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Jorgensen et al. (2007) showed that 6-32% volunteers of other varieties b elonged to varieties cultivated 4-17 years ago. In Canada, F1 and backcross hybrid generations between of he rbicide-resistant Brassica napus and B. r apa were d etected ove r-six years in th e field even in absence of the glyphosate selection pressure (Warwick et al. 2008). Herbicide-tolerant seedlings o f oi lseed r ape e merged t en years a fter a f ield t rial c onducted i n 1995, w hich confirmed t he l ong-term p ersistence o f t ransgenic s eeds i n t he s eed ba nk i n S weden (D’Hertefeldt e t a l. 200 8), a nd m ulti-herbicide-resistant oi lseed r ape vol unteers w ere found five to eight years after the last GM cultivar was grown in a multi-years farm-scale study in France (Méssean et al., 2007). Table 1.2. Long-term persistence of volunteers and feral plants in and/ or outside of cultivated fields References Years Sites Types GM GM rate Jorgensen et al. 2007 4-17 oilseed r fields ape Oilseed rape varieties Non 6-32% D’Hertefeldt e t a l. 10 oilseed r ape Herbicide-tolerant s eedlings Yes 39% 2008 fields from seed bank Warwick et al. 2008 6 oilseed r ape Herbicide-tolerant h ybrids Yes 2.5% fields between B.napus and B. rapa Méssean et al., 2007 5-8 Convention Herbicide-tolerant volunteers yes 0-18% oilseed r fields ape 1.3 Consequences of the hybridization Halfhill e t a l. ( 2005) f ound t hat h ybridization, w ith or w ithout Bt-transgene i ntrogression, resulted i n l ess ve getative gr owth a nd s eed pr oduction t han B. r apa and le ss c ompetitive populations. D i e t al. ( 2009) f ound t hat t ransgenic h ybrids be tween Bt-transgenic B. napus and B. juncea had intermediate composite fitness compared to their parents without herbivory. Generally, th e h ybrids s how in termediate mo rphologic characteristics c ompared t o their parents (e.g. leaf hairness of hybrids formed between B. napus and Hirschfeldia incan by Lefol e t a l. 1996; i ntermediate l eaf and f lower m orphology of h ybrids f ormed be tween B. carinata and B. rapa by Choudhary et al. 2000). The cytoplasm of crops or wilds introgress 21
into hybrids would affect the morphology of the progeny (e.g. Lefol et al. 1996; Zhang et al. 2002; Guéritaine et al. 2002; Chang et al. 2007). For example, after seven backcrossing of B. carinata to B. napus and B. juncea, plants with the cytoplasm of B. napus flowered later, had shorter filaments and longer pistils, lower pollen amount, lower seed set, lower petal length and w idth a nd di fferent petal c olor compared w ith pl ants w ith t he c ytoplasm of B. j uncea, suggesting that the cytoplasm is involved in the development of flower organs (Chang et al. 2007). In the case of B napus x R. raphanistrum crosses, F1 hybrids formed between oilseed rape and wild radish a re mainly allotriploids and present low f ertility (3. 2 s eeds/ plant for Darmency e t a l. 1998; 11 s eeds/ pl ant f or C hèvre e t a l.1998). T he F 1 h ybrids s howed significantly r educed s eedling emergence, emergence d elay, l ower survival r ate, an d decreased d ry m atter co mpared w ith t heir p arents ( Guéritaine et al. 2 003). H owever, t he progeny of F1 (F2, BC1) grew well and produced viable seeds (Darmency et al. 1998), and the fertility increased after repeated backcross to wild radish. In addition, the progeny having wild radish cytoplasm were more likely to propagate in agronomic and natural sites and had a fitness value 100 times more than that having oilseed rape cytoplasm (Guéritaine et al. 2002). 1.4 Consequences of the transgene A transgene transferred from oilseed rape to wild relatives might be neutral, but it is likely to increase o r d ecrease t he f itness o f r eceiving hybrid/ b ackcross p lants (Table 1.3) . T his depends on t he transgenic character, its selection and costs, how and where the transgene is introgressed in the genome, and the population composition (Felber et al. 2007). In response to insect herbivory, the Bt-transgene for insect resistance was shown to confer high relative fitness i n oi lseed r ape i n s everal s tudies ( Stewart e t a l. 1997; R amachandran e t a l. 2000; Letourneau and Hagen 2009). In turn, no fitness cost in the absence of the selection pressure was detected in the Brassiceae (Mason et al. 2003; Ramachandran et al. 2000; Di et al., 2009) and in sunflower (Snow et al. 2003). 22
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: populations is reduced as the dista
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
Page 75 and 76: 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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