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400 J.D.Rhodes,J.F.Thain,D.C.Wildon rations of the electrical events recorded by the surface and bath electrodes would normally lead to their classification as variation potentials with the implication that they reflect depolarisations in the underlying tissues due to chemicals that have travelled in the xylem from the wound site. However, given our current ignorance of detailed mechanisms at the cell level, they could reflect the summation of action potentials travelling at slightly different times in the different STE/CC complexes of the petiole. 26.3 Conclusions and Future Prospects For severe wounding, the results are consistent with the distribution of elicitors of PI synthesis and electrical activity by hydraulic dispersal in the xylem. We conclude that the electrical events are likely to be responses to chemicals transported in the xylem by hydraulic dispersal from the wound site, rather than action potentials propagated from the wound site. Following a small crushing wound, the pattern of systemic PI synthesis was consistent with the transport of a chemical elicitor in the phloem from the wound site. The lack of an electrical event following a small wound indicates that electrical signals are not an essential part of the systemic signalling system that induces PI synthesis. The fact that a severe wound leads to large rapid action-potential-like depolarisations in the cells of the STE/CC complexes in the petiole, but not in other cell types, is intriguing. Do these depolarisations have a function? In the absence of detailed studies at the cell level, the description of wound-induced electrical events recorded with surface-contact electrodes as either variation potentials or action potentials could carry misleading implications about their cellular mechanisms. A significant step forward in plant signalling would be the identification of the cellular mechanism of wound-induced electrical events; this could be achieved using a similar approach to that described here and in Rhodes et al. (1996). However, since the mapping of its genome, Arabidopsis may be the preferred model organism; Arabidopsis does show electrical events following severe wounds (our unpublished observations). The identification of the ion channels responsible for the electrical events could be studied by the use of a range of agonists and inhibitors which could be introduced via the xylem stream and their electrical consequences in the phloem could be monitored using microelectrodes. Acknowledgements. The work from our laboratory was funded by the Biotechnology and Biological Sciences Research Council, UK.
26 Signals and Signalling Pathways in Plant Wound Responses 401 <strong>References</strong> Baker DA, Milburn JA (1989) Appendix Table X: speed of assimilate transport in the phloem of plant species analysed by differing techniques. In: Baker DA, Milburn JA (eds) Transport of photoassimilates. Longman, Harlow, pp 354–355 Baydoun EA-H, Fry SC (1985) The immobility of pectic substances in injured tomato leaves and its bearing on the identity of the wound hormone. Planta 165:269–276 Bowles D (1998) Signal transduction in the wound response of tomato plants. Philos Trans R Soc Lond Ser B 353:1495–1510 Davies E (2004) New functions for electrical signals in plants. New Phytol 161:607–610 Green TR, Ryan CA (1972) Wound-induced proteinase inhibitor in plant leaves: a possible defense mechanism against insects. Science 175:776–777 Herde O, Fuss H, Peña-Cortès H, Fisahn J (1995) Proteinase inhibitor II gene expression induced by electrical stimulation and control of photosynthetic activity in tomato leaves. Plant Cell Physiol 36:737–742 Mackie GO (1965) Conduction in nerve-free epithelia of siphonophores. Am Zool 5:439–453 Malone M (1996) Rapid, long-distance signal transmission in higher plants. Adv Bot Res 22:163–228 Mancuso S (1999) Hydraulic and electrical transmission of wound-induced signals in Vitis vinifera. Aust J Plant Physiol 26:55–61 Moyen C, Johannes E (1996) Systemin transiently depolarizes the tomato mesophyll cell membrane and antagonizes fusicoccin-induced extracellular acidification of mesophyll tissue. Plant Cell Environ 19:464–470 Nelson CE, Walker-Simmons M, Makus D, Zuroske G, Graham J, Ryan CA (1983) Regulation of synthesis and accumulation of proteinase inhibitors in leaves of wounded tomato plants. In: Hedin PA (ed) Plant resistance to insects. ACS symposium series no 208. American Chemical Society, Washington, DC, pp 103–122 Passioura JB (1972) The effect of root geometry on the yield of wheat growing on stored water. Aust J Agric Res 23:745–752 Peña-Cortès H, Fisahn J, Willmitzer L (1995) Signals involved in wound-induced proteinase inhibitor II gene expression in tomato and potato plants. Proc Natl Acad Sci USA 92:4106–4113 Rhodes JD, Thain JF, Wildon DC (1996) The pathway for systemic electrical signal conduction in the wounded tomato plant. Planta 200:50–57 Rhodes JD, Thain JF, Wildon DC (1999) Evidence for physically distinct systemic signalling pathways in the wounded tomato plant. Ann Bot 84:109–116 Rigby NM, MacDougall AJ, Needs PW, Selvendran RR (1994) Phloem translocation of a reduced oligogalacturonide in Ricinus communis L. Planta 193:536–541 Stankovic B, Davies E (1996) Both action potentials and variation potentials induce proteinase inhibitor gene expression in tomato. FEBS Lett 390:275–279 Stankovic B, Davies E (1997) Intercellular communication in plants: electrical stimulation of proteinase inhibitor gene expression in tomato. Planta 202:402–406 Stratmann JW (2003) Long distance run in the wound response – jasmonic acid is pulling ahead. Trends Plant Sci 8:247–250 Thain JF, Wildon DC (1996) Electrical signalling in plants. In: Smallwood M, Knox JP, Bowles DJ (eds) Membranes: specialized functions in plants. Bios, Oxford, pp 301–317 Thain JF, Gubb IR, Wildon DC (1995) Depolarization of tomato leaf cells by oligogalacturonide elicitors. Plant Cell Environ 18:211–214 van Bel AJE (2003) The phloem, a miracle of ingenuity. Plant Cell Environ 26:125–149 Wildon DC, Thain JF, Minchin PEH, Gubb IR, Reilly AJ, Skipper YD, Doherty HM, O’Donnel PJ, Bowles DJ (1992) Electrical signalling and systemic proteinase inhibitor induction in the wounded plant. Nature 360:62–65
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František Baluška · Stefano Manc
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Dr. František Baluška University
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VI Preface turn, reward the ants by
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VIII Preface of olfactory response.
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Contents 1 The Green Plant as an In
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Contents XIII 6 Signals and Targets
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Contents XV 11 Amino Acid Transport
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Contents XVII 15 Regulation of Plan
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Contents XIX 21.3 Conclusions and P
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Contents XXI 27.3.2 Catechin Induce
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XXIV Contributors Correa-Aragunde,
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XXVI Contributors Lamattina, L. (e-
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XXVIII Contributors Song, C. Depart
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1 The Green Plant as an Intelligent
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38 P.W. Barlow Thestimulipresentedt
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40 P.W. Barlow that a tropism is su
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42 P.W. Barlow the auxin flow into
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44 P.W. Barlow afferentnervousimpul
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46 P.W. Barlow analysis. In particu
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48 P.W. Barlow reception of his boo
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50 P.W. Barlow Iijima M, Kono Y (19
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66 P.M. Neumann Finally, I examined
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68 P.M. Neumann evolutionary progre
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70 P.M. Neumann conclusion is that
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72 P.M. Neumann resources from matu
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6 Signals and Targets Triggered by
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6TargetsofSI 79 by Yang 2002) has p
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6TargetsofSI 81 al. 2002). Thus, SI
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6TargetsofSI 83 Fig.6.2. PrABP80 ha
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6TargetsofSI 85 many of the genes e
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6TargetsofSI 87 [Ca 2+ ]i may signa
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6TargetsofSI 89 is crosstalk betwee
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6TargetsofSI 91 Hepler PK, Vidali L
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6TargetsofSI 93 Snowman BN, Kovar D
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96 T. Nürnberger, B. Kemmerling Wh
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124 M.L. Lanteri et al. 9.1.1 Auxin
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134 M.L. Lanteri et al. Bellamine J
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136 M.L. Lanteri et al. Pagnussat G
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138 S.J. Murch H 3C CH3 CH3 N H 2C
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140 S.J. Murch edible plants (Manch
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142 S.J. Murch However, over the la
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144 S.J. Murch of monoamine, amino
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146 S.J. Murch On Guam and in other
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148 S.J. Murch 10.4 Conclusions and
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150 S.J. Murch Lindstrom H, Luthman
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11 Amino Acid Transport in Plants a
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11 AA transport in plant versus neu
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188 M. Gilliham et al. Fig.13.1. Ar
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190 M. Gilliham et al. as compatibl
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192 M. Gilliham et al. apex (Zhang
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198 M. Gilliham et al. 2005). It is
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17 Touch-Responsive Behaviors and G
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18 Oscillations in Plants Sergey Sh
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18 Oscillations in Plants 271 prehe
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278 K.Trebacz,H.Dziubinska,E.Krol W
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292 R. Stahlberg, R.E. Cleland, E.
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318 E.Davies,B.Stankovic 1992; Beel
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320 E.Davies,B.Stankovic Hentze MW,
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322 J. Fromm, S. Lautner in the ran
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324 J. Fromm, S. Lautner cells (Sam
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326 J. Fromm, S. Lautner 22.5 Ion C
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334 S. Mancuso, S. Mugnai like the
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336 S. Mancuso, S. Mugnai 2,000 nM
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338 S. Mancuso, S. Mugnai the fourt
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342 S. Mancuso, S. Mugnai 23.6 Airb
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346 S. Mancuso, S. Mugnai Fort C, F
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Page 452 and 453: 422 V.Ninkovic,R.Glinwood,J.Petters
Page 466 and 467: 436 Subject Index defense 67, 95-10
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