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and flower compared to other populations. It is necessary to find appropriate molecular markers of oilseed rape species for confirming the introgression in populations and genes that imply the modification of selected traits, for example flower color for attracting pollinators and silique form for dispersal and survival (Chapter 4.2). 2. Competition interaction and individual fitness As it is inevitable that populations composed transgenic and non-transgenic individuals after the occurrence of introgressions (Ramachandran et al. 2000; Vacher et al. 2004; Moon et al. 2007), their relative competition capacity will affect the individual growth and reproduction and further affect population dynamics. Insect-resistant plants had competitive advantage when competing with insect- susceptible plants under insects. This relative advantage was increased with the percentage of healthy plants in susceptible populations. However, as the increasing of percentage of resistant plants in populations, the intra-class competition among resistant plants affects their inter-class competition between resistant and susceptible plants. In addition, this competition advantage of resistant plants was magnified under harsh conditions, like limiting resources, high plant density. The effects of competition and herbivory on plant fitness were additive. Competition interaction among plants in populations was changed because of the presence of introgressed resistant plants. (Chapter 3.3) In case of the introgression succeed, the transgene will be treated as other genes in wild relatives, and it could arrive at equilibrium situation or fixed or disappeared that depends on the selection pressure. Hence, which traits and in which condition a transgene confers fitness advantage should be detected. A transgene transferred from transgenic crops to wild relatives might be neutral, but it is likely to increase or decrease the fitness of receiving hybrid/ backcross plants (Stewart et al. 1997; Ramachandran et al. 2000; Mason et al. 2003; Di et al., 2009; Letourneau and Hagen 2009). This depends on the transgenic character, its selection and costs, the context of transgene introgressed in the genome, and the population composition. 201
Transgenic F1 produced higher biomass, lower seed number and weight and reproductive allocation than mustard and transgenic oilseed rape (Chapter 2.2). Herbicide- resistant BC1 was associated with greater plant and produced more seeds than their susceptible counterparts (Chapter 2.3). Insect-resistant plants produced higher plant fitness: higher biomass and seed weight and more seeds (Chapter 3.3). However, Bt-transgenic BC2 showed similar fitness compared to susceptible BC2 when they were planted together in the presence of insects because of the protection of transgenic plants on non-transgenic plants (“halo effect”) (Chapter 3.4). The ancient introgressed wild radish plants did not show significant higher fitness but intermediate values compared with others (Chapter 4). Moreover, the fitness effects of transgenes were affected by other factors, such as morphological traits (seed size, flower color etc.), competition, herbivory, and resources availability. Large-seeded plants in transgenic oilseed rape showed higher plant fitness than small-seeded ones (Chapter 2). Ancient introgressed wild radish plants with white petal produced more seeds than plants with white petal (Chapter 4). Increased plant density significantly decreased plant biomass and seed output (Chapter 2). Competition magnifyied the fitness advantage of the insect-resistant plants, but as their frequency increases, neighbor competition limits their growth. High resources availability and low herbivory induced compensatory growth in susceptible plants, and accordingly decreased the fitness differences between susceptible and resistant plants (Chapter 3.3). The mixed cultivation with Bt-transgenic plants and non-transgenic plants increased production of both transgenic and non-transgenic plants (Chapter 3.4). 3. Population production Population equilibrium might be broken in case of the invasion of transgenic plants or the transferring of transgenes in wild populations because of the competition interaction between transgenic and non-transgenic plants. In long-term, population might diverge from others after successively introgression because of the trasgenes conferring new traits. The total vegetative and reproductive production of mixed populations of healthy and damaged plants was the same as that of pure populations of either plant type, but 202
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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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compétition et le devenir des popu
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difficult to predict. Similarly, it
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Table 3.1: ANOVA results of the eff
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Table 3.2: Parameters of linear reg
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herbivores than when exposed to Bt
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esistant hybrid populations might i
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Letourneau D .K., H agen J .A., 200
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Germination rate 0.6 0.5 0.4 0.3 0.
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3.3 Article 4: Compétition entre p
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Introduction The invasion of transg
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Whether a popul ation w ill e volve
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while ensuring that the same ratio
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silique number, seed number, seed w
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Our ga rden e xperiments s howed t
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experiment, harsh growth conditions
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References Agrawal AA. 2004. R esis
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density- dependent c osts of vi ral
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Table 3.5. F values of the analysis
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Table 3.7. Results of Helmert contr
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Fig. 3.4. Examples of experimental
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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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Page 157 and 158: Field experiment Number of siliques
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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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Page 189 and 190: Hybridisation within Brassica and a
Page 191 and 192: etween weedy Brassica rapa and oils
Page 193 and 194: Hybridization between oilseed rape
Page 195 and 196: Nosil P. 2008. Speciation with gene
Page 197 and 198: 32:305-32 Schadler, M., R. Brandl,
Page 199 and 200: Thalmann C, Guadagnuolo R, Felber F
Page 201 and 202: Zheng XM and Ge S. 2010. Ecological
Page 203 and 204: Therefore, after a review of the st
Page 205: more easily sieved out by harvester
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