Plastids
The
plastid (Greek: πλαστός; plastós: formed, molded – plural plastids) is a major
organelle found in the cells of plants and algae. Plastids are the site of
manufacture 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 cell's color. They possess a
double-stranded DNA molecule, which is circular, like that of prokaryotes.
Plastids in plants
Those
plastids which contain pigments can carry out photosynthesis. Plastids can also
store products like starch and can synthesise 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.[1] 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.
Plant cells with visible chloroplasts
In
plants, plastids may differentiate into several forms, depending upon which
function they play in the cell. Undifferentiated plastids (proplastids) may
develop into any of the following variants.
- 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 senescence
- Leucoplasts colourless plastids: for monoterpene synthesis; leucoplasts sometimes differentiate into more specialized plastids:
- Amyloplasts: for starch storage and detecting gravity
- 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.
Plastids in algae
In
algae, the term leucoplast is used for all
unpigmented plastids and their function differs from the leucoplasts of plants.
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.
Glaucocystophytic algae contain muroplasts, which
are similar to chloroplasts except that they have a cell wall that is similar
to that of prokaryotes. Rhodophytic algae contain rhodoplasts, which are
red chloroplasts that allow the algae to photosynthesise to a depth of up to
268 m.[2]
Inheritance of plastids
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.[3]
Origin of plastids
Plastids
are thought to have originated from endosymbiotic cyanobacteria. The symbiosis evolved around 1.5
billion years ago and enabled eukaryotes to carry out oxygenic photosynthesis.[4] 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 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
instead contain stroma and grana thylakoids, structures found only in plants and
in closely related green algae. 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, 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.
From :
http://en.wikipedia.org/wiki/Plastids
https://www.youtube.com/watch?v=cldx7bObtto
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