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Table 1.5. M orphology was us ed t o m easure i ntrogression i n doc umented c ases of n atural hybridization and introgression of crops and wild relatives (One part of Table 1 in Jarvis and Hodgkin 1999) Crop References Technique(s) used to measure introgression Cabbages, mustards, rapes (Brassic spp.) Jorgensen & Andersen (1994) Morphology, c RAPDs ytology, i sozymes, Perrino & Hammer (1985) Morphology Snogerup et al. (1990) Morphology Stace (1991) Morphology Worede (1986) Morphology Radish (Raphanus sativus L.) Hammer & Perrino (1995) Morphology Klinger et al. (1992) Morphology, isozymes Klinger & Ellstrand (1994) Morphology The e volution of m orphology t raits i n a popul ation de pends a t l east i n pa rt on introgression and natural selection. For example, petal color of wild radish, a recessive singlelocus trait (Stanton et al. 1989), could be as an indirect evidence for crop-to-wild gene flow Lee and Snow 1998; Snow et al. 2001). Wild radish (Raphanus raphanistrum) in California had yellow petal color, whereas cultivated radish (R. sativus) and their hybrids had white to pink petals (Lee and Snow 1998; Kercher and Conner 1996). Butterflies (Pieris rapae) and syrphid f lies pr efer vi sit yellow f lowers ( Stanton e t a l. 1989; Lee a nd Snow 1998) , w hile bumble bees show a slight preference for white and pink colors (Lee and Snow 1998). Hence, pollinators could contribute to the loss of white petal allele when bumble bees were infrequent at f ields, a lthough S now e t a l. ( 2010) f ound t his a llele ha ve existed 10 year i n artificial populations. In a ddition, C ampbell e t a l. ( 2009) f ound t hat t he w hite pe tal i s l inked t o l ate flowering, a heritable trait. While selection acts against late-flowering plants that failed to set fruit or never bolted, this linkage also could lose the white petal allele. Lehtilä and Holmén Bränn ( 2007) f ound t hat t he l ines w ith l arger f lowers s tarted t o f lower later a nd pr oduced 27
more flowers and larger plant size than the lines with small flowers. Therefore, if white color petal allele is lost during the process of introgression when pollinators are selecting for yellow petals, l arger f lower an d l arger pl ant w ill be i nfrequent i n a popul ation. B ased on t his, i f transgenes related with white petal are transferred from the crop to the wild plant, the risk of transgene in trogression is lo w in f ields w here p ollinators s elect f or yellow p etals, a nd inversely, it is high. Thus, in turn, introgression of transgenes from crops to wild r elatives also could be influenced by morphological traits. Seed size is another example. Gene flow and introgression might be hampered by small seed size of hybrids between crops and wild relatives (Wei and Darmency 2008) because certain studies indicated that interspecific hybrids are present only among seeds with a diameter less than 1.6 m m (eg. Eber et al. 1994; Baranger et al. 1995; Chèvre e t a l. 2000) . For e xample, m ost crosses b etween o ilseed r ape ( Brassica napus ) a nd wild r elatives o nly p roduced s mall s eeds, s uch as B. j uncea (Bing et a l., 1996) , B. r apa (Jørgensen and Andersen, 1994), Hirschfeldia incana and R. raphanistrum (Eber et al., 1994; Chadoeuf et al., 1998). Wei and Darmency (2008) employed male sterile B. napus and five wild relatives to obtain hybrids and found all seeds of four hybrids were small. Small seeds perform d isadvantage b ecause s eed m ass affects t he em ergence and i nitial s eedling s ize (Aparicio et al. 2002; Westoby et al., 2002). This could decrease the plant fitness of smallseeded pl ants ( Gardner and va nderlip 1989; V erdu a nd T raveset, 2005) , e specially und er particular hazard conditions (such as high density, shade, drought or herbivory). 1.5.2 Consequences of interspecific introgression on population dynamics Gene f low a nd i ntrogression pl ays a c rucial r ole i n t he e volution of natural popul ation (Postma a nd N oordwijk 2005; Lenormand 2002 ), be cause gene f low c ould c ounteract t he negative effects of genetic drift, inbreeding (Ebert et al. 2002), genetic differentiation (Postma and Noordwijk 2005), and local selection (Lenormand 2002), on genetic variation and thus on the evolution of local adaptations. Over short time-scales, invading h ybrid populations may experience s trong s election ( Sakai e t a l., 20 01; A llendorf & Lundquist, 2003) . T he evolutionary t rajectory of w eed popul ations m ay persistently be a ltered b y t he l ong-term introgression of c rop alleles into weed populations, such as with sunflower (Whitton et al., 1997), oi lseed r ape ( Hansen e t a l., 2001) and r adish ( Snow e t al., 200 1). T he i nteraction 28
Page 1 and 2: UNIVERSITE DE BOURGOGNE THÈSE Pour
Page 3 and 4: REMERCIEMENTS My heartfelt thanks g
Page 5 and 6: forming many of the ideas I present
Page 7 and 8: Abstract In the framework of commer
Page 9 and 10: 2.3 A rticle 2: Les r étrocroiseme
Page 11 and 12: Table 3.2. Parameters of linear reg
Page 13 and 14: experiment. BC1NS is separated into
Page 15 and 16: Fig. 4.7. Mean and standard error (
Page 17 and 18: INTRODUCTION GENERALE Dans le cadre
Page 19 and 20: chapitre 1 dressant l’état des c
Page 21 and 22: CHAPTER 1 LITERATURE REVIEW ON THE
Page 23 and 24: oilseed rape and B. oleracea is ver
Page 25 and 26: populations is reduced as the dista
Page 27 and 28: into hybrids would affect the morph
Page 29 and 30: Snow e t a l. ( 1999) f ound no f i
Page 31: 1.5 Consequences of introgression I
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 and 78: Productivity of the resistant proge
Page 79 and 80: References [1] A .A. Snow, D .A. A
Page 81 and 82: 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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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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