Plastid

The plastid (Greek: πλαστός; plastós:

formed, molded – plural plastids) is a
membrane-bound organelle[1] found in
the cells of plants, algae, and some other
eukaryotic organisms. They are
considered endosymbiotic
Cyanobacteria, related to the
Gloeomargarita.[2] Plastids were
discovered and named by Ernst Haeckel,
but A. F. W. Schimper was the first to
provide a clear definition. Plastids are the
site of manufacture and storage of
important chemical compounds used by
the cells of autotrophic eukaryotes. They
often contain pigments used in
photosynthesis, and the types of
pigments in a plastid determine the cell's
color. They have a common evolutionary
origin and possess a double-stranded
DNA molecule that is circular, like that of
the circular chromosome of prokaryotic
cells.

Plastid

Scientific classification

Domain: Bacteria

Phylum: Cyanobacteria

Clade: Plastids
Plant cells with visible chloroplasts.

In plants

Leucoplasts in plant cells.

Plastids that contain chlorophyll can
carry out photosynthesis and are called
chloroplasts. Plastids can also store
products like 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.[3] All plastids are
derived from proplastids, which are
present in the meristematic regions of
the plant. Proplastids and young
chloroplasts commonly divide by binary
fission, but more mature chloroplasts
also have this capacity.

In plants,
undifferentiated
plastids
(proplastids) may
differentiate into
several forms, depending upon which
function they perform in the cell. They
may develop into any of the following
variants:[4]

Chloroplasts: green plastids for

photosynthesis; see also etioplasts, the
predecessors of chloroplasts
Chromoplasts: coloured plastids for
pigment synthesis and storage
Gerontoplasts: control the dismantling
of the photosynthetic apparatus during
plant senescence
Leucoplasts: colourless plastids for
monoterpene synthesis; leucoplasts
sometimes differentiate into more
specialized plastids:
Amyloplasts: for starch storage
and detecting gravity (for
geotropism)
Elaioplasts: for storing fat
Proteinoplasts: for storing and
modifying protein
 Tannosomes: for synthesizing and
producing tannins and
polyphenols

Depending on their morphology and

function, plastids have the ability to
differentiate, or redifferentiate, between
these and other forms.

Each plastid creates multiple copies of a

circular 75–250 kilobase plastome. The
number of genome copies per plastid is
variable, ranging from more than 1000 in
rapidly dividing cells, which, in general,
contain few plastids, to 100 or fewer in
mature cells, where plastid divisions
have 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. Plant nuclear 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 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.
In 2014, evidence of possible plastid
genome loss was found 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 Rafflesia and Polytomella
yielded no results, however the
conclusion that their plastomes are
entirely missing is still controversial.[5]
Some scientists argue that plastid
genome loss is unlikely since even non-
photosynthetic plastids contain genes
necessary to complete various
biosynthetic pathways, such as heme
biosynthesis.[5][6]
In algae
In algae, the term leucoplast is used for
all unpigmented plastids. Their function
differs from the leucoplasts of plants.
Etioplasts, amyloplasts and
chromoplasts 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.

Glaucophyte algae contain muroplasts,

which are similar to chloroplasts except
that they have a peptidoglycan cell wall
that is similar to that of prokaryotes. Red
algae contain rhodoplasts, which are red
chloroplasts that allow them to
photosynthesize to a depth of up to 268
m.[4] The chloroplasts of plants differ
from the rhodoplasts of red algae in their
ability to synthesize starch, which is
stored in the form of granules within the
plastids. In red algae, floridean starch is
synthesized and stored outside the
plastids in the cytosol.[7]

Inheritance
Most plants inherit the plastids from only
one parent. In general, angiosperms
inherit plastids from the female gamete,
whereas 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. Approximately 20% of
angiosperms, including alfalfa (Medicago
sativa), normally show biparental
inheritance of plastids.[8]

DNA damage and repair

Plastid DNA of maize seedlings is
subject to increased damage as the
seedlings develop.[9] The DNA is
damaged in oxidative environments
created by photo-oxidative reactions and
photosynthetic/respiratory electron
transfer. Some DNA molecules are
repaired while DNA with unrepaired
damage appears to be degraded to non-
functional fragments.

DNA repair proteins are encoded by the

cell's nuclear genome but can be
translocated to plastids where they
maintain genome stability/integrity by
repairing the plastid's DNA.[10] As an
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.[11]

Origin
Plastids are thought to be endosymbiotic
cyanobacteria. This primary
endosymbiotic event is hypothesized to
have occurred around 1.5 billion years
ago[12] and enabled eukaryotes to carry
out oxygenic photosynthesis.[13] Three
evolutionary lineages have since
emerged in which the plastids are named
differently: chloroplasts in green algae
and plants, rhodoplasts in red algae and
muroplasts in the glaucophytes. The
plastids differ both in their pigmentation
and in their ultrastructure. For example,
chloroplasts in plants and green algae
have lost all phycobilisomes, the 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.

In contrast to primary plastids derived

from primary endosymbiosis of a
prokaryoctyic cyanobacteria, complex
plastids originated by secondary
endosymbiosis in which a eukaryotic
organism engulfed another eukaryotic
organism that contained a primary
plastid. [14] When a eukaryote engulfs a
red or a green alga and retains the algal
plastid, that plastid 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, 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,
thief.

Paulinella chromatophora
Paulinella has a similar organelle which
does not belong to the plastids
discussed in this text. It is a
chromatophore, which is a more recently
acquired endosymbiotic β-
Cyanobacteria.[15] This endosymbiosis
event occurred much more recent, and is
only the second known primary
endosymbiosis event of cyanobacteria.

See also
Mitochondrion

References
1. Sato N (2006). "Origin and Evolution
of Plastids: Genomic View on the
 Unification and Diversity of Plastids".
In Wise RR, Hoober JK (eds.). The
Structure and Function of Plastids.
Advances in Photosynthesis and
Respiration. 23. Springer
Netherlands. pp. 75–102.
doi:10.1007/978-1-4020-4061-0_4 .
ISBN 978-1-4020-4060-3.
2. Moore KR, Magnabosco C, Momper
L, Gold DA, Bosak T, Fournier GP
(2019). "An Expanded Ribosomal
Phylogeny of Cyanobacteria
Supports a Deep Placement of
Plastids" . Frontiers in Microbiology.
10: 1612.
doi:10.3389/fmicb.2019.01612 .
PMC 6640209 . PMID 31354692 .
3. 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
4. Wise, Robert R. (2006). "1. The
Diversity of Plastid Form and
Function". Advances in
Photosynthesis and Respiration. 23.
Springer. pp. 3–26. doi:10.1007/978-
1-4020-4061-0_1 . ISBN 978-1-4020-
4060-3.
5. "Plants Without Plastid Genomes" .
The Scientist. Retrieved 2015-09-26.
 . Barbrook AC, Howe CJ, Purton S
(February 2006). "Why are plastid
genomes retained in non-
photosynthetic organisms?". Trends
in Plant Science. 11 (2): 101–8.
doi:10.1016/j.tplants.2005.12.004 .
PMID 16406301 .
7. Viola R, Nyvall P, Pedersén M (July
2001). "The unique features of starch
metabolism in red algae" .
Proceedings. Biological Sciences.
268 (1474): 1417–22.
doi:10.1098/rspb.2001.1644 .
PMC 1088757 . PMID 11429143 .
. Zhang Q (March 2010). "Why does
biparental plastid inheritance revive
 in angiosperms?". Journal of Plant
Research. 123 (2): 201–6.
doi:10.1007/s10265-009-0291-z .
PMID 20052516 .
9. Kumar RA, Oldenburg DJ, Bendich AJ
(December 2014). "Changes in DNA
damage, molecular integrity, and
copy number for plastid DNA and
mitochondrial DNA during maize
development" . Journal of
Experimental Botany. 65 (22): 6425–
39. doi:10.1093/jxb/eru359 .
PMC 4246179 . PMID 25261192 .
10. Oldenburg DJ, Bendich AJ (2015).
"DNA maintenance in plastids and
mitochondria of plants" . Frontiers in
 Plant Science. 6: 883.
doi:10.3389/fpls.2015.00883 .
PMC 4624840 . PMID 26579143 .
11. Odahara M, Kishita Y, Sekine Y
(August 2017). "MSH1 maintains
organelle genome stability and
genetically interacts with RECA and
RECG in the moss Physcomitrella
patens". The Plant Journal. 91 (3):
455–465. doi:10.1111/tpj.13573 .
PMID 28407383 .
12. Ochoa de Alda JA, Esteban R, Diago
ML, Houmard J (September 2014).
"The plastid ancestor originated
among one of the major
cyanobacterial lineages". Nature
 Communications. 5: 4937.
Bibcode:2014NatCo...5.4937O .
doi:10.1038/ncomms5937 .
PMID 25222494 .
13. Hedges SB, Blair JE, Venturi ML,
Shoe JL (January 2004). "A
molecular timescale of eukaryote
evolution and the rise of complex
multicellular life" . BMC Evolutionary
Biology. 4: 2. doi:10.1186/1471-
2148-4-2 . PMC 341452 .
PMID 15005799 .
14. Chan CX, Bhattachary D (2010). "The
Origin of Plastids" . Nature
Education. 3 (9): 84.
15. Marin, Birger; Nowack, Eva CM;
Glöckner, Gernot; Melkonian, Michael
(2007). "The ancestor of the
Paulinella chromatophore obtained a
carboxysomal operon by horizontal
gene transfer from a Nitrococcus-like
γ-proteobacterium" . BMC
Evolutionary Biology. 7 (1): 85.
doi:10.1186/1471-2148-7-85 .
PMC 1904183 . PMID 17550603 .

Further reading
Hanson MR, Köhler RH. "A Novel View of
Chloroplast Structure" . Plant Physiology
Online. Archived from the original on 2005-
06-14.
Wycliffe P, Sitbon F, Wernersson J, Ezcurra I,
Ellerström M, Rask L (October 2005).
"Continuous expression in tobacco leaves
of a Brassica napus PEND homologue
blocks differentiation of plastids and
development of palisade cells". The Plant
Journal. 44 (1): 1–15. doi:10.1111/j.1365-
313X.2005.02482.x . PMID 16167891 .
Birky CW (2001). "The inheritance of genes
in mitochondria and chloroplasts: laws,
mechanisms, and models" (PDF). Annual
Review of Genetics. 35: 125–48.
doi:10.1146/annurev.genet.35.102401.090
231 . PMID 11700280 . Archived from the
original (PDF) on 2010-06-22. Retrieved
2009-03-01.
Chan CX, Bhattacharya D (2010). "The
origins of plastids" . Nature Education. 3
 (9): 84.
Bhattacharya D, ed. (1997). Origins of Algae
and their Plastids. New York: Springer-
Verlag/Wein. ISBN 978-3-211-83036-9.
Gould SB, Waller RF, McFadden GI (2008).
"Plastid evolution" . Annual Review of Plant
Biology. 59: 491–517.
doi:10.1146/annurev.arplant.59.032607.09
2915 . PMID 18315522 .
Keeling PJ (March 2010). "The
endosymbiotic origin, diversification and
fate of plastids" . Philosophical
Transactions of the Royal Society of London.
Series B, Biological Sciences. 365 (1541):
729–48. doi:10.1098/rstb.2009.0103 .
PMC 2817223 . PMID 20124341 .

External links
 Transplastomic plants for
biocontainment (biological
confinement of transgenes) — Co-
extra research project on coexistence
and traceability of GM and non-GM
supply chains
Tree of Life Eukaryotes

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