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Plant cells with visible chloroplasts.

Plastids are major organelles found in the cells of plants and algae. Plastids are the site of manufacture and storage of important chemical compounds used by the cell. Plastids often contain pigments used in photosynthesis, and the types of pigments present can change or determine the cell's colour.

Contents

Plastids in plants

Plastids are responsible for photosynthesis, storage of products like starch and for the synthesis have the ability to differentiate, or redifferentiate, between these and other forms. All plastids are derived from proplastids (formerly "eoplasts", eo-: dawn, early), which are present in the meristematic regions of the plant. Proplastids and young chloroplasts commonly divide, but more mature chloroplasts also have this capacity.

Plastids types en.svg

In plants, plastids may differentiate into several forms, depending upon which function they need to play in the cell. Undifferentiated plastids (proplastids) may develop into any of the following plastids:


Each plastid creates multiple copies of the circular 75-250 kilo bases plastome. The number of genome copies per plastid is flexible, ranging from more than 1000 in rapidly dividing cells, which generally contain few plastids, to 100 or fewer in mature cells, where plastid divisions has given rise to a large number of plastids. The plastome contains about 100 genes encoding ribosomal and transfer ribonucleic acids (rRNAs and tRNAs) as well as proteins involved in photosynthesis and plastid gene transcription and translation. However, these proteins only represent a small fraction of the total protein set-up necessary to build and maintain the structure and function of a particular type of plastid. Nuclear genes encode the vast majority of plastid proteins, and the expression of plastid genes and nuclear genes is tightly co-regulated to allow proper development of plastids in relation to cell differentiation.

Plastid DNA exists as large protein-DNA complexes associated with the inner envelope membrane and called 'plastid nucleoids'. Each nucleoid particle may contain more than 10 copies of the plastid DNA. The proplastid contains a single nucleoid located in the centre of the plastid. The developing plastid has many nucleoids, localized at the periphery of the plastid, bound to the inner envelope membrane. During the development of proplastids to chloroplasts, and when plastids convert from one type to another, nucleoids change in morphology, size and location within the organelle. The remodelling of nucleoids is believed to occur by modifications to the composition and abundance of nucleoid proteins.

Many plastids, particularly those responsible for photosynthesis, possess numerous internal membrane layers.

In plant cells long thin protuberances called stromules sometimes form and extend from the main plastid body into the cytosol and interconnect several plastids. Proteins, and presumably smaller molecules, can move within stromules. Most cultured cells that are relatively large compared to other plant cells have very long and abundant stromules that extend to the cell periphery.

Plastids in algae

In algae, the term leucoplast (leukoplast) is used for all unpigmented plastids. Their function differ from the leukoplasts in plants. Etioplast, amyloplast and chromoplast are plant-specific and do not occur in algae. Plastids in algae and hornworts may also differ from plant plastids in that they contain pyrenoids.

Inheritance of plastids

Most plants inherit the plastids from only one parent. Angiosperms generally inherit plastids from the female gamete, while many gymnosperms inherit plastids from the male pollen. Algae also inherit plastids from only one parent. The plastid DNA of the other parent is thus completely lost.

In normal intraspecific crossings (resulting in normal hybrids of one species), the inheritance of plastid DNA appears to be quite strictly 100% uniparental. In interspecific hybridisations, however, the inheritance of plastids appears to be more erratic. Although plastids inherit mainly maternally in interspecific hybridisations, there are many reports of hybrids of flowering plants that contain plastids of the father.

Origin of plastids

Plastids are thought to have originated from endosymbiotic cyanobacteria. They developed around 1500 million years ago and allowed eukaryotes to carry out oxygenic photosynthesis.[1] Due to a split-up into three evolutionary lineages, the plastids are named differently: chloroplasts in green algae and plants, rhodoplasts in red algae and cyanelles in the glaucophytes. The plastids differ by their pigmentation, but also in ultrastructure. The chloroplasts e.g. have lost all phycobilisomes, the light harvesting complexes found in cyanobacteria, red algae and glaucophytes, but - only in plants and in closely related green algae - contain stroma and grana thylakoids. The glaucocystophycean plastid - in contrast to the chloroplasts and the rhodoplasts - is still surrounded by the remains of the cyanobacterial cell wall. All these primary plastids are surrounded by two membranes.

Complex plastids start by secondary endosymbiosis, when a eukaryote engulfs a red or green alga and retains the algal plastid, which is typically surrounded by more than two membranes. In some cases these plastids may be reduced in their metabolic and/or photosynthetic capacity. Algae with complex plastids derived by secondary endosymbiosis of a red alga include the heterokonts, haptophytes, cryptomonads, and most dinoflagellates (= rhodoplasts). Those that endosymbiosed a green alga include the euglenids and chlorarachniophytes (= chloroplasts). The Apicomplexa, a phylum of obligate parasitic protozoa including the causative agents of malaria (Plasmodium spp.), toxoplasmosis (Toxoplasma gondii), and many other human or animal diseases also harbor a complex plastid (although this organelle has been lost in some apicomplexans, such as Cryptosporidium parvum, which causes cryptosporidiosis). The 'apicoplast' is no longer capable of photosynthesis, but is an essential organelle, and a promising target for antiparasitic drug development.

Some dinoflagellates and sea slugs, particularly 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. These captured plastids are known as kleptoplastids.

Sources

  • Continuous expression in tobacco leaves of a Brassica napus PEND homologue blocks differentiation of plastids and development of palisade cells Wycliffe et al., 2005. The Plant Journal Volume 44 Issue 1 Page 1. PMID: 16167891
  • Birky, C. W. 2001. The inheritance of genes in mitochondria and chloroplasts: laws, mechanisms and models. Annual Review of Genetics 35: 125-148. URL

See also

Mitochondrion

References

  1. ^ Hedges, S. Blair et al. (2004) "A molecular timescale of eukaryote evolution and the rise of complex multicellular life" BMC Evolutionary Biology 4:2

Further reading

  • Bhattacharya, D. (Ed.) 1997 Origins of Algae and their Plastids. Springer-Verlag/Wein, New York. ISBN 3-211-83036-7
  • Tree of Life Eukaryotes

External links


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Simple English

A plastid is a self-reproducing organelle of plants and algae. A plastome is the DNA genome of a plastid.[1]p341

Examples of plastids are:

  • Chloroplasts: photosynthesis; other plastids may have developed from chloroplasts.
  • Chromoplasts: pigment synthesis and storage.
  • Leucoplasts: make terpenes such as resin.
    • Elaioplasts: fat storage.
    • Proteinoplasts: store & modify protein.
    • Amyloplasts: starch storage

Genetics and evolution

File:Plagiomnium affine
Chloroplasts visible in the cells of Plagiomnium affine

Plastids are one of the many different types of organelles in the cell. In endosymbiont theory, plastids originated as cyanobacteria. This was first suggested by Mereschkowsky in 1905[2][3] after an observation by Schimper in 1883 that chloroplasts closely resemble cyanobacteria.[4]

Almost all chloroplasts are thought to derive directly or indirectly from a single endosymbiotic event.[5] Mitochondria also derive from symbiosis, but chloroplasts are found only in plants and protista. The chloroplast is surrounded by a double-layered composite membrane with an intermembrane space; further, it has reticulations, or many infoldings, filling the inner spaces. The chloroplast has its own DNA, which codes for redox proteins involved in electron transport in photosynthesis.[6]

References

  1. King R.C. Stansfield W.D. & Mulligan P.K. 2006. A dictionary of genetics, 7th ed. Oxford.
  2. Mereschkowsky C (1905). [Expression error: Unexpected < operator "Über Natur und Ursprung der Chromatophoren im Pflanzenreiche"]. Biol Centralbl 25: 593–604. 
  3. Khakhina L.N. 1992. Concepts of symbiogenesis: a historical and critical survey of the research of Russian botanists. Yale, New Haven CN.
  4. Schimper AFW (1883). [Expression error: Unexpected < operator "Über die Entwicklung der Chlorophyllkörner und Farbkörper"]. Bot. Zeitung 41: 105–14, 121–31, 137–46, 153–62. 
  5. Patrick J. Keeling (2004). "Diversity and evolutionary history of plastids and their hosts". American Journal of Botany 91: 1481–1493. doi:10.3732/ajb.91.10.1481. http://www.amjbot.org/cgi/content/full/91/10/z1481. 
  6. Krause K (September 2008). [Expression error: Unexpected < operator "From chloroplasts to "cryptic" plastids: evolution of plastid genomes in parasitic plants"]. Curr. Genet. 54 (3): 111–21. doi:10.1007/s00294-008-0208-8. PMID 18696071. 

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