacteria
Phylum: Cyanobacteria
Clade: Plastid

A plastid (from

endosymbiotic cyanobacteria.[1]

Examples of plastids include chloroplasts (used for photosynthesis); chromoplasts (used for synthesis and storage of pigments); leucoplasts (non-pigmented plastids some of which can differentiate); and apicoplasts (non-photosynthetic plastids of apicomplexa derived from secondary endosymbiosis).

A permanent primary endosymbiosis event occurred about 1.5 billion years ago in the

amoeboids of the cyanobacteria genera Prochlorococcus and Synechococcus, or the "PS-clade".[4][5] Secondary and tertiary endosymbiosis events have also occurred in a wide variety of organisms; and some organisms developed the capacity to sequester ingested plastids—a process known as kleptoplasty
.

biological pigments such as used in photosynthesis or which determine a cell's color. Plastids in organisms that have lost their photosynthetic properties are highly useful for manufacturing molecules like the isoprenoids.[8]

In land plants

Plastid types
Leucoplasts in plant cells.

Chloroplasts, proplastids, and differentiation

In

chloroplasts
; (see top graphic).

Other plastids can synthesize

mesophyll tissue. Plastids function to store different components including starches, fats, and proteins.[9]

All plastids are derived from proplastids, which are present in the

binary fission
, but more mature chloroplasts also have this capacity.

Plant proplastids (undifferentiated plastids) may differentiate into several forms, depending upon which function they perform in the cell, (see top graphic). They may develop into any of the following variants:[10]

Leucoplasts differentiate into even more specialized plastids, such as:

Depending on their morphology and target function, plastids have the ability to differentiate or redifferentiate between these and other forms.

Plastomes and Chloroplast DNA/ RNA; plastid DNA and plastid nucleoids

Each plastid creates multiple copies of its own unique genome, or

plastome, (from 'plastid genome')—which for a chlorophyll plastid (or chloroplast) is equivalent to a 'chloroplast genome', or a 'chloroplast DNA'.[11][12] The number of genome copies produced per plastid is variable, ranging from 1000 or more in rapidly dividing new cells
, encompassing only a few plastids, down to 100 or less in mature cells, encompassing numerous plastids.

A plastome typically contains a

differention
of plastids.

Many plastids, particularly those responsible for photosynthesis, possess numerous internal membrane layers. Plastid DNA exists as protein-DNA complexes associated as localized regions within the plastid's inner envelope

membrane; and these complexes are called 'plastid nucleoids
'. Unlike the nucleus of a eukaryotic cell, a plastid nucleoid is not surrounded by a nuclear membrane. The region of each nucleoid may contain more than 10 copies of the plastid DNA.

Where the proplastid (undifferentiated plastid) contains a single nucleoid region located near the centre of the proplastid, the developing (or differentiating) plastid has many nucleoids localized at the periphery of the plastid and bound to the inner envelope membrane. During the development/ differentiation of proplastids to chloroplasts—and when plastids are differentiating from one type to another—nucleoids change in morphology, size, and location within the organelle. The remodelling of plastid nucleoids is believed to occur by modifications to the abundance of and the composition of nucleoid proteins.

In normal plant cells long thin protuberances called stromules sometimes form—extending from the plastid body into the cell cytosol while interconnecting several plastids. Proteins and smaller molecules can move around and through the stromules. Comparatively, in the laboratory, most cultured cells—which are large compared to normal plant cells—produce very long and abundant stromules that extend to the cell periphery.

In 2014, evidence was found of the possible loss of plastid genome in Rafflesia lagascae, a non-photosynthetic parasitic flowering plant, and in Polytomella, a genus of non-photosynthetic green algae. Extensive searches for plastid genes in both taxons yielded no results, but concluding that their plastomes are entirely missing is still disputed.[13] Some scientists argue that plastid genome loss is unlikely since even these non-photosynthetic plastids contain genes necessary to complete various biosynthetic pathways including heme biosynthesis.[13][14]

Even with any loss of plastid genome in Rafflesiaceae, the plastids still occur there as "shells" without DNA content,[15] which is reminiscent of hydrogenosomes in various organisms.

In algae and protists

Plastid types in algae and protists include:

The plastid of photosynthetic Paulinella species is often referred to as the 'cyanelle' or chromatophore, and is used in photosynthesis.[17][18] It had a much more recent endosymbiotic event, in the range of 140–90 million years ago, which is the only other known primary endosymbiosis event of cyanobacteria.[19][20]

Etioplasts, amyloplasts and chromoplasts are plant-specific and do not occur in algae.[citation needed] Plastids in algae and hornworts may also differ from plant plastids in that they contain pyrenoids.

Inheritance

In reproducing, most plants inherit their plastids from only one parent. In general,

gymnosperms inherit plastids from the male pollen
. Algae also inherit plastids from just one parent. Thus the plastid DNA of the other parent is completely lost.

In normal intraspecific crossings—resulting in normal hybrids of one species—the inheriting of plastid DNA appears to be strictly uniparental; i.e., from the female. In interspecific hybridisations, however, the inheriting is apparently more erratic. Although plastids are inherited mainly from the female in interspecific hybridisations, there are many reports of hybrids of flowering plants producing plastids from the male. Approximately 20% of angiosperms, including alfalfa (Medicago sativa), normally show biparental inheriting of plastids.[21]

DNA damage and repair

The plastid DNA of maize seedlings is subjected to increasing damage as the seedlings develop.[22] The DNA damage is due to oxidative environments created by photo-oxidative reactions and photosynthetic/ respiratory electron transfer. Some DNA molecules are repaired but DNA with unrepaired damage is apparently degraded to non-functional fragments.

nuclear genome and then translocated to plastids where they maintain genome stability/ integrity by repairing the plastid's DNA.[23] For example, in chloroplasts of the moss Physcomitrella patens, a protein employed in DNA mismatch repair (Msh1) interacts with proteins employed in recombinational repair (RecA and RecG) to maintain plastid genome stability.[24]

Origin

Plastids are thought to be descended from

light harvesting complexes found in cyanobacteria, red algae and glaucophytes, but instead contain stroma and grana thylakoids
. The glaucocystophycean plastid—in contrast to chloroplasts and rhodoplasts—is still surrounded by the remains of the cyanobacterial cell wall. All these primary plastids are surrounded by two membranes.

The plastid of photosynthetic Paulinella species is often referred to as the 'cyanelle' or chromatophore, and had a much more recent endosymbiotic event about 90–140 million years ago; it is the only known primary endosymbiosis event of cyanobacteria outside of the Archaeplastida.[17][18] The plastid belongs to the "PS-clade" (of the cyanobacteria genera Prochlorococcus and Synechococcus), which is a different sister clade to the plastids belonging to the Archaeplastida.[4][5]

In contrast to primary plastids derived from primary endosymbiosis of a prokaryoctyic cyanobacteria, complex plastids originated by secondary

antiparasitic drug
development.

Some

dinoflagellates and sea slugs, in particular of the genus Elysia, take up algae as food and keep the plastid of the digested alga to profit from the photosynthesis; after a while, the plastids are also digested. This process is known as kleptoplasty, from the Greek, kleptes (κλέπτης
), thief.

Plastid development cycle

An illustration of the stages of inter-conversion in plastids

In 1977 J.M Whatley proposed a plastid development cycle which said that plastid development is not always unidirectional but is instead a complicated cyclic process. Proplastids are the precursor of the more differentiated forms of plastids, as shown in the diagram to the right.[28]

See also

  • Mitochondrion – Organelle in eukaryotic cells responsible for respiration
  • Cytoskeleton – Network of filamentous proteins that forms the internal framework of cells
  • Photosymbiosis – A type of symbiotic relationship where one organism is capable of photosynthesis

Notes

  1. ^ Sometimes Ernst Haeckel is credited to coin the term plastid, but his "plastid" includes nucleated cells and anucleated "cytodes"[7] and thus totally different concept from the plastid in modern literature.

References

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  6. ^ Schimper, A.F.W. (1882) "Ueber die Gestalten der Stärkebildner und Farbkörper" Botanisches Centralblatt 12(5): 175–178.
  7. ^ Haeckel, E. (1866) "Morphologische Individuen erster Ordnung: Plastiden oder Plasmastücke" in his Generelle Morphologie der Organismen Bd. 1, pp. 269-289
  8. ^ Picozoans Are Algae After All: Study | The Scientist Magazine®
  9. ^ Kolattukudy, P.E. (1996) "Biosynthetic pathways of cutin and waxes, and their sensitivity to environmental stresses", pp. 83–108 in: Plant Cuticles. G. Kerstiens (ed.), BIOS Scientific publishers Ltd., Oxford
  10. ^ .
  11. .
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  13. ^ a b "Plants Without Plastid Genomes". The Scientist. Retrieved 2015-09-26.
  14. PMID 16406301
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  15. ^ "DNA of Giant 'Corpse Flower' Parasite Surprises Biologists". April 2021.
  16. PMID 11429143
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  27. ^ Chan CX, Bhattachary D (2010). "The Origin of Plastids". Nature Education. 3 (9): 84.
  28. JSTOR 2431207
    .

Further reading

External links