See discussions, stats, and author profiles for this publication at: http://www.researchgate.net/publication/225594620 Plant signalling peptides ARTICLE in ACTA PHYSIOLOGIAE PLANTARUM · FEBRUARY 2003 Impact Factor: 1.52 · DOI: 10.1007/s11738-003-0043-y CITATIONS DOWNLOADS VIEWS 5 89 86 3 AUTHORS: Justyna Wiśniewska Alina Trejgell Nicolaus Copernicus University Nicolaus Copernicus University 15 PUBLICATIONS 3,006 CITATIONS 28 PUBLICATIONS 31 CITATIONS SEE PROFILE SEE PROFILE Andrzej Tretyn Nicolaus Copernicus University 189 PUBLICATIONS 1,037 CITATIONS SEE PROFILE Available from: Andrzej Tretyn Retrieved on: 03 August 2015 ACTA PHYSIOLOGIAE PLANTARUM Vol. 25. No. 1. 2003:105-122 review Plant signalling peptides Justyna Wigniewska, Alina Trejgell, Andrzej Tretyn Nicolaus Copernicus University, Institute of General and Molecular Biology, Department of Biotechnology, Gagarina 9, 87-100 Torud, Poland Key words: CLAVATA3, ENOD40, insulin-like mal systems these intercellular communications protein, natriuretic peptide, phytosulfokine, signal- are mainly mediated by chemical signals such as ling peptides, systemin, RALF. steroids, peptides and other small bioactive com- pounds. Peptides are probably the most commonly used signal molecules in animal systems. This is Abstract most likely to be a result of several factors: the ease Biochemical and genetic studies have identifiedpeptides that with which structural variation can be introduced; play crucial roles in plant growth and development,including the availability of a common secretion mechanism defence mechanisms in response to wounding by pests, the control of cell division and expansion, and pollen self-incom- for peptides; and also the fact that the activity of patibility. The first two signalling peptides to be described in peptide signalling molecules can be controlled by plants were tomatosystemin and phytosulfokine(PSK). There common processing and modification mecha- is also biochemicalevidence that natriureticpeptide-like mole- nisms. Dozens of signalling peptides have been cules, immunologically-relatedto those found in animals,may exist in plants. Another example of signalling peptide is identified and most are recognized by specific re- ENOD40, a product of a gene, whichbecame activeearly in the ceptors anchored in the plasmalema of the recipient root nodulationprocess followingRhizobium infection of le- cells (Bisseling 1999). gumes. Other predicted bioactive peptides or oligopeptides have been identifiedby means of genetic, rather then biochem- Hormone is a term once applied in both botanical ical methods. The Arabidopsis CLAVATA3 protein is re- and zoological contexts. For animal physiologists it quired for the correctorganization of the shoot apical meristem and the pollen S determinant S-locus cysteine-rich protein denotes any molecule, usually of small molecular (SCR also called S-locus protein 11, SP11). mass, secreted outside cells and carried to specific The plant signallingpeptides discovered so far are involvedin target cells/organs by whose response they bring various processes and play an importantrole in communication about a specific and adaptive physiological re- between cells or organs, respectively.This reviewwill focuson sponse. Hormones tend to be either water-soluble these peptides and their role in intercellular signalling. peptides and proteins, or steroids. In animals, poly- peptide hormones classically fall into two major Introduction categories: endocrine hormones (Douglass et aL 1984) and membrane-anchored growth factors In multicellular biological systems, cell-cell inter- (Massague and Pandiella 1993). The endocrine actions are indispensable for coordinating cell polypeptide hormones, and some growth factors, growth and differentiation in various organs. In ani- are synthesized as preprohormones in the secretory 105 J. WI~NIEWSKA, A. TREJGELL & A. TRETYN pathway of specialized cells and are sequestered in Plants are using small (phyto)hormones such as: vesicles until released into extracellular compart- auxins, cytokinins, gibberellins, abscisic acid, ethyl- ments in response to appropriate physiological cues ene (Kende and Zeevaart 1997) and brassinoste- (Douglass et al. 1984). Processing of polypeptide roids (Bishop and Koncz 2002) for cell-to-cell hormones from larger prohormone precursors in communication. They work at low concentrations vesfcles is mediated by members of the endopep- (excluding nutritious activity), and regulate growth tidase family. The hormones are then transported to and development of plants by modifying a gene ex- target cells, either nearby or long distances away, pression and/or metabolic processes. Besides these where they are recognized by specific receptors to "classical" hormones plants may also use oligo- initiate signalling pathways. Some well-known ex- and polypeptides as signalling molecules, which amples are: growth hormone, adrenocorticotropic may play similar role to peptide hormones from ani- hormone, oxytocin, vasopressin, calcitonin, para- mal and yeasts. thormone, glucagons, insulin and endorphins (Douglass et al. 1984, Massague and Pandiella 1993). Polypeptide hormones are usually small, be- Systemin ing derived from precursors that are 100-500 aa in Systemin, a polypeptide isolated from plant tissues length (Douglass et al. 1984). Some of these poly- (tomato leaves), is synthesized as a result of me- peptides are produced as a single copy from a pre- chanic damage to plant organs or injury caused by cursor, such as insulin, but some precursors harbor an insect. It is able to move fast in plant and to in- several copies of the hormone, such as proence- duce genes encoding proteinase inhibitor I and II - phalin, while still other precursors are processed to one of the elements of a cascade leading to immu- produce multiple hormones having different phy- nological reaction (Pearce et al. 1991). siological roles, such as pro-opiomelanocortin (Massague and Pandiella 1993). The variability of Systemin regulates the activation of more than 20 structural conformations is critical for the high defensive genes. During systemic reaction, the sti- specificity of receptor binding that exists among mulation of MAP kinase (Mitogen Activated Pro- hundreds of polypeptide hormones (Ryan et al. tein Kinase) is followed by synthesis of trypsine in- 2002). hibitors (Meindl et al. 1998, Pearce et al. 2001b). As a result of direct action of systemin on uninjured m different scenario for polypeptide synthesis and cells, the changes in cytoplasmic concentration of release is found with membrane-anchored precur- Ca 2+ ions are observed. These changes cause the sors of growth factors. Examples include TGF-ot activation of phospholipase A (PLAe), which ca- (Transforming Growth Factor, 50 aa derived from a talyses the degradation of membrane ingredients 160 aa precursor), EGF (Epidermal Growth Factor, leading to releasing of a free linolenic acid from 53 aa derived from a 1207 aa precursor), and membranes and converting it to jasmonic acid TNF-~t (Tumor Necrosis Factor, 157 aa derived (Ryan 2000). Simultaneous increase in cytoplasmic from a 233 aa precursor) (Massague and Pandiella Ca 2+ concentration leads to modulation of plasma 1993). The precursors of most members of this membrane H+-ATPase and in consequence to class of polypeptides are generally between 150 to membrane depolarization and changing of ion 650 aa in length, although some are larger. They are fluxes (Moyen et al. 1998, Ryan and Pearce 1998). synthesized through the secretory pathway; the In plants defensive mechanisms polyphenol oxi- proproteins, however, are not processed in vesicles dase, calmodulin and H202 are also involved. Ac- but are anchored in vesicle membranes. The vesi- cumulation of these compounds is observed after cles fuse with the plasma membrane of the cell, ex- wounding or treatment with systemin or metyle- posing the polypeptide to the extracellular space. jasmonate (Constabel and Ryan 1998, Schaller Upon appropriate cues, membrane-associated pro- 1999, Bergey and Ryan 1999, Ryan 2000, Oroz- teases are activated that release the factors, which co-Cardenas et al. 2001). Increased polygalactu- can be recognized by their receptors (Douglass et ronase activity under influence of systemin and as a al. 1984, Massague and Pandiella 1993). result of leaf injury is also observed (Bergey et al. 106 PLANT SIGNALLING PEPTIDES 1999). In plants and animals there are large analo- (Dombrowski et al. 1999). It was affirmed how- gies in signal transduction triggered either by ever, that extracts from tomato leaves lack the po- wounding or pathogen. In case of plants, the signal tential enzymes that could separate systemin from molecule conveying this information is systemin, prosystemin. whereas jasmonic acid is responsible for direct physiological response (Ryan 2000). In animals, Li and Howe (2001) isolated another prosystemin this function is played by cytokines and prosta- (prosys B) from tomato leaves, which varied from glandins (Bergey et al. 1996, Barciszewski and the earlier one in positions of a few amino acids and Legocki 1997, Stratmann et al. 2000). resulted from an alternative splicing. Systemin is a polypeptide consisting of 18-amino It was shown recently that the polypeptides similar acids with the sequence to systemin also function in other species of Sola- AVQSKPPSKRDPPKMQTD, naceous family beside tomato (potato, black night- which is active when present at fentomoles per shade and bell pepper) (Constabel et aL 1998). The plant (Bergey