<str<strong>on</strong>g>Proceedings</str<strong>on</strong>g> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> <str<strong>on</strong>g>Third</str<strong>on</strong>g> <str<strong>on</strong>g>Internati<strong>on</strong>al</str<strong>on</strong>g> <str<strong>on</strong>g>C<strong>on</strong>ference</str<strong>on</strong>g> <strong>on</strong> <strong>Invasive</strong> SpartinaChapter 1: Spartina Biology<str<strong>on</strong>g>the</str<strong>on</strong>g> same chloroplast genome as S. alterniflora, which wasc<strong>on</strong>sidered <str<strong>on</strong>g>the</str<strong>on</strong>g> maternal genome d<strong>on</strong>or when it hybridizedwith S. maritima (Ferris et al. 2001).Ano<str<strong>on</strong>g>the</str<strong>on</strong>g>r sterile hybrid has been described in southwestFrance (Basque regi<strong>on</strong>) <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> Spanish border and has beencalled S. x neyrautii (Foucault, 1897). As <str<strong>on</strong>g>the</str<strong>on</strong>g> two hybrids S.x neyrautii and S. x townsendii display markedmorphological differences, <str<strong>on</strong>g>the</str<strong>on</strong>g>y were first c<strong>on</strong>sidered asreciprocal hybrids (Marchant 1977). However, Baumel et al.(2003) found that S. x neyrautii also has <str<strong>on</strong>g>the</str<strong>on</strong>g> same chloroplastgenome as S. alterniflora, which indicates that hybridizati<strong>on</strong>occurred in <str<strong>on</strong>g>the</str<strong>on</strong>g> same directi<strong>on</strong> (with S. maritima as <str<strong>on</strong>g>the</str<strong>on</strong>g> maleparent) in England and in southwest France. Both parentalspecies (S. maritima and S. alterniflora) have been foundgenetically depauperate in western Europe (Raybould et al.1991a; Baumel et al. 2003; Yannic et al. 2004), so <str<strong>on</strong>g>the</str<strong>on</strong>g> twohybrids have inherited similar parental genotypes that maybe distinguished by a few molecular markers (Baumel et al.2003). Moreover, Salm<strong>on</strong> et al. (2005) have found that <str<strong>on</strong>g>the</str<strong>on</strong>g>setwo taxa display very similar genetic and epigeneticdynamics.THE YOUNG INVASIVE S. ANGLICAThe neoallopolyploid S. anglica is a perfect example <str<strong>on</strong>g>of</str<strong>on</strong>g><str<strong>on</strong>g>the</str<strong>on</strong>g> fitness that can be immediately gained followingduplicati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> a hybrid genome: This species displays largerecological amplitude than its parents; as a pi<strong>on</strong>eer species, itis able to col<strong>on</strong>ize a vacant niche fur<str<strong>on</strong>g>the</str<strong>on</strong>g>r down <str<strong>on</strong>g>the</str<strong>on</strong>g> shore,and to tolerate several hours <str<strong>on</strong>g>of</str<strong>on</strong>g> immersi<strong>on</strong> under sea water(Thomps<strong>on</strong>, 1991). Spartina anglica is characterized byhigher physiological tolerance to anoxic soil c<strong>on</strong>diti<strong>on</strong>s, andit is able to enhance <str<strong>on</strong>g>the</str<strong>on</strong>g> sediment oxygenati<strong>on</strong> (Lee 2003).This species now has a worldwide distributi<strong>on</strong> (northwesternEurope, China, Australia, North America), as aresult <str<strong>on</strong>g>of</str<strong>on</strong>g> both natural propagati<strong>on</strong> and deliberateintroducti<strong>on</strong>s for land reclamati<strong>on</strong>. This propagati<strong>on</strong> isfacilitated by high seed producti<strong>on</strong> and impressivevegetative features such as str<strong>on</strong>g rhizomes, that increase <str<strong>on</strong>g>the</str<strong>on</strong>g>sediment accumulati<strong>on</strong>. The rapid spread <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> introducedplants has lead to various attempts to c<strong>on</strong>trol or eradiate <str<strong>on</strong>g>the</str<strong>on</strong>g>species (Hedge et al. 2010; Hacker and Dethier 2010).However, after a variable period <str<strong>on</strong>g>of</str<strong>on</strong>g> rapid expansi<strong>on</strong>, S.anglica populati<strong>on</strong>s seem to experience “dieback” in oldercol<strong>on</strong>ized sites, which is probably caused by age-relateddecline <str<strong>on</strong>g>of</str<strong>on</strong>g> vigor or by competiti<strong>on</strong> with o<str<strong>on</strong>g>the</str<strong>on</strong>g>r species afterniche elevati<strong>on</strong> (Gray 2010; An et al. 2010).Spartina anglica benefits from <str<strong>on</strong>g>the</str<strong>on</strong>g> reuni<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> twodivergent (homeologous) genomes from S. alterniflora andS. maritima (Baumel et al. 2002a), <str<strong>on</strong>g>the</str<strong>on</strong>g>refore it ischaracterized by high levels <str<strong>on</strong>g>of</str<strong>on</strong>g> intergenomic heterozygosityat nuclear loci (Guenegou et al. 1988; Raybould et al.1991b). However, c<strong>on</strong>trasting with most o<str<strong>on</strong>g>the</str<strong>on</strong>g>r known cases<str<strong>on</strong>g>of</str<strong>on</strong>g> recent alloploid formati<strong>on</strong> that have been explored withmolecular markers (Soltis et al. 2004; Abbott and Lowe2004), S. anglica has underg<strong>on</strong>e a genetic bottleneck as aresult <str<strong>on</strong>g>of</str<strong>on</strong>g> ei<str<strong>on</strong>g>the</str<strong>on</strong>g>r a unique hybridizati<strong>on</strong> event, or multipleevents involving similar parental genotypes (Ainouche et al.2004b). The populati<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> S. anglica display c<strong>on</strong>sistentmorphological variati<strong>on</strong>, particularly in differentsuccessi<strong>on</strong>al stages where pi<strong>on</strong>eer populati<strong>on</strong>s have beenfound to exhibit phenotypes with smaller plants and smallerinflorescence than in mature populati<strong>on</strong>s (Thomps<strong>on</strong> et al.1991a). This variati<strong>on</strong> has been found to be caused mainlyby phenotypic plasticity ra<str<strong>on</strong>g>the</str<strong>on</strong>g>r than by geneticdifferentiati<strong>on</strong> (Thomps<strong>on</strong> et al. 1991b, c).The British populati<strong>on</strong>s appear genetically depauperatefor allozyme markers (Raybould et al. 1991b). Baumel et al.(2001) have explored <str<strong>on</strong>g>the</str<strong>on</strong>g> populati<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina anglica inFrance using RAPD (Random Amplificati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> PolymorphicDNA) and ISSR (InterSimple Sequence Repeat) markers,and have found that most populati<strong>on</strong>s are composed <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>same major genotype that is identical to <str<strong>on</strong>g>the</str<strong>on</strong>g> first generati<strong>on</strong>hybrid S. x townsendii. Table 1 summarizes molecularinvestigati<strong>on</strong>s that have been c<strong>on</strong>ducted in populati<strong>on</strong>s fromboth <str<strong>on</strong>g>the</str<strong>on</strong>g> native range <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> species (western Europe) andmore recently col<strong>on</strong>ized c<strong>on</strong>tinents (Australia). Most <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>introduced populati<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> S. anglica in <str<strong>on</strong>g>the</str<strong>on</strong>g> world arebelieved to have originated from Pool Harbor, sou<str<strong>on</strong>g>the</str<strong>on</strong>g>rnEngland (Hubbard 1965). In sou<str<strong>on</strong>g>the</str<strong>on</strong>g>rn England, S. xtowsendii has been sampled in Hy<str<strong>on</strong>g>the</str<strong>on</strong>g> (for comparis<strong>on</strong> withS. anglica genotypes), and <str<strong>on</strong>g>the</str<strong>on</strong>g> populati<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> S. anglica havebeen sampled in Lymingt<strong>on</strong>, <str<strong>on</strong>g>the</str<strong>on</strong>g> former site where S. anglicawas recorded, Fawley (near Hy<str<strong>on</strong>g>the</str<strong>on</strong>g>), Keyhaven, Sand Bank,and Studland (Pool Harbour). Samples from Ireland (BullIsland) were also analyzed. Forty-three out <str<strong>on</strong>g>of</str<strong>on</strong>g> 45 S. anglicasamples from England and Ireland have <str<strong>on</strong>g>the</str<strong>on</strong>g> same multilocusRAPD genotype (Table 1); <strong>on</strong>ly two individuals exhibitdifferent genotypes: <strong>on</strong>e individual, sampled in Fawleydiffers from <str<strong>on</strong>g>the</str<strong>on</strong>g> major genotype by <str<strong>on</strong>g>the</str<strong>on</strong>g> absence <str<strong>on</strong>g>of</str<strong>on</strong>g> <strong>on</strong>eRAPD marker, whereas two o<str<strong>on</strong>g>the</str<strong>on</strong>g>r individuals fromLymingt<strong>on</strong> are distinguished by <str<strong>on</strong>g>the</str<strong>on</strong>g> loss <str<strong>on</strong>g>of</str<strong>on</strong>g> two DNAfragments. A few ISSR, IRAP and REMAP markers arepolymorphic (Table 1), which allowed Baumel et al. (2002b)to distinguish 13 genotypes that were encountered at lowfrequency in some populati<strong>on</strong>s; <str<strong>on</strong>g>the</str<strong>on</strong>g>se genotypes are verysimilar to <str<strong>on</strong>g>the</str<strong>on</strong>g> major <strong>on</strong>e, as most <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>m differ by <strong>on</strong>ly <strong>on</strong>e(lost) marker.The French populati<strong>on</strong>s have been sampled in variouslocati<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> Atlantic coast (Baumel et al. 2001),including Baie des Veys (Normandy) <str<strong>on</strong>g>the</str<strong>on</strong>g> first col<strong>on</strong>ized sitein France where S. anglica was recorded in 1906. Mostindividuals sampled in France also bel<strong>on</strong>g to <str<strong>on</strong>g>the</str<strong>on</strong>g>predominant genotype. One small populati<strong>on</strong> from Kerdruc(sou<str<strong>on</strong>g>the</str<strong>on</strong>g>rn Brittany) was found fixed for two mutati<strong>on</strong>s, with<str<strong>on</strong>g>the</str<strong>on</strong>g> absence <str<strong>on</strong>g>of</str<strong>on</strong>g> <strong>on</strong>e RAPD (Baumel et al. 2001) and <strong>on</strong>eISSR marker (this study). The few slightly different o<str<strong>on</strong>g>the</str<strong>on</strong>g>rgenotypes (Table 1) were found to be randomly distributed,and at low frequency, am<strong>on</strong>g <str<strong>on</strong>g>the</str<strong>on</strong>g> o<str<strong>on</strong>g>the</str<strong>on</strong>g>r French populati<strong>on</strong>s.In Saint-Armel (sou<str<strong>on</strong>g>the</str<strong>on</strong>g>rn Brittany), S. anglica growssympatrically with its parental species S. maritima.-17-
Chapter 1: Spartina Biology<str<strong>on</strong>g>Proceedings</str<strong>on</strong>g> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> <str<strong>on</strong>g>Third</str<strong>on</strong>g> <str<strong>on</strong>g>Internati<strong>on</strong>al</str<strong>on</strong>g> <str<strong>on</strong>g>C<strong>on</strong>ference</str<strong>on</strong>g> <strong>on</strong> <strong>Invasive</strong> SpartinaTable 1: Inter-individual molecular variati<strong>on</strong> in native and introduced Spartina anglica populati<strong>on</strong>s investigated using three multilocus methods:RAPD (Random Amplificati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> Polymorphic DNA, 14 primers), ISSR (Inter Simple Sequence Repeats, 4 primers), IRAP (Inter-Retrotranspos<strong>on</strong> Amplified Polymorphism, 7 primer pairs), REMAP (Retrotranspos<strong>on</strong> Microsatellite Amplified Polymorphism, 4 primer pairs).The mean individual number sampled per populati<strong>on</strong> = 10. N= number <str<strong>on</strong>g>of</str<strong>on</strong>g> samples analyzed per regi<strong>on</strong>; pm = polymorphic markers; m=number<str<strong>on</strong>g>of</str<strong>on</strong>g> markers recorded; mg = number <str<strong>on</strong>g>of</str<strong>on</strong>g> genotypes that differ from <str<strong>on</strong>g>the</str<strong>on</strong>g> “major” genotype identified by Baumel et al. (2001) and identical to S. xtownsendii.RAPD ISSR IRAP/REMAPN pm/ m mg/N N pm/m mg/N N pm/m mg/NEngland &Ireland45 3/146 2/45 20 0/1440 b 3/920/204/40 40 b 10/296 9/40France129 a 6/146 6/12920129 a 2/921/142/201/1295 2/296 2/5Australia 20 0/146 0/20 160 0/21 0/160 - - -a = data from Baumel et al. (2001); b = data from Baumel et al. (2002b).Intermediate, morphologically variable individuals may beencountered (Guenegou and Levasseur, pers<strong>on</strong>alcommunicati<strong>on</strong>), but no genetic exchange was foundbetween <str<strong>on</strong>g>the</str<strong>on</strong>g> two species according to cytological, allozyme(M.T. Misset and G. Allard, unpublished data) andmolecular (Baumel et al. 2001) data.In Australia, 160 samples were collected in various sitesfrom Victoria (Corner Inlet including <str<strong>on</strong>g>the</str<strong>on</strong>g> mouth <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>Albert River and Anders<strong>on</strong> Inlet, Gippsland) and Tasmania(Franklin River in Port Sorell Bay, Duck River nearSmitht<strong>on</strong>, Perkins Bay, Tamar River and Little Swanportestuary). In Victoria, various attempts at eradicati<strong>on</strong> havebeen performed using <str<strong>on</strong>g>the</str<strong>on</strong>g> herbicide Fusillade; we havesampled n<strong>on</strong>-treated z<strong>on</strong>es and also collected rare survivingplants from treated areas. Morphological variati<strong>on</strong> is alsoencountered in <str<strong>on</strong>g>the</str<strong>on</strong>g> Australian populati<strong>on</strong>s where two distinctphenotypes (short plants <str<strong>on</strong>g>of</str<strong>on</strong>g> 10 – 20 cm high <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> <strong>on</strong>e handand taller plants measuring up to 30 cm <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> o<str<strong>on</strong>g>the</str<strong>on</strong>g>r hand)were collected in <str<strong>on</strong>g>the</str<strong>on</strong>g> Albert River (Victoria). In LittleSwanport, 20 seedlings have been collected in order toanalyse samples resulting unambiguously from sexualreproducti<strong>on</strong>. We found that all <str<strong>on</strong>g>the</str<strong>on</strong>g> individuals investigated inAustralia and Tasmania display <str<strong>on</strong>g>the</str<strong>on</strong>g> same multilocus RAPD orISSR genotype (Table 1), which is identical to <str<strong>on</strong>g>the</str<strong>on</strong>g> majorgenotype encountered in Europe and to S. x townsendii.It appears that populati<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> S. anglica in both itsnative range and more recently col<strong>on</strong>ized areas are mainlycomposed <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> major genotype that is identical to <str<strong>on</strong>g>the</str<strong>on</strong>g> firstgenerati<strong>on</strong> hybrid S. x townsendii. Some variati<strong>on</strong> may beencountered, however, that result from a few mutati<strong>on</strong>soccurring in local populati<strong>on</strong>s and corresp<strong>on</strong>ding in mostcases to fragment loss. Ayres and Str<strong>on</strong>g (2001) also notedfragment loss in S. anglica. They encountered variati<strong>on</strong> inRAPD and ISSR analyses including samples <str<strong>on</strong>g>of</str<strong>on</strong>g> S. anglicafrom England and Australia, and observed variable bandlossat five S. maritima-specific loci. However, Salm<strong>on</strong> et al.(2005) have reported preferential fragment loss <str<strong>on</strong>g>of</str<strong>on</strong>g> S.alterniflora-specific markers using AFLP (AmplifiedFragment Length Polymorphism) in both S. x townsendii andS. anglica. It should be noted that <str<strong>on</strong>g>the</str<strong>on</strong>g> estimati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> diversitymay be ei<str<strong>on</strong>g>the</str<strong>on</strong>g>r underestimated or overestimated according toboth sampling (number <str<strong>on</strong>g>of</str<strong>on</strong>g> individuals analysed and number<str<strong>on</strong>g>of</str<strong>on</strong>g> markers examined over <str<strong>on</strong>g>the</str<strong>on</strong>g> genome) and <str<strong>on</strong>g>the</str<strong>on</strong>g> techniqueused: We can see (Table 1) that although globally limited,more variati<strong>on</strong> can be detected from ISSR and IRAP–REMAP markers, that target repetitive (Simple SequenceRepeats and Retrotranspos<strong>on</strong>s), potentially more variable,parts <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> genome. Moreover, c<strong>on</strong>sidering that <str<strong>on</strong>g>the</str<strong>on</strong>g>se PCRbasedmarkers are dominant, variati<strong>on</strong> resulting fromsegregating heterozygous genotypes cannot be ruled out.Ano<str<strong>on</strong>g>the</str<strong>on</strong>g>r potential source <str<strong>on</strong>g>of</str<strong>on</strong>g> variati<strong>on</strong> is <str<strong>on</strong>g>the</str<strong>on</strong>g> structuraldynamic that may affect recently formed allopolyploidgenomes (Wendel 2000), but in <str<strong>on</strong>g>the</str<strong>on</strong>g> case <str<strong>on</strong>g>of</str<strong>on</strong>g> S. anglica, thisdynamic does not seem to affect <str<strong>on</strong>g>the</str<strong>on</strong>g> genotype initiallyformed in England to <str<strong>on</strong>g>the</str<strong>on</strong>g> extent that was reported in <str<strong>on</strong>g>the</str<strong>on</strong>g>literature in o<str<strong>on</strong>g>the</str<strong>on</strong>g>r allopolyploid model systems (Baumel etal. 2002b; Ainouche et al. 2004b).We have recently explored <str<strong>on</strong>g>the</str<strong>on</strong>g> epigenetic alterati<strong>on</strong>sthat may affect <str<strong>on</strong>g>the</str<strong>on</strong>g> allopolyploid genome <str<strong>on</strong>g>of</str<strong>on</strong>g> S. anglica(Salm<strong>on</strong> et al. 2005). Epigenetic mechanisms causeexpressi<strong>on</strong> changes that do not result from <str<strong>on</strong>g>the</str<strong>on</strong>g> modificati<strong>on</strong><str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> DNA sequence, and have been shown to result fromvarious, n<strong>on</strong>-exclusive processes including DNA or hist<strong>on</strong>emethylati<strong>on</strong>, hist<strong>on</strong>e acetylati<strong>on</strong>, and chromatin compacti<strong>on</strong>-18-
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