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256 E. McCormack et al. interpretation that darkness and perhaps the other stimuli that induce expression of the TCH genes are related in some way. One possibility is that all these stimuli share the property of causing mechanical perturbations andinthiswaytriggeracommonmechanosensorypathwaythatleadsto gene upregulation of expression (Braam 2000). If the diverse stimuli that trigger TCH upregulation of expression act throughacommonpathway,thenonewouldexpectthatasinglecis regulatory element would be responsible for conferring the complex expression regulation. Indeed, a 102 base-pair region located upstream of the TCH4 transcriptional start site is sufficient to confer touch, darkness, cold heat and epi-brassinolide upregulation of expression to reporter genes (Iliev et al. 2002). Sequences within this region share some similarities to sequences identified as important for conferring cold and touch inducibility to the CBF2 gene (Zarka et al. 2003). The mechanism by which these sequences mayacttoregulategeneexpressioniscurrentlyunknown. Distinct methods have been used to monitor TCH expression during plant morphogenesis. TCH::reporter transgenics have been characterized in addition to the more direct methods of immunolocalization and reverse transcription PCR (Sistrunk et al. 1994; Antosiewicz et al. 1995, 1997; Xu et al. 1995; Delk et al. 2005). In general, TCH expression is enriched at sites that may be predicted to experience mechanical stress. TCH protein accumulates and/or TCH::reporter transgenes are expressed in the ruptured seed coat, branch points, the root–shoot junction, elongating hypocotyls and roots, and developing trichomes and silique abscission zones. In addition, plants subjected to enhanced weight on the inflorescence have increased TCH2 and TCH4 expression (Ko et al. 2004). These data indicate that TCH expression may not only be upregulated in response to externally applied mechanical perturbations such as touch, but also by mechanical forces that become manifest during normal plant morphogenesis. 17.5 Conclusions and Future Prospects Plants perceive much more of their environment than is often apparent to the casual observer. Touch can induce profound rapid responses and more slowly acquired growth alterations. Rapid touch-induced plant movement in specialized plants is often associated with predation or protection. The speed of these responses is an essential component of the response in these situations. Plants that acclimate over their lifetime to touch or wind stimuli also undergo dramatic touch-induced changes, but these, at least overtly, occurslowlyovertime.Molecularresponsesinnonspecializedplantscan, however, occur quite rapidly. In Arabidopsis, changes in gene expression are
17 Touch-Responsive Behaviors and Gene Expression in Plants 257 seen within minutes after touch, and over 700 genes have altered transcript levels within 30 min. The prevalence of touch-responsive genes and the rapidity with which such changes in gene expression occur are indications that nonspecialized plants do possess capacity for rapid responses to touch. Recent research reveals the types of genes upregulated by touch and implicate Ca 2+ signaling, cell wall modification and disease resistance as potential downstream responses. Further work is needed to reveal the physiological relevance of these touch-induced changes in gene expression. Furthermore, touch-induced genes are powerful molecular tools for thedissectionoftheperceptionandresponsepathwaysthatenableplants to perceive mechanical stimuli and manifest appropriate responses. Perception mechanisms and intercellular and intracellular transduction machinery and signals await future discovery. Acknowledgements. Our research in this area is supported by the Department of Energy (grant no. DE-FG02-03ER15394) and the National Science Foundation (grant nos. IBN-9982654, IBN-0321532 and IBN-0313432). <strong>References</strong> Antosiewicz DM, Polisensky DH, Braam J (1995) Cellular localization of the Ca 2+ binding TCH3 protein of Arabidopsis. Plant J 8:623–636 Antosiewicz DM, Purugganan MM, Polisensky DH, Braam J (1997) Cellular localization of Arabidopsis xyloglucan endotransglycosylase-related proteins during development and after wind stimulation. Plant Physiol 115:1319–1328 Barnett JR, Asante AK (2000) The formation of cambium from callus in grafts of woody species. In: Savidge RA, Barnett JR, Napier R (eds) Cell and molecular biology of wood formation. BIOS, Oxford, pp 155–167 Batiza AF, Schulz T, Masson PH (1996) Yeast respond to hypotonic shock with a calcium pulse. J Biol Chem 271:23357–23362 Biddington NL (1986) The effects of mechanically-induced stress in plants – a review. Plant Growth Regul 4:103–123 Blechert S, Bockelmann C, Füsslein M, v Schrader T, Stelmach BA, Niesel U, Weiler EW (1999) Structure-activity analyses reveal the existence of two separate groups of active octadecanoids in elicitation of the tendril-coiling response of Bryonica dioica Jacq. Planta 207:470–479 Braam J (1992) Regulated expression of the calmodulin-related TCH genes in cultured Arabidopsis cells: induction by calcium and heat shock. Proc Natl Acad Sci USA 89:3213– 3216 Braam J (2000) The Arabidopsis TCH genes: regulated in expression by mechanotransduction? In: Cherry JH, Locy RD, Rychter A (eds) Plant tolerance to abiotic stresses in agriculture: role of genetic engineering, vol 83, Kluwer, Dordrecht, pp 29–37 Braam J (2005) In touch: plant responses to mechanical stimuli. New Phytol 165:373–389 Braam J, Davis RW (1990) Rain-, wind- and touch-induced expression of calmodulin and calmodulin-related genes in Arabidopsis. Cell 60:357–364 Burdon-Sanderson J (1873) Note on the electrical phenomena which accompany stimulation of leaf of Dionaea muscipula. Proc Philos Trans R Soc Lond 21:495–496
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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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2 Neurobiological View of Plants an
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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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4 How Can Plants Choose the Most Pr
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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 77 6.1.2 Self-Incompat
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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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194 M. Gilliham et al. 13.3.3 Are A
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196 M. Gilliham et al. sufficiently
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198 M. Gilliham et al. 2005). It is
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204 M. Gilliham et al. Zheng Y, Mel
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310 E.Davies,B.Stankovic It is assu
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312 E.Davies,B.Stankovic 21.2 Evide
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316 E.Davies,B.Stankovic Relative m
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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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328 J. Fromm, S. Lautner hastobedon
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330 J. Fromm, S. Lautner Beilby MJ,
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332 J. Fromm, S. Lautner Williams S
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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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340 S. Mancuso, S. Mugnai “slow-w
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342 S. Mancuso, S. Mugnai 23.6 Airb
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344 S. Mancuso, S. Mugnai that tree
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346 S. Mancuso, S. Mugnai Fort C, F
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348 S. Mancuso, S. Mugnai Pophof B,
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372 E. Wagner et al. Exudation [µl
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374 E. Wagner et al. metabolism at
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376 E. Wagner et al. The circadian
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378 E. Wagner et al. Fig.25.5. Time
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380 E. Wagner et al. Fig.25.7. Patt
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384 E. Wagner et al. Fig.25.11. a K
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386 E. Wagner et al. 25.9 Conclusio
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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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406 L.G. Perry et al. Fig.27.1. A s
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408 L.G. Perry et al. phytotoxins i
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410 L.G. Perry et al. monooxygenase
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412 L.G. Perry et al. 27.4 (±)-Cat
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414 L.G. Perry et al. (±)-catechin
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416 L.G. Perry et al. mineralizatio
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418 L.G. Perry et al. Dyer AR (2004
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420 L.G. Perry et al. Singh HP, Bat
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422 V.Ninkovic,R.Glinwood,J.Petters
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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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