Stromule
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A stromule is a
microscopic The microscopic scale () is the scale of objects and events smaller than those that can easily be seen by the naked eye, requiring a lens or microscope to see them clearly. In physics, the microscopic scale is sometimes regarded as the scale betwe ...
structure found in
plant cell Plant cells are the cells present in Viridiplantae, green plants, photosynthetic eukaryotes of the kingdom Plantae. Their distinctive features include primary cell walls containing cellulose, hemicelluloses and pectin, the presence of plastids ...
s. Stromules ( stroma-filled tubules) are highly dynamic structures extending from the surface of all
plastid A plastid is a membrane-bound organelle found in the Cell (biology), cells of plants, algae, and some other eukaryotic organisms. Plastids are considered to be intracellular endosymbiotic cyanobacteria. Examples of plastids include chloroplasts ...
types, including proplastids,
chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...
s, etioplasts, leucoplasts,
amyloplast Amyloplasts are a type of plastid, double-enveloped organelles in plant cells that are involved in various biological pathways. Amyloplasts are specifically a type of leucoplast, a subcategory for colorless, non-pigment-containing plastids. Amylo ...
s, and
chromoplast Chromoplasts are plastids, heterogeneous organelles responsible for pigment synthesis and storage in specific Photosynthesis, photosynthetic eukaryotes. It is thought (according to symbiogenesis) that like all other plastids including chloroplast ...
s. Protrusions from and interconnections between plastids were observed in 1888 ( Gottlieb Haberlandt) and 1908 ( Gustav Senn) and have been described sporadically in the literature since then. Stromules were recently rediscovered in 1997 and have since been reported to exist in a number of
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit ...
species including ''
Arabidopsis thaliana ''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small plant from the mustard family (Brassicaceae), native to Eurasia and Africa. Commonly found along the shoulders of roads and in disturbed land, it is generally ...
'',
wheat Wheat is a group of wild and crop domestication, domesticated Poaceae, grasses of the genus ''Triticum'' (). They are Agriculture, cultivated for their cereal grains, which are staple foods around the world. Well-known Taxonomy of wheat, whe ...
,
rice Rice is a cereal grain and in its Domestication, domesticated form is the staple food of over half of the world's population, particularly in Asia and Africa. Rice is the seed of the grass species ''Oryza sativa'' (Asian rice)—or, much l ...
and
tomato The tomato (, ), ''Solanum lycopersicum'', is a plant whose fruit is an edible Berry (botany), berry that is eaten as a vegetable. The tomato is a member of the nightshade family that includes tobacco, potato, and chili peppers. It originate ...
, but their role is not yet fully understood. This highly dynamic nature is caused by the close relationship between plastid stromules and actin microfilaments, which are anchored to the stromule extensions, either in a longitudinal fashion to pull from the stromule and guide the plastid in a given direction or in a hinge fashion allowing the plastid to rest anchored in a given place. The actin microfilaments also define the stromule shape through their interactions. This dynamic random walk-like movement is probably caused by
Myosin Myosins () are a Protein family, family of motor proteins (though most often protein complexes) best known for their roles in muscle contraction and in a wide range of other motility processes in eukaryotes. They are adenosine triphosphate, ATP- ...
XI proteins as a recent work found. Other
organelles In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' th ...
are also associated to stromules, as
mitochondria A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is us ...
, which have been observed associated and sliding over stromule tubes. Plastids and
mitochondria A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is us ...
need to be spatially close as some
metabolic pathways In biochemistry, a metabolic pathway is a linked series of chemical reactions occurring within a cell. The reactants, products, and intermediates of an enzymatic reaction are known as metabolites, which are modified by a sequence of chemical ...
like
photorespiration Photorespiration (also known as the oxidative photosynthetic carbon cycle or C2 cycle) refers to a process in plant physiology, plant metabolism where the enzyme RuBisCO oxygenates RuBP, wasting some of the energy produced by photosynthesis. Th ...
require the association of both organelles to recycle
glycolate Glycolic acid (or hydroxyacetic acid; chemical formula ) is a colorless, odorless and hygroscopic crystalline solid, highly soluble in water. It is used in various skin-care products. Glycolic acid is widespread in nature. A glycolate (sometimes s ...
and detoxify the
ammonium Ammonium is a modified form of ammonia that has an extra hydrogen atom. It is a positively charged (cationic) polyatomic ion, molecular ion with the chemical formula or . It is formed by the protonation, addition of a proton (a hydrogen nucleu ...
produced during photorespiration. Stromules are usually 0.35–0.85 Î¼m in diameter and of variable length, from short beak-like projections to linear or branched structures up to 220 Î¼m long. They are enclosed by the inner and outer plastid envelope membranes and enable the transfer of molecules as large as
RuBisCO Ribulose-1,5-bisphosphate carboxylase/oxygenase, commonly known by the abbreviations RuBisCo, rubisco, RuBPCase, or RuBPco, is an enzyme () involved in the light-independent (or "dark") part of photosynthesis, including the carbon fixation by wh ...
(~560 k Da) between interconnected plastids. Stromules occur in all cell types, but stromule morphology and the proportion of plastids with stromules vary from tissue to tissue and at different stages of plant development. In general, smaller plastids produce shorter stromules, although the largest plastids, mesophyll chloroplasts produce relatively short stromules, indicating that other factors control stromule formation. In general, stromules are more abundant in cells containing non-green plastids, and in cells containing smaller plastids. The primary function of stromules is still unresolved, although the presence of stromules markedly increases the plastid surface area, potentially increasing transport to and from the cytosol. Other functions of stromules, such as transfer of macromolecules between plastids and
starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diet ...
granule formation in cereal
endosperm The endosperm is a tissue produced inside the seeds of most of the flowering plants following double fertilization. It is triploid (meaning three chromosome sets per nucleus) in most species, which may be auxin-driven. It surrounds the Embryo#Pla ...
, may be restricted to particular tissues and cell types.


See also

*
Membrane nanotube A tunneling nanotube (TNT) or membrane nanotube is a term that has been applied to cytoskeletal protrusions that extend from the plasma membrane which enable different animal cells to connect over long distances, sometimes over 100 Micrometre, Î ...


Sources


A Novel View of Chloroplast Structure
a good introduction on stromules, with nice fluorescence pictures of
chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...
s and stromules. * : A good review on stromules (unfortunately access is restricted to subscribers)


References

{{Reflist Plant physiology Cell anatomy