Stromule Formation

  • John C. GrayEmail author
Part of the Advances in Photosynthesis and Respiration book series (AIPH, volume 36)


Stromules are highly dynamic stroma-filled tubules extending from the surface of plastids in all multicellular plants. Although stromules may interconnect two, or more, plastids and allow the transfer of stromal proteins as large as Rubisco (∼550 kDa) between plastids, their function is still largely a matter of conjecture. They may increase the plastid surface area to facilitate movement of materials into or out of plastids, be involved in sensing the cellular environment, and/or have signaling functions due to close apposition of stromules and nuclei, plasma membrane and other cell organelles. Stromule formation appears unrelated to chloroplast division or to light-intensity-dependent chloroplast movement. Stromules are most easily observed by confocal microscopy of cells expressing plastid-targeted fluorescent proteins, and the definition of stromules is based on such observations. Identification of stromules in electron microscope images is problematic, and ideally requires examination of thin serial sections. In leaves of both monocots and dicots, stromules are most abundant in epidermal cell-types, such as trichomes, guard cells and pavement cells, and are more difficult to observe in mesophyll cells containing large closely packed chloroplasts. Stromule formation and movement depends on the actin cytoskeleton and requires the ATPase activity of myosin XI proteins. A 42-amino-acid-residue region of myosin XI that directs myosin XI to the chloroplast periphery offers prospects for identification of stromule components required for stromule mobility. Stromules are affected by various environmental factors and by biotic stresses. Stromules are induced by water stress, acting via abscisic acid signaling pathways, and by viral infection. Stromule abundance is also affected by light and temperature.


Green Fluorescent Protein Guard Cell Mesophyll Cell Bundle Sheath Subsidiary Cell 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



Abscisic acid;


1-amino­cyclopropane-1-carboxylic acid;




2,3-butanedione 2-monoxime;


Bimolecular fluorescence complementation;


Cyan fluorescent protein;


Differential interference contrast;


Green fluorescent protein;


Outer envelope protein of 14 kDa;


Polyethylene glycol;


Translocon of the inner chloroplast envelope membrane;


Tobacco mosaic virus;


Translocon of the outer chloroplast envelope membrane;


Triose phosphate–phosphate translocator;


Virus-induced gene silencing;


Yellow fluorescent protein



I am extremely grateful to Jim Sullivan, Michael Hansen and Senthil Natesan for providing previously unpublished images of stromules, and to John Carr for discussion of virus infection. The work in my laboratory on stromules has been supported by research grants from the Biotechnology and Biological Sciences Research Council (BBSRC).


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© Springer Science+Business Media Dordrecht 2013

Authors and Affiliations

  1. 1.Department of Plant SciencesUniversity of CambridgeCambridgeUK

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