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oth s howed f itness cost [ 25], but t here a re many ALS m utations e ndowing h erbicide- resistance s o t hat ge neralizations c ould not be made. In our e xperiment, t he r esistant B C1 (BC1NR) ha d hi gher frequency of B. napus morphology a nd l arger f lower s ize t han susceptible B C1 (BC1NS). P lants w ith B. napu s morphology a re m ore r obust, which c ould help BC1NR plants to establish in a mixed stand. Similarly, larger flowers like oilseed rape flowers c ould attract mo re f oraging in sects, w hich is a gain a w ay to b etter d isperse th e B. napus characteristics – and t he h erbicide-resistance – in r uderal a nd r oadside pl ant communities. As for the higher performances of BC1NS* compared to BC1NS, we suppose that the largest plants producing the highest number of flowers were more frequently visited by t he h ouseflies c arrying f oreign pol len, w hich c ould r esult i n t he pr oduction of r esistant seeds, i n c ontrast t o t he s mallest pl ants. T his ph enomenon c ould m ake t he di spersal of t he resistance gene more rapid again. Finally, the presence of herbicide-resistant plants in the progeny of susceptible F2, BC1N and conventional B. napus indicated that the first generation progeny of the hybrid is capable of tr ansmitting th e h erbicide-resistance gene t o B. napus volunteers and f eral popul ations, which provides an unexpected way to the transfer of transgenes via pollen dispersal from GM crops to non-GM crops [26]. It is also a source for further gene flow and introgression [8]. Our results show that the B. napus-like BC1 plants produced with B. napus pollen are v ery likely to establish and persist in the fields and along roadsides as false feral crop population. They look l ike B. napus volunteers but di splay additional ge netic va riability t hat a llow f or further adaptation of the plants and propagate the herbicide-resistance gene. In particular, they could have inherited from the wild B. juncea long-term seed viability in the soil, which can make them even weedier [2]. This phenomenon should be considered in biosafety assessment of genetically modified oilseed rape with herbicide-resistance traits. Acknowledgements The authors thank Alain FLEURY and Zhixi TANG for their assistance in the experiments. This work was supported by a PhD joint fellowship between France (CNOUS, No. 20072315) and China (Natural Science Foundation, grant No. 30970432). 73
References [1] A .A. Snow, D .A. A ndow, P . G epts, E .M. H allerman, A . P ower, J .M. T iedje, L.L. Wolfenbarger, G enetically engineered o rganisms and the environment: current status and recommendations, Ecol. Appl. 15 (2005) 377–404. [2] L.M. Hall, M.H. Habibur Rahman, R.H. Gulden, A.G. Thomas, Volunteer oilseed rape – Will h erbicide-resistance tr aits a ssist f erality? in : J.B. G ressel, ( Ed.), C rop f erality and volunteerism, CRC Press, Boca Raton, (2005), pp. 59-79. [3] S. Pivard, K. Adamczyk, J. Lecomte, C. Lavigne, A. Bouvier, A. Deville, P.H. Gouyon, S. Huet, Where do the feral oilseed rape populations come from? A large-scale study of their possible origin in a farmland area, J. Appl. Ecol. 45 (2008) 476-485. [4] A . M esséan, C . S ausse, J . G asquez, H . D armency, O ccurrence o f genetically m odified oilseed rape seeds in the harvest of subsequent conventional oilseed rape over time, Eur. J. Agron. 27 (2007) 115–122. [5] N .C. E llstrand, H .C. P rentice, J .F. H ancock, G ene flow a nd i ntrogression f rom domesticated plants into their wild relatives, Ann. Rev. Ecol. Syst. 30 (1999) 539–563. [6] J. Allainguillaume, M. Alexander, J.M. Bullock, M. Saunders, C.J. Allender, G. King, C.S. Ford, M .J. W ilkinson, F itness of h ybrids be tween r apeseed ( Brassica n apus) a nd w ild Brassica rapa in natural habitats, Mol. Ecol. 15 (2006) 1175–1184. [7] A .L. K nispel, S .M. McLachlan, R .C. Van A cker, L.F. Friesen, G ene f low a nd m ultiple herbicide resistance in escaped canola populations. Weed. Sci. 56 (2008) 72–80. [8] S.I. Warwick, A. Légère, M.J. Simard, T. James, Do escaped transgenes persist in nature? The case of an herbicide resistance transgene in a weedy Brassica rapa population, Mol. Ecol. 17 (2008) 1387–1395. [9] T.P. Hauser, R.B. Jorgensen, H. Ostergard, Fitness of backcross and F2 hybrids between weedy Brassica rapa and oilseed rape (B. napus), Heredity 81 (1998) 436–443. [10] B. Zhu, J.R. Lawrence, S.I. Warwick, P. Mason, L. Braun, M.D. Halfhill, C.N. Jr Stewart, Inheritance of GFP-Bt transgenes from Brassica napus in backcrosses with three wild B. rapa accessions, Environ. Biosafety. Res. 3 (2004) 45–54. [11] M.J. Crawley, R.S. Hails, M. Rees, D. Kohn, J. Buxton, Ecology of transgenic oilseed rape in natural habitats, Nature 363 (1993) 620-623. [12] S. F rello, K .R. H ansen, J . Jensen, R .B. J ørgensen, Inheritance of rapeseed ( Brassica napus) s pecific R APD markers an d a t ransgene i n t he cr oss Brassica j uncea×Brassica 74
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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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Page 29 and 30: Snow e t a l. ( 1999) f ound no f i
Page 31 and 32: 1.5 Consequences of introgression I
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Page 35 and 36: Moreover, beside the introgression
Page 37 and 38: their parental plants. Moon et al.
Page 39 and 40: 2.1 Introduction CHAPTER 2 CONDITIO
Page 41 and 42: ARTICLE 1 The effect of seed size o
Page 43 and 44: plant growth was evaluated without
Page 45 and 46: These two experiments were kept wee
Page 47 and 48: interactions were found among the t
Page 49 and 50: 2006) or m ore f it t han their pa
Page 51 and 52: they b ear b eneficial t ransgenes
Page 53 and 54: Diggle PK, Abrahamson NJ, Baker RL
Page 55 and 56: Ramachandran S , B untin G D, A ll
Page 57 and 58: Table 2.2. F -values f rom a f ive-
Page 59 and 60: Days to flowering Biomass Per. of s
Page 61 and 62: 2.3 Article 2: Les rétrocroisement
Page 63 and 64: 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: Productivity of the resistant proge
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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
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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 117 and 118: References Agrawal AA. 2004. R esis
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
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No. viable seeds Biomass 50000 4000
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Photo 3.3. Cotton bollworm (Helicov
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Introduction Spontaneous introgress
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screening as transgenic BC2 (trBC2)
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2002), but certain studies showed n
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Acknowledgements We t hank Z hixi T
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92, 368-374. Rieseberg, LH, and Bur
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Table 3.10. F-values of three-way A
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Seed number Biomass 140 120 100 80
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CHAPTER 4 RECHERCHE DES CONSEQUENCE
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Photo 4.1. Wild radish in a herbici
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Introduction Plant divergence is ge
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or t he popul ation of wild r adish
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estimated. All the seeds of every h
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Field experiment Number of siliques
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aphanistrum from hybrids and R. sat
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competition diminished the differen
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Conner, J. K., Rice A. M., Stewart
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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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