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308 R. Stahlberg, R.E. Cleland, E. Van Volkenburgh Stankovic B, Zawadzki T, Davies E (1997) Characterization of the variation potential in sunflower. Plant Physiol 115:1083–1088 Stankovic B, Witters DL, Zawadzki T, Davies E (1998) Action and variation potentials in sunflower: an analysis of their relationship and distinguishing characteristics. Physiol Plant 103:51–58 Theilet C, Delpeyroux F, Fiszman M, Reigner P, Esnault R (1982) Influence of excision shockonontheproteinmetabolismofVicia faba L. meristematic root cells. Planta 155:478–485 Tinz-Fuchtmeyer A, Gradmann D (1990) Laser-interferometric re-examination of rapid conductance of excitation in Mimosa pudica. J Exp Bot 41:15–19 Tsaplev YB, Zatsepina GN (1980) The electrical nature of the propagation of the variation potential in Tradescantia. Biofizika 25:723–728 Umrath K (1959) Der Erregungsvorgang. In: Ruhland W (ed) Handbuch der Pflanzenphysiologie, vol 17. Springer, Berlin Heidelberg New York, pp 24–110 Van Sambeek JW, Pickard BG (1976) Mediation of rapid electrical, metabolic, transpirational and photosynthetic changes by factors released from wounds. Can J Bot 54:2662–2671 Volkov AG, Haak RA (1995) Bioelectrochemical signals in potato plants. Russ J Plant Physiol 42:17–23 Westgate ME, Steudle E (1985) Water transport in the midrib of tissue of maize leaves. Direct measurement of the propagation of changes in cell turgor across a plant tissue. Plant Physiol 78:183–191 Wildon DC, Thain JF, Minchin PEH, Gubb IR Reilly AJ, Skipper YD, Doherty HM, O’Donnell PJ, Bowles DJ (1992) Electrical signaling and systemic proteinase inhibitor induction in the wounded plant. Nature 360:62–65 Zawadzki T, Davies E, Dziubinska H, Trebasz K (1991) Characteristics of action potentials in Helianthus annuus. Physiol Plant 83:601–604 Zerrenthin U, Stahlberg R (1981) Die Nutzung der bioelektrischen Kurzzeitreaktion auf mechanische Reizung zur Ermittlung von Kennwerten und Sortenunterschioeden bei Getreidekeimpflanzen. Arch Acker- Pflanzenbau und Boden 25:197–203
21 Electrical Signals, the Cytoskeleton, and Gene Expression: a Hypothesis on the Coherence of the Cellular Responses to Environmental Insult Eric Davies, Bratislav Stankovic Abstract When plant tissue is abiotically injured by crushing, cutting, heat-wounding, electrical stimulation, or by several other means, the injured (perceiving) tissue generates electrical signals (action potentials and variation potentials) and transmits them to distant (responding) tissue. Here they evoke apparently disparate responses, such as callose formation, closing of plasmodesmata, stoppage of cytoplasmic streaming, inhibition of ribosome movement along messenger RNA (mRNA), and ultrarapid but transient accumulation of over 100 transcripts, which are degraded without being translated. These apparently disparate responses can be reconciled by one fundamental hypothesis that assumes that “the plant does not know what hit it” and thus “expecting the worst” mounts a holistic defense response against its most potent nemesis, a putative viral invasion. We postulate that the basis for this response is calcium influx into the cytoplasm via voltage-gated channels (action potential) associated with the microtubules, or via mechano-sensitive channels (variation potential) associated with microfilaments. The calcium interacts with calcium and/or calmodulin-dependent cytoskeleton-associated protein kinases. This causes the phosphorylation of myosin, which stops cytoplasmic streaming, and of elongation factor 2F, which slows elongation and termination and causes ribosomes to pile up on polyribosomes. This decreases protein synthesis, but protects preexisting “host” transcripts from degradation. The phosphorylation signal then passes into the nucleus, where it phosphorylates RNA polymerase II, which goes into overdrive (i.e., does not stop at accuracy checkpoints), thus causing the synthesis of large amounts of mismade mRNA. The mRNA is transported into the cytoplasm, where it is scanned (checked for accuracy) by ribosomes, and found to be incorrect. This surveillance mechanism stimulates ribonuclease activity, which degrades the free (non-polysome-associated), mismade RNA, but leaves the original, “host” transcripts unscathed since they are protected by ribosomes. The ribonuclease also (and here is the crux of the matter) attacks other free mRNAs, including viral mRNAs, so these are disposed of before they can be translated. Within minutes this reaction is over, cytoplasmic steaming resumes, translation continues, ribosomes are released and so can be used to translate new (correctly made) transcripts. 21.1 Introduction to the Hypothesis There are several different kinds of electrical activities in plants, including action potentials (APs), variation potentials (VPs), voltage spikes or voltage transients, and rhythmic electrical activities but their role is far from understood (Davies 1987b, 2004; Davies et al. 1991). They all involve ion fluxes across membranes and bring about changes in membrane potential. Communication in Plants F. Baluška, S. Mancuso, D. Volkmann (Eds.) © Springer-Verlag Berlin Heidelberg 2006
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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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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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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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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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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 91 Hepler PK, Vidali L
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150 S.J. Murch Lindstrom H, Luthman
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15 Regulation of Plant Growth and D
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26 Signals and Signalling Pathways
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404 L.G. Perry et al. properties (S
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418 L.G. Perry et al. Dyer AR (2004
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436 Subject Index defense 67, 95-10
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438 Subject Index sieve-tube-elemen
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