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the da maged nt rBC2 pl ants pos sibly i nducing defense m echanisms a nd m oved t owards healthy and attractive trBC2 plants. Alternately, it c ould be a further evidence o f the lower innate fitness of the transgenic plants, leaving more and more resources to their neighbor. Population production with varied proportion of transgenic plants Population production, biomass and seed output, increased with the percentage of transgenic BC2 in populations under insect pressure, perhaps because of the generalized protection effect as proposed above. It was not the case when the populations were not subjected to herbivory, again w ith a m aximum of s eed pr oduction at a 50% pr oportion of t ransgenic pl ants. Th e increased population production in the presence of insects is important to predict the increase of the colonizing ability of the B. juncea populations hosting more and more insect-resistant plants. F ew s tudies f ocused on the d ynamics of resistant pl ants i n s usceptible popul ations, although some studies showed that herbivory strongly affected population dynamics of plants (e.g. P almisano a nd Fox 1997) . R amachandran e t a l. (2000) s howed t hat i nsects d ecreased total biomass but did not affect seed yield in mixed and pure stands of Bt-transgenic and/ or non-transgenic oi lseed r ape. T hey found t hat no s ignificant di fferences oc curred i n t otal biomass and seed production when there was no i nsect, and that biomass and seed yield was significantly different among mixed plantings for insect-infested plots with a maximum at the proportion of 50% i n t wo f ield e xperiments, but t hat i t w as not di fferent i n a nother f ield experiment in the presence of insects. In our e xperiments simulating insect-resistant plants through clipping leaves, the increase of healthy plants in a population of damaged plants did not increased the population production but resulted in a maximum seed output when healthy plants dom inated t he popul ation ( 75% of he althy plants) (Liu e t a l.2010b). E nhanced s eed production c ould m ake a popul ation m ore a ble t o s pread t o ne ighboring ha bitats, e venly displacing other species suffering of insect damages. Thus, the occurrence of insect attack is the major determinant of the fate and dispersal of the transgene: while pests are over-frequent in ar able l ands co vered b y the s ame cr op o ver h ectares, t he o ccurrence o f s uch i nsect infestations is not so permanent in wild habitats, which could seriously hamper the spread of the Bt transgene in and among wild populations, as observed in the case of virus-resistance genes i n s quash ( Fuchs e t a l 2004a ; 2004b) . Laughlin et a l. ( 2009) showed t hat vi rusresistance transgenes and conventional crop genes can increase fecundity of wild plants under virus pressure, and viruses play a role in the population dynamics of wild squash (Cucurbita pepo). 133
Acknowledgements We t hank Z hixi T ang, X in J ing a nd H anbin W u for th eir a ssistances in e xperiment manipulating. This work is supported by a project of the Natural Science Foundation of China (grant no. 30970432) and a PhD joint fellowship between China and France (CNOUS, No. 20072315). References Agrawal AA. 1998. Induced responses to herbivory and increased plant performance. Science, 279:1201–1202. Agrawal A A. 199 9. Induced r esponses t o he rbivory i n w ild r adish: effects on s everal herbivores and plant fitness. Ecology, 80: 1713–1723 Al Mouemar A and Darmency H. 2004. Lack of stable inheritance of introgressed transgene from oilseed rape in wild radish. Environ. Biosafety Res. 3, 209–214 Arnold M L, Hodges S A ( 1995) A re na tural h ybrids f it or unf it r elative t o t heir pa rents ? Trends in Ecology and Evolution, 10, 67–71. Boalt E. and Lehtilâ K. 2007. Tolerance to apical and foliar damage: costs and mechanisms in Raphanus raphanistrum. Oikos 116: 2071-2081 Burke JM, and Arnold ML. 2001. Genetics and the fitness of hybrids. Annu. Rev. Genet. 35: 31-52 Campbell LG, and Snow AA. 2007. Competition alters life history and increases the relative fecundity of crop-wild radish hybrids (Raphanus spp.). New Phytologist 173: 648–660. Darmency, H. 1994. The impact of hybrids between genetically modified crop plants and their related species: introgression and weediness. Molecular Ecology. 3, 37–40. 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. Escarré J, Lepart, J, Sans X, Sentuc JJ. and Gorse V. 1999. Effects of herbivory on the growth and reproduction of Picris hieracioides in the Mediterranean region. Journal of Vegetation Science 10: 101-110 Frello S, Hansen KR, Jensen J, Jørgensen RB. 1995. Inheritance of rapeseed (Brassica napus) 134
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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
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Introduction In t he f ramework o f
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eceived p ollens f rom wild B. j un
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Statistical analysis Mean values ar
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Characteristics of the backcrosses
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Table 2.7. Mean (±95% CL) of plant
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plants w ith B. napus morphology t
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Productivity of the resistant proge
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References [1] A .A. Snow, D .A. A
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insight into adaptive life-history
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CHAPTER 3 EFFETS DE LA RESISTANCE A
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3.2 A rticle 3 : S imulation d e l
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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
Page 115 and 116: experiment, harsh growth conditions
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
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: 2002), but certain studies showed n
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
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Page 161 and 162: competition diminished the differen
Page 163 and 164: Conner, J. K., Rice A. M., Stewart
Page 165 and 166: Rattenbury J A. 1962. C yclic h ybr
Page 167 and 168: Fig. 4.1. Four r egions where w ild
Page 169 and 170: Fig. 4.3. Flower phot os showing pe
Page 171 and 172: Fig. 4.5. Plot of the first two axe
Page 173 and 174: Plant weight No. of seeds per plant
Page 175 and 176: Emergence rate Plant weight No.of a
Page 177 and 178: CONCLUSION 172
Page 179 and 180: véritable s uccès ad aptatif d an
Page 181 and 182: Dans l e cas d es p remières gén
Page 183 and 184: REFERENCES Abbott RJ, Comes HP.2007
Page 185 and 186: Bing D J, D owney R K and R akow G
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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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