Plasmodesma

Plasmodesma
Plasmodesmata (singular: plasmodesma) are microscopic channels which traverse the cell walls of plant cells[2] [3] and some algal cells, enabling transport and communication between them. Species that have plasmodesmata include members of the Charophyceae, Charales and Coleochaetales (which are all algae), as well as all embryophytes, better known as land plants.[4] Unlike animal cells, every plant cell is surrounded by a polysaccharide cell wall. Neighbouring plant cells are therefore separated by a pair of cell walls and the intervening lamella, forming an extracellular domain known as the apoplast. Although cell walls are permeable to small soluble proteins and other solutes, plasmodesmata enable direct, regulated, symplastic intercellular transport of substances between cells. There are two forms of plasmodesmata: primary ones are formed during cell division and secondary ones can form between mature cells.[5] Similar structures, called gap junctions[6] and membrane nanotubes, interconnect animal cells[7] and stromules form between plastids in plant cells.[8]

Plasmodesma allow molecules to travel between plant cells through the symplastic pathway

Formation
Plasmodesmata are formed when portions of the endoplasmic reticulum are trapped across the middle lamella as new cell wall is laid down between two newly divided plant cells and these eventually become the cytoplasmic connections between cells (primary plasmodesmata). Here the wall is not thickened further, and depressions or thin areas known as pits are formed in the walls. Pits normally pair up between adjacent cells. Alternatively, plasmodesmata can be inserted into existing cell walls

The structure of a primary plasmodesma. CW=Cell wall CA=Callose PM=Plasma membrane ER=Endoplasmic reticulum DM=Desmotubule Red circles=Actin Purple circles and spokes=Other unidentified [1] proteins

Plasmodesma between non-dividing cells (secondary plasmodesmata)[9]

Structure
Plasmodesmatal plasma membrane
A typical plant cell may have between 103 and 105 plasmodesmata connecting it with adjacent cells[10] equating to between 1 and 10 per m2.[11] Plasmodesmata are approximately 50-60nm in diameter at the mid-point and are constructed of three main layers, the plasma membrane, the cytoplasmic sleeve, and the desmotubule.[10] They can transverse cell walls that are up to 90nm thick.[11] The plasma membrane portion of the plasmodesma is a continuous extension of the cell membrane or plasmalemma [12] It is similar in structure to the cellular phospholipid bilayers.

Cytoplasmic sleeve
The cytoplasmic sleeve is a fluid-filled space enclosed by the plasmalemma and a continuous extension of the cytosol. Trafficking of molecules and ions through plasmodesmata occurs through this passage. Smaller molecules (e.g. sugars and amino acids) and ions can easily pass through plasmodesmata by diffusion without the need for additional chemical energy. Proteins can also pass through the cytoplasmic sleeve (for example Green fluorescent protein).[13] It is not yet known how the selective transport of larger molecules, such as proteins, occurs. One hypothesis is that the polysaccharide callose accumulates around the neck region of plasmodesmata to form a collar, reducing their diameter and thereby controlling permeability to substances in the cytoplasm.[12]

Desmotubule
The desmotubule is a tube of appressed endoplasmic reticulum that runs between two adjacent cells [14] Some molecules are known to be transported through this channel,[15] but it is not thought to be the main route for plasmodesmatal transport. Around the desmotubule and the plasma membrane areas of an electron dense material have been seen, often joined together by spoke-like structures that seem to split the plasmodesma into smaller channels [14] These structures may be composed of myosin[16] [17] [18] and actin,[17] [19] which are part of the cell's cytoskeleton. If this is the case these proteins could be used in the selective transport of large molecules between the two cells.

Plasmodesma

Transport
Plasmodesmata have been shown to transport proteins (including transcription factors), short interfering RNA, messenger RNA and viral genomes from cell to cell. One example of a viral movement proteins is the tobacco mosaic virus MP-30. MP-30 is thought to bind to the virus's own genome and shuttle it from infected cells to uninfected cells through plasmodesmata.[13] Flowering Locus T protein moves from leaves to the shoot apical meristem through plasmodesmata to initiate flowering. The size of molecules that can pass through plasmodesmata is determined by the size exclusion limit. This limit is highly variable and can is subject to active modification.[5] MP-30 is able to increase the size exclusion limit from 700 Daltons to 9400 Daltons thereby aiding its movement through a plant.[20]

Several models for possible active transport through plasmodesmata exist. It has been suggested that such transport is mediated by interactions with proteins localized on the desmotubule, and/or by chaperones partially unfolding proteins, allowing them to fit through the narrow passage. A similar mechanism may be involved in transporting viral nucleic acids through the plasmodesmata.[21]

Tobacco mosaic virus movement protein 30 localizes to plasmodesmata.

References
[1] Maule, Andrew (December 2008). "Plasmodesmata: structure, function and biogenesis". Current Opinion in Plant Biology 11 (6): 680686. doi:10.1016/j.pbi.2008.08.002. PMID18824402. [2] Oparka, K. J. (2005) Plasmodesmata. Blackwell Pub Professional. ISBN 10: 1405125543 ISBN 13: 9781405125543 [3] Plasmodesmata (www.dictionary.com) (http:/ / dictionary. reference. com/ browse/ Plasmodesmata) [4] Graham, LE; Cook, ME; Busse, JS (2000), Proceedings of the National Academy of Sciences 97, 4535-4540. [5] http:/ / www. pubmedcentral. nih. gov/ articlerender. fcgi?artid=1692983 The shoot apical meristem: the dynamics of a stable structure. Jan Traas and Teva Vernoux : Philos Trans R Soc Lond B Biol Sci. 2002 June 29; 357(1422): 737747. (page 744) [6] Bruce Alberts (2002). Molecular biology of the cell (4th ed.). New York: Garland Science. ISBN0-8153-3218-1. [7] Gallagher KL, Benfey PN (January 2005). "Not just another hole in the wall: understanding intercellular protein trafficking" (http:/ / www. genesdev. org/ cgi/ pmidlookup?view=long& pmid=15655108). Genes Dev. 19 (2): 18995. doi:10.1101/gad.1271005. PMID15655108. . [8] Gray JC, Sullivan JA, Hibberd JM, Hansen MR (2001). "Stromules: mobile protrusions and interconnections between plastids". Plant Biology 3: 22333. doi:10.1055/s-2001-15204. [9] Lucas W., Ding, B. and Van der Schoot, C. (1993) Tansley Review No.58 "Plasmodesmata and the supracellular Nature of Plants" New Phytologist, Vol. 125, No. 3, pp. 435-476, Stable URL: http:/ / www. jstor. org/ stable/ 2558257 [10] AW Robards (1975) Plasmodesmata. Annual Review of Plant Physiology 26, 13-29 [11] "22". Molecular Cell Biology (4 ed.). 2000. pp.998. ISBN0-7167-3706-X. [12] AW Robards (1976) Plasmodesmata in higher plants. In: Intercellular communications in plants: studies on plasmodesmata. Edited by BES Gunning and AW Robards Springer-Verlag Berlin pps 15-57. [13] http:/ / www. ingentaconnect. com/ content/ bsc/ pce/ 2003/ 00000026/ 00000001/ art00007 Plasmodesmata and the control of symplastic transport A. G. ROBERTS & K. J. OPARKA [14] RL Overall, J Wolfe, BES Gunning (1982) Intercellular communication in Azolla roots: I. Ultrastructure of plasmodesmata. Protoplasma 111: 134-150 [15] LC Cantrill, RL Overall and PB Goodwin (1999) Cell-to-cell communication via plant endomembranes. Cell Biology International 23: 653661 [16] JE Radford and RG White (1998) Localization of a myosinlike protein to plasmodesmata. Plant Journal 14: 743-750 [17] LM Blackman and RL Overall (1998) Immunolocalisation of the cytoskeleton to plasmodesmata of Chara corallina. Plant Journal 14: 733-741 [18] S Reichelt, AE Knight, TP Hodge, F Baluska, J Samaj, D Volkmann and J Kendrick-Jones (1999) Characterization of the unconventional myosin VIII in plant cells and its localization at the post-cytokinetic cell wall. Plant Journal 19: 555569 [19] RG White, K Badelt, RL Overall and M Vesk (1994) Actin associated with plasmodesmata. Protoplasma 180: 169-184

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[20] http:/ / www. sciencemag. org/ cgi/ content/ abstract/ 246/ 4928/ 377 Science 20 October 1989: Vol. 246. no. 4928, pp. 377 - 379 Movement Protein of Tobacco Mosaic Virus Modifies Plasmodesmatal Size Exclusion Limit SHMUEL WOLF, WILLIAM J. LUCAS, CARL M. DEOM,and ROGER N. BEACHY [21] http:/ / jpkc. zju. edu. cn/ k/ 437/ content/ 05. pdf Plant Physiology lectures, chapter 5

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