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population more able to spread to neighboring populations. Because the advantage of insect- resistant pl ants i ncreases unde r ha rsh c onditions, i t i s likely t hat r esistant pl ants w ould dominate t he popul ation a nd t hat t he popul ation w ould e volve f rom 0 t o 100 % r esistant within a short time. Letourneau and Hagen (2009) have found that B. rapa plants (including those protected and exposed to Bt -susceptible herbivores) produced more seeds in disked soil or field margins than in natural v egetation. If this population were to spread into disturbed habitats, the fate of adjacent wild populations could quickly depend on the prevalence of the invading insect-resistant plants. Implication for the invasion of transgenic volunteers into wild populations The current study shows that sufficient simulated herbivory decreases plant fitness, showing that h ealthy p lants h ave a cl ear co mpetitive a dvantage. T his ad vantage v aried w ith t he percentage of resistant p lants i n t he popul ation, and w as m ore obvi ous under l ow r esource availability than when resources were high. In addition, the interaction between herbivory and competition ha d a n a dditive e ffect on pl ant f itness. H owever, pl ants had t he a bility f or compensatory growth f ollowing i njury b y h erbivory. Low c ompetition a nd s ufficient resources support regrowth in semi-natural environments. Thus, at the final equilibrium (i.e. resistance gene f ixed i n a popul ation) ve getative a nd reproductive p roduction of wild populations was not significantly altered. The overall productive output of the pure resistant population w as t he s ame as t he i nsect-attacked pure popul ation ( Figure 3.8), m eaning t hat changing t he popul ation f rom s usceptible t o i nsect-resistant w ould not r esult i n di fferent demographic k inetics. T herefore, t hese results s uggest t hat, al though i nsect-resistant p lants can invade and completely replace wild susceptible populations, which still is one of major concerns on c ommercial r elease o f t ransgenic crops, t here will be no change t o t he pl ant community e quilibrium due t o t he i ntrinsic c hange i n t he B. j uncea demography, unl ess herbivorous insects switch to another species. This could result in more resources available in the habitat for the insect-resistant species. Acknowledgements This work was supported by a PhD joint fellowship between France and China (CNOUS, No. 20072315) and a fund from the National Natural Science Foundation of China(NSFC grant No. 30970432). 111
References Agrawal AA. 2004. R esistance and susceptibility of milkweed: Competition, root herbivory, and plant genetic variation. Ecology 85: 2118-2133. Baldwin IT. 1990. H erbivory s imulations i n e cological r esearch. Trends i n E cology and Evolution 5: 91–93. Bing DJ, Downey RK, Rakow GFW. 1996. H ybridizations among Brassica napus, B. rapa and B. j uncea and t heir t wo w eedy relatives B.nigra and Sinapis ar vensis under op en pollination conditions in the field. Plant Breeding 115: 470-473. Boalt E , Lehtilâ K . 200 7. Tolerance t o a pical a nd f oliar da mage: c osts a nd m echanisms i n Raphanus raphanistrum. Oikos 116: 2071-2081. Chase JM, Abrams PA, Grover JP, Diehl S, Chesson P, Holt RD, Richards SA, Nisbet RM, Case TJ. 2002. The interaction between predation and competition: a review and synthesis. Ecology Letters 5: 302-315. Crawley MJ, Brown SL. 1995. Seed limitation and the dynamics of feral oilseed rape on the M25 motorway. Proceedings of the Royal Society of London B 259: 49–54. Crawley MJ, Brown SL. 2004. Spatially structured population dynamics in feral oilseed rape. Proceedings of the Royal Society of London B 271: 1909–1916. Damgaard C , K jaer C . 2009. C ompetitive interactions a nd t he e ffect of he rbivory on Bt- Brassica napus, Brassica rapa and Lolium perenne. Journal of Applied Ecology 46: 107- 1079 D’Hertefeldt T, Jørgensen R B, P ettersson L B. 2008. L ong-term p ersistence o f G M o ilseed rape in the seedbank. Biology Letters 3: 314–317. Di K, Stewart CN Jr, Wei W, Shen BC, Tang ZX, Ma KP. 2009. Fitness and maternal effects in h ybrids f ormed b etween t ransgenic o ilseed r ape ( Brassica napus L.) a nd w ild br own mustard [B. juncea (L.) Czern et Coss.] in the field. Pest Management Science 65: 753–760. Ellstrand N C, P rentice H C, H ancock J F. 1 999. G ene flow a nd introgression f rom domesticated plants into their wild relatives. Annual Review of Ecology and Systematics 30: 539–563. Ellstrand N C, S chirenbeck K A. 2000. H ybridization a s a s timulus f or t he e volution of invasiveness in plants? Proc Natl Acad Sci USA 97:7043–7050. Frello S, Hansen KR, Jensen J, Jørgensen RB. 1995. Inheritance of rapeseed (Brassica napus) specific R APD m arkers an d a t ransgene i n t he cr oss Brassica j uncea×Brassica napus . 112
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UNIVERSITE DE BOURGOGNE THÈSE Pour
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REMERCIEMENTS My heartfelt thanks g
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forming many of the ideas I present
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Abstract In the framework of commer
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2.3 A rticle 2: Les r étrocroiseme
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Table 3.2. Parameters of linear reg
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experiment. BC1NS is separated into
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Fig. 4.7. Mean and standard error (
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INTRODUCTION GENERALE Dans le cadre
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chapitre 1 dressant l’état des c
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CHAPTER 1 LITERATURE REVIEW ON THE
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oilseed rape and B. oleracea is ver
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populations is reduced as the dista
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into hybrids would affect the morph
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Snow e t a l. ( 1999) f ound no f i
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1.5 Consequences of introgression I
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more flowers and larger plant size
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Moreover, beside the introgression
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their parental plants. Moon et al.
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2.1 Introduction CHAPTER 2 CONDITIO
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ARTICLE 1 The effect of seed size o
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plant growth was evaluated without
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These two experiments were kept wee
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interactions were found among the t
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2006) or m ore f it t han their pa
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they b ear b eneficial t ransgenes
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Diggle PK, Abrahamson NJ, Baker RL
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Ramachandran S , B untin G D, A ll
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Table 2.2. F -values f rom a f ive-
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Days to flowering Biomass Per. of s
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2.3 Article 2: Les rétrocroisement
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Photo.2.3. S praying chlorsulfuron
Page 65 and 66: Introduction In t he f ramework o f
Page 67 and 68: eceived p ollens f rom wild B. j un
Page 69 and 70: Statistical analysis Mean values ar
Page 71 and 72: Characteristics of the backcrosses
Page 73 and 74: Table 2.7. Mean (±95% CL) of plant
Page 75 and 76: plants w ith B. napus morphology t
Page 77 and 78: Productivity of the resistant proge
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Page 81 and 82: insight into adaptive life-history
Page 83 and 84: CHAPTER 3 EFFETS DE LA RESISTANCE A
Page 85 and 86: 3.2 A rticle 3 : S imulation d e l
Page 87 and 88: compétition et le devenir des popu
Page 89 and 90: difficult to predict. Similarly, it
Page 91 and 92: Table 3.1: ANOVA results of the eff
Page 93 and 94: Table 3.2: Parameters of linear reg
Page 95 and 96: herbivores than when exposed to Bt
Page 97 and 98: esistant hybrid populations might i
Page 99 and 100: Letourneau D .K., H agen J .A., 200
Page 101 and 102: Germination rate 0.6 0.5 0.4 0.3 0.
Page 103 and 104: 3.3 Article 4: Compétition entre p
Page 105 and 106: Introduction The invasion of transg
Page 107 and 108: Whether a popul ation w ill e volve
Page 109 and 110: while ensuring that the same ratio
Page 111 and 112: silique number, seed number, seed w
Page 113 and 114: Our ga rden e xperiments s howed t
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Page 119 and 120: density- dependent c osts of vi ral
Page 121 and 122: Table 3.5. F values of the analysis
Page 123 and 124: Table 3.7. Results of Helmert contr
Page 125 and 126: Fig. 3.4. Examples of experimental
Page 127 and 128: Flowering date Seed weight Biomass
Page 129 and 130: No. viable seeds Biomass 50000 4000
Page 131 and 132: Photo 3.3. Cotton bollworm (Helicov
Page 133 and 134: Introduction Spontaneous introgress
Page 135 and 136: screening as transgenic BC2 (trBC2)
Page 137 and 138: 2002), but certain studies showed n
Page 139 and 140: Acknowledgements We t hank Z hixi T
Page 141 and 142: 92, 368-374. Rieseberg, LH, and Bur
Page 143 and 144: Table 3.10. F-values of three-way A
Page 145 and 146: Seed number Biomass 140 120 100 80
Page 147 and 148: CHAPTER 4 RECHERCHE DES CONSEQUENCE
Page 149 and 150: Photo 4.1. Wild radish in a herbici
Page 151 and 152: Introduction Plant divergence is ge
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Page 155 and 156: estimated. All the seeds of every h
Page 157 and 158: Field experiment Number of siliques
Page 159 and 160: aphanistrum from hybrids and R. sat
Page 161 and 162: competition diminished the differen
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Page 165 and 166: Rattenbury J A. 1962. C yclic h ybr
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Fig. 4.1. Four r egions where w ild
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Fig. 4.3. Flower phot os showing pe
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Fig. 4.5. Plot of the first two axe
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Plant weight No. of seeds per plant
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Emergence rate Plant weight No.of a
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CONCLUSION 172
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véritable s uccès ad aptatif d an
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Dans l e cas d es p remières gén
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REFERENCES Abbott RJ, Comes HP.2007
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Bing D J, D owney R K and R akow G
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carinata and Brassica rapa. Plant B
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Hybridisation within Brassica and a
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etween weedy Brassica rapa and oils
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Hybridization between oilseed rape
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Nosil P. 2008. Speciation with gene
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32:305-32 Schadler, M., R. Brandl,
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Thalmann C, Guadagnuolo R, Felber F
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Zheng XM and Ge S. 2010. Ecological
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Therefore, after a review of the st
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more easily sieved out by harvester
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Transgenic F1 produced higher bioma
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