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1.1 Hybridizations between Brassica napus and its wild relatives In order to understand the possible risks of gene flow from Brassica napus to wild relatives, it is first important to understand the relationship between the species of its primary gene pool amongst others belonging to the Brassicaceae, consisting of over 3000 species in 370 genera. Three species, B. napus (AACC, 2n = 4x=38), B. juncea (AABB, 2n = 36), and B. carinata (BBCC, 2n = 34), are allotetraploids derived from three diploid species B. nigra (AA, 2n = 16), B. rapa (CC, 2n = 20), and B. oleracea (BB, 2n = 18) (Fig. 1.1). Fig. 1.1. “U-triangle” r epresenting t he genomic relationships b etweenBrassica s pecies ( redrawn from U 1935) Almost a ll h ybridization a mong B. napus and t he f ive ot her s pecies c ould oc cur, despite i t pr esented va rying di fficulty. T he h ybridization i s i nfluenced b y e xternal f actors, including w ind di rection a nd s peed, i nsect m ovement ( Scheffler e t al. 19 93), l ocations and flowers positions at plants (Becker et al. 1992). Natural crosses can frequently occur among the three oilseed Brassicas, B. napus, B. rapa and B. juncea (Bing et al. 1991,1996a; Leckie et al. 1993; Jorgensen and Andersen 1994), and transgene flow from B. napus into B. rapa (Bing et al. 1991, Leckie et al. 1993) or B. juncea (Bing et al. 1991, Frello et al. 1995) may occur t hrough n atural p ollination. A lthough t he f requency of s uccessful c rosses be tween 17
oilseed rape and B. oleracea is very low, hybridization could be achieved by hand pollination. In t he U nited K ingdom, s pontaneous h ybrids be tween B. napus and B. o leracea were observed i n t he B. oleracea wild popul ation, us ing f low c ytometry a nd c rop-specific microsatellite markers (Ford et al. 2006). No natural crosses are reported between the three species an d B. ni gra (Bing et a l. 1996 a), but male s terile B. napus produce s pontaneous hybrids w hen pl aced i n a B. ni gra stand ( Wei and D armency 2008) , although t here i s no indication a bout t he r eciprocal c ross. Hybrids f ormed be tween B. r apa and B. ni gra was obtained by ovule culture (Bing et al. 1996b). Hybridization of B. napus× B. carinata and B. juncea × B. carinata have been a chieved b y ha nd pollination (Chang et al. 2007). H ybrids between B. rapa and B. carinata were only obtained when B. carinata was used as the female parent (Choudhary et al. 2000). No natural crosses occurred between the three species and Sinapis arvensis (Bing et al. 1996a, Lefol et al. 1996), indicating that direct transgene escape from B. napus to S. arvensis appears very unlikely (Moyes et al. 2002).Spontaneous hybridization between B. napus and wild radish (Raphanus raphanistrum, Rr, 2n=18) could occur in the nature, and some hybrids were reported on male sterile B. napus as mother in Canada and Denmark (Eber et al. 1994; Baranger et a l. 1995; A mmitzboll a nd J orgensen 2006) a nd a t l ow f requency on R . raphanistrum as mother in Australia and France (Darmency et al. 1998; Chèvre et al. , 2000; Rieger et al. 2001 ). The frequency of hybridization depended on environmental conditions, oilseed rape variety, wild radish populations and their density (Eber et al. 1994; Baranger et al. 1995; Darmency et al. 1998; Chèvre et al. 1996, 2000; Rieger et al. 2001; Ammitzboll and Jorgensen 2006). Genetic polymorphism for pollen germination and ovule fertilisation also represented va riation of pr e-zygotic b arriers to in terspecific h ybridisation ( Guéritaine a nd Darmency 2001). Several ove rviews on h ybridization be tween B. napus and its w ild r elatives a nd associated species are available in Rieger et al. (1999), FitzJohn et al. (2007), Andersson and de Vicente (2010). 1.2 Dispersal of transgenes Extensive transgene flow between oilseed rape varieties has been detected in Canada resulting in the generation of volunteers that contain three types of herbicide resistance (Hall et al. 2000; Beckie et al. 2003; Knispel et al. 2008). The flow of transgenes could have occurred within a 18
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: CHAPTER 1 LITERATURE REVIEW ON THE
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 and 32: 1.5 Consequences of introgression I
Page 33 and 34: more flowers and larger plant size
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
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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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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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