Plastid

starch and can synthesize fatty acids and terpenes, which

can be used for producing energy and as raw material
for the synthesis of other molecules. For example, the
components of the plant cuticle and its epicuticular wax
are synthesized by the epidermal cells from palmitic acid,
which is synthesized in the chloroplasts of the mesophyll
tissue.[2] All plastids are derived from proplastids, which
are present in the meristematic regions of the plant. Pro-
plastids and young chloroplasts commonly divide by bi-
nary ssion, but more mature chloroplasts also have this
capacity.

Plant cells with visible chloroplasts.

The plastid (Greek: ; plasts: formed,

molded plural plastids) is a major double-membrane
organelle[1] found in the cells of plants, algae, and some
other eukaryotic organisms. Plastids are the site of man-
ufacture and storage of important chemical compounds
used by the cell. They often contain pigments used in
photosynthesis, and the types of pigments present can
change or determine the cells color. They have a com-
mon evolutionary origin and possess a double-stranded
DNA molecule that is circular, like that of prokaryotic
cells.

In plants, plastids may dierentiate into several forms, de-

1 Plastids in plants pending upon which function they play in the cell. Undif-
ferentiated plastids (proplastids) may develop into any of
the following variants:[3]

Chloroplasts green plastids: for photosynthesis; see

also etioplasts, the predecessors of chloroplasts
Chromoplasts coloured plastids: for pigment syn-
thesis and storage
Gerontoplasts: control the dismantling of the pho-
tosynthetic apparatus during plant senescence
Leucoplasts colourless plastids: for monoterpene
synthesis; leucoplasts sometimes dierentiate into
more specialized plastids:
Amyloplasts: for starch storage and detecting
gravity
Leucoplasts in plant cells.
Elaioplasts: for storing fat
Those plastids that contain chlorophyll can carry out Proteinoplasts: for storing and modifying
photosynthesis. Plastids can also store products like protein

1
2 4 ORIGIN OF PLASTIDS

Tannosomes: for synthesizing and producing 2 Plastids in algae

tannins and polyphenols
In algae, the term leucoplast is used for all unpigmented
Depending on their morphology and function, plastids plastids and their function diers from the leucoplasts
have the ability to dierentiate, or redierentiate, be- of plants. Etioplasts, amyloplasts and chromoplasts are
tween these and other forms. plant-specic and do not occur in algae. Plastids in algae
and hornworts may also dier from plant plastids in that
Each plastid creates multiple copies of a circular 75
they contain pyrenoids.
250 kilobase plastome. The number of genome copies
per plastid is variable, ranging from more than 1000 in Glaucocystophytic algae contain muroplasts, which
rapidly dividing cells, which, in general, contain few plas- are similar to chloroplasts except that they have a
tids, to 100 or fewer in mature cells, where plastid divi- peptidoglycan cell wall that is similar to that of
sions have given rise to a large number of plastids. The prokaryotes. Rhodophytic algae contain rhodoplasts,
plastome contains about 100 genes encoding ribosomal which are red chloroplasts that allow the algae to photo-
[3]
and transfer ribonucleic acids (rRNAs and tRNAs) as synthesise to a depth of up to 268 m. The chloroplasts
well as proteins involved in photosynthesis and plastid of plants dier from the rhodoplasts of red algae in their
gene transcription and translation. However, these pro- ability to synthesize starch, which is stored in the form
teins only represent a small fraction of the total protein of granules within the plastids. In red algae, oridean
set-up necessary to build and maintain the structure and starch is synthesized and stored outside the plastids in the
[6]
function of a particular type of plastid. Plant nuclear cytosol.
genes encode the vast majority of plastid proteins, and the
expression of plastid genes and nuclear genes is tightly co-
regulated to coordinate proper development of plastids in 3 Inheritance of plastids
relation to cell dierentiation.
Plastid DNA exists as large protein-DNA complexes as- Most plants inherit the plastids from only one parent. In
sociated with the inner envelope membrane and called general, angiosperms inherit plastids from the female ga-
'plastid nucleoids. Each nucleoid particle may contain mete, whereas many gymnosperms inherit plastids from
more than 10 copies of the plastid DNA. The proplastid the male pollen. Algae also inherit plastids from only one
contains a single nucleoid located in the centre of the plas- parent. The plastid DNA of the other parent is, thus,
tid. The developing plastid has many nucleoids, localized completely lost.
at the periphery of the plastid, bound to the inner enve-
In normal intraspecic crossings (resulting in normal hy-
lope membrane. During the development of proplastids
brids of one species), the inheritance of plastid DNA
to chloroplasts, and when plastids convert from one type
appears to be quite strictly 100% uniparental. In inter-
to another, nucleoids change in morphology, size and lo-
specic hybridisations, however, the inheritance of plas-
cation within the organelle. The remodelling of nucleoids
tids appears to be more erratic. Although plastids inherit
is believed to occur by modications to the composition
mainly maternally in interspecic hybridisations, there
and abundance of nucleoid proteins.
are many reports of hybrids of owering plants that con-
Many plastids, particularly those responsible for photo- tain plastids of the father. Approximately 20% of an-
synthesis, possess numerous internal membrane layers. giosperms, including alfalfa (Medicago sativa), normally
In plant cells, long thin protuberances called stromules show biparental inheritance of plastids.[7]
sometimes form and extend from the main plastid body
into the cytosol and interconnect several plastids. Pro-
teins, and presumably smaller molecules, can move within 4 Origin of plastids
stromules. Most cultured cells that are relatively large
compared to other plant cells have very long and abun- Plastids are thought to have originated from
dant stromules that extend to the cell periphery. endosymbiotic cyanobacteria. This symbiosis evolved
In 2014, evidence of possible plastid genome loss around 1.5 billion years ago[8] and enabled eukaryotes
was found in Raesia lagascae, a non-photosynthetic to carry out oxygenic photosynthesis.[9] Three evo-
parasitic owering plant, and in Polytomella, a genus of lutionary lineages have since emerged in which the
non-photosynthetic green algae. Extensive searches for plastids are named dierently: chloroplasts in green
plastid genes in both Raesia and Polytomella yielded algae and plants, rhodoplasts in red algae and muroplasts
no results, however the conclusion that their plastomes in the glaucophytes. The plastids dier both in their
are entirely missing is still controversial.[4] Some scien- pigmentation and in their ultrastructure. For example,
tists argue that plastid genome loss is unlikely since even chloroplasts have lost all phycobilisomes, the light
non-photosynthetic plastids contain genes necessary to harvesting complexes found in cyanobacteria, red algae
complete various biosynthetic pathways, such as heme and glaucophytes, but instead contain stroma and grana
biosynthesis.[4][5] thylakoids, structures found only in plants and closely

related green algae. 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.
Complex plastids start by secondary endosymbiosis
(where a eukaryotic organism engulfs another eukary-
otic organism that contains a primary plastid resulting
in its endosymbiotic xation),[10] when a eukaryote en-
gulfs a red or green alga and retains the algal plas-
tid, which is typically surrounded by more than two
membranes. In some cases these plastids may be re-
duced in their metabolic and/or photosynthetic capac-
ity. Algae with complex plastids derived by secondary
endosymbiosis of a red alga include the heterokonts,
haptophytes, cryptomonads, and most dinoagellates (=
rhodoplasts). Those that endosymbiosed a green alga in-
clude the euglenids and chlorarachniophytes (= chloro-
plasts). The Apicomplexa, a phylum of obligate para-
sitic 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 or-
ganelle, and a promising target for antiparasitic drug de-
velopment.
Some dinoagellates and sea slugs, in particular of the [10] Chan, C. X. & Bhattachary, D. (2010). The Origin of
genus Elysia, take up algae as food and keep the plastid of
the digested alga to prot from the photosynthesis; after
a while, the plastids are also digested. This process is
known as kleptoplasty, from the Greek, kleptes, thief.
5 See also
Mitochondrion
6 References
[1] Sato, N. (2006). Origin and Evolution of Plastids: Ge-
nomic View on the Unication and Diversity of Plas-
tids. In R.R. Wise; J.K. Hoober. The Structure and
Function of Plastids. 23. Springer Netherlands. pp.
75102. ISBN 978-1-4020-4060-3. doi:10.1007/978-1-
4020-4061-0_4.
[2] 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
Scientic publishers Ltd., Oxford
[3] Wise, Robert R. (2006). 1. The Diversity of Plas-
tid Form and Function. Advances in Photosynthesis
and Respiration (PDF). 23. Springer. pp. 326.
doi:10.1007/978-1-4020-4061-0_1.
3
[4] Plants Without Plastid Genomes | The Scientist Maga-
zine". The Scientist. Retrieved 2015-09-26.
[5] Barbrook, Adrian C.; Howe, Christopher J.; Pur-
ton, Saul (2006). Why are plastid genomes re-
tained in non-photosynthetic organisms?". Trends in
Plant Science. 11 (2): 101108. PMID 16406301.
doi:10.1016/j.tplants.2005.12.004.
[6] Viola, R.; Nyvall, P.; Pedersn, M. (2001). The
unique features of starch metabolism in red algae.
Proceedings of the Royal society of London, B. 268:
14171422. PMC 1088757 . PMID 11429143.
doi:10.1098/rspb.2001.1644.
[7] Zhang, Q.; Sodmergen (2010). Why does biparental
plastid inheritance revive in angiosperms?". Journal of
Plant Research. 123 (2): 201206. PMID 20052516.
doi:10.1007/s10265-009-0291-z.
[8] Ochoa De Alda, Jess A. G.; Esteban, Roco; Di-
ago, Mara Luz; Houmard, Jean (2014). The
plastid ancestor originated among one of the ma-
jor cyanobacterial lineages. Nature Communications.
5: 4937. Bibcode:2014NatCo...5E4937O. PMID
25222494. doi:10.1038/ncomms5937.
[9] Hedges SB, Blair JE, Venturi ML, Shoe JL (January
2004). A molecular timescale of eukaryote evolu-
tion and the rise of complex multicellular life. BMC
Evol. Biol. 4: 2. PMC 341452 . PMID 15005799.
doi:10.1186/1471-2148-4-2.
Plastids. Nature Education. 3 (9): 84.
7 Sources
A Novel View of Chloroplast Structure: contains
uorescence images of chloroplasts and stromules as
well as an easy to read chapter.
Wyclie P, Sitbon F, Wernersson J, Ezcurra I, Eller-
strm M, Rask L (October 2005). Continuous
expression in tobacco leaves of a Brassica napus
PEND homologue blocks dierentiation of plas-
tids and development of palisade cells. Plant J. 44
(1): 115. PMID 16167891. doi:10.1111/j.1365-
313X.2005.02482.x.
Birky CW (2001). The inheritance of
genes in mitochondria and chloroplasts: laws,
mechanisms, and models. Annu. Rev.
Genet. 35: 12548. PMID 11700280.
doi:10.1146/annurev.genet.35.102401.090231.
PDF
8 Further reading
Chan CX, Bhattacharya D (2010). The origins of
plastids. Nature Education. 3 (9): 84.
4 9 EXTERNAL LINKS

Bhattacharya, D., ed. (1997). Origins of Algae and

their Plastids. New York: Springer-Verlag/Wein.
ISBN 3-211-83036-7.

Gould SB, Waller RR, McFadden GI (2008). Plas-

tid evolution. Annu Rev Plant Biol 59: 491517.

Keeling P (2010). The endosymbiotic origin, diver-

sication and fate of plastids. Philos Trans R Soc
Lond B Biol Sci. 365(1541): 729-48

9 External links
Transplastomic plants for biocontainment (biologi-
cal connement of transgenes) Co-extra research
project on coexistence and traceability of GM and
non-GM supply chains

Tree of Life Eukaryotes

 5

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