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ARTICLE 6 The role of long-term coexistence with oilseed rape on the morphological and life-history trait variability of Raphanus raphanistrum populations Yongbo LIU, Wei WEI, KePing MA, AM CHEVRE, Henri DARMENCY Summary • Plant di vergence a nd adaptation i s c hallenged b y the oc currence of gene f low and introgression f rom r elated s pecies. A pa rticular c ase c onsists i n g ene f low f rom genetically modified (GM) crops to their wild relatives. • Wild radish (Raphanus raphanistrum) populations were sampled from geographically distant r egions, of w hich one ha d a l ong hi story of oi lseed r ape c ultivation. P lants were cultivated together, and their morphological, growth and reproductive traits were measured. • The morphological traits discriminated the wild radish populations from the different regions. T he popul ations w ere di fferent f or growth a nd r eproduction a mongst t hem, but the differences changed when the plants were grown in an oilseed rape field or in intra-specific competition condition. • The geographic distance cannot account for all the differences among regions. There is no direct evidence of introgression with oilseed rape, but this hypothesis could fit the data. The longer coexistence time one wild radish populations with oilseed rape in one region had no dramatic effect on the studied traits. Key words: introgression, life history, morphology divergence, natural selection, wild radish 145
Introduction Plant divergence is generated by isolation by distance (genetic drift) or by changes in mating systems and adaptation driven via natural selection (Hey 2006; Ross-Ibarra et al. 2009; Zheng and G e 2010) . In c ontrary, gene f low pl ays a ke y r ole i n m aintaining c ohesion a mong geographically s eparated popul ations a nd m akes s pecies t o e volve a s uni tes ( Morjan and Rieseberg 2004; Abbott and Comes 2007; Hey 2006; Nosil 2008). Introgression of wild plants by genes from crops could result in poor divergence when gene flow is strong enough, with no correlation b etween genetic d ifferences and geographic d istances (Kercher a nd C onner 1996; Sahli et al. 2008). Although it is a natural phenomenon associated to crop development, gene flow and introgression f rom c rops to th eir w ild r elatives r eceive n ow in creasing a ttention in th e framework of the commercial release of genetically modified (GM) crops. Because GM crops confer new traits, such as resistance to insects, herbicides and diseases, they could result in undesired consequences of e ventual t ransgene i ntrogression t hrough t he di spersal of interspecific h ybrids and their progeny. Introgression after hybridization between GM crops and their wild relatives may result in increasing: (1) genetic variation that promotes adaptive evolution ( Rattenbury 1 962; P ostma a nd V an N oordwijk 2005), ( 2) weediness of w ild species ( Darmency 199 4; P ilson a nd P rendeville 2004) , ( 3) c ompetitiveness or c olonizing abilities through hybrid vigor or new traits (Ellstrand et al. 1999; Ramachandran et al. 2000), (4) i nvasiveness vi a e nhanced f itness ( Vacher et a l. 2004) , a nd ( 5) a bility t o m atch new environment (Campbell et al. 2006; Whitney et al. 2006), which eventually could result in the displacement o f n atural c ommunity e quilibriums, in cluding th e elimination o f d esirable species through competition (Tiedje et al. 1989). Several studies have found long-term introgression from crops to their wild relatives, such as in rice (Suh et al. 1997), radish (Snow et al. 2001; Hedge et al. 2006), oilseed rape (Hansen et al. 2001) and sunflower (Whitton et al. 1997; Linder et al. 1998). The crop-wild introgression ma y a lter morphological a nd lif e history tr aits a s w ell as th e e volutionary trajectory of w ild popul ations ( Lee a nd S now1998; S now e t a l. 2001; Lenormand 2002; Campbell e t a l. 2009a ), w hich c ould l ead t o popul ations e xhibiting pe culiar t raits a nd population dynamics. In addition, local natural selection exerted on new genetic backgrounds could make introgressed wild populations to converge to or diverge from their parental taxa, 146
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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
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
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Page 137 and 138: 2002), but certain studies showed n
Page 139 and 140: Acknowledgements We t hank Z hixi T
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: Photo 4.1. Wild radish in a herbici
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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
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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
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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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