Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. ity to hybridize and form viable F viable form and to hybridize ity demonstrated that cultivated fl fl between studies fl cultivated with insympatry grow may and world ofthe parts inmany distributed are which cies genus ofnovel fl risk environmental of to evaluation reviewed the was initiate tion interspecifi reported and relatives offl phylogeny the occurrence, the tives, fl transgenic from introgression gene for potential The populations. their on or weedy relatives and the it impact may have fl gene for potential the is novel trait ofany release the for consideration important An products. nutraceutical and bio-industrial of production the for platform acrop as ated ploidy level, and sexual compatibility. sexual level, and ploidy distribution, sympatry, fl concurrent species on depending America, native to North species inseveral possible is relatives weedy or or weedy relatives, gene fl the evidence hybridization of reported with wild However, on reported. or based studied been not either has relatives wild other many with CROP ( Flax pallescens, fl L. bens, L. angustifolium, L. corymbiferum, L. decum- of species nine least at ax. Interspecifi ax.

SCIENCE Linum usitatissimum Linum Linum occosum, L. hirsutum, L nervosum, L. L. Lnervosum, hirsutum, L. occosum, Weedy and Wild Relatives: An AvenueWeedy Relatives: An Wild and for and , contains approximately 230 spe- 230 approximately contains VOL c hybridization and cytogenetic cytogenetic and c hybridization L. tenue Potential Hybridization of Flax with with of Flax Hybridization Potential ABSTRACT . species congeneric and ax 48 Movement Genes? of Engineered , MAY ). Hybridization offl ). Hybridization – Linum fl from ow L.) is being evalu- being L.) is JUNE ax in Canada. The The inCanada. ax abil- the has ax ax to wild rela-

( 2008 Amit J. Jhala, Linda M. Hall,* and Jocelyn C. Hall C. Jocelyn and Hall,* M. Linda J. Jhala, Amit L. africanum, 1 c hybridiza- c plants with wild to ow 825 ax to wild owering, owering, axwild ax Sept. 2007. *Corresponding author ([email protected]). author 2007. *Corresponding Sept. 6 Received Canada. T6G 2E9 AB, Edmonton, Univ.ter, ofAlberta, Cen- Sciences Biological Sciences, Dep. ofBiological Hall, C. Jocelyn Canada. T6G 2P5 AB, Edmonton, Univ. ofAlberta, Building, estry 410 Food, and Agriculture/For- Agriculture Alberta Hall, M. Linda Canada. T6G 2P5 AB, Edmonton, Univ. ofAlberta, Science, tional Nutri- and Food Dep. ofAgricultural, Hall, M. Linda and J.Jhala Amit has been obtained by the publisher. the by herein obtained been contained has material the reprinting for and printing for from Permission writing in publisher. the permission without system, retrieval and storage information any or recording, photocopying, including mechanical, or any in electronic means, any by or form transmitted or reproduced be may periodical this of part No reserved. rights All 53711 WI USA Madison, Rd., Segoe S. 677 America of Society Science © Crop doi: 10.2135/cropsci2007.09.0497 (2008). 48:825–840 Sci. Crop in Published F DNA; amplifi random RAPD, ofCanada; Resources Gene PGRC, Plant Station; Introduction Plant Regional Central North Inspection Food modifi genetically Canadian GM, CFIA, Organization; Biology Group; Molecular European EMBO, Agency; Phylogeny Angiosperm APG, Abbreviations: Zealand, and Australia (Rosberg, 1996). (Rosberg, Australia and Zealand, fl transported colonists New European America, to ax North era, Colonial the 1904). During Candolle, (De Finland and sia 1200 B.C. Flax(Ros- cultivation East to 2500 Near from period the the 1997) 1996; during berg, Stephens, from by AryansEurope flinto ax extended transporting north for sible to Rus- Matthews, 1908). 1904; Candolle, (De years at>4500 were dated mummy-cloths It is believed Egyptian from fabrics Flax Pharaohs. ofthe time the during use that Phoenicians (Cooke, 1903) in where Egypt it was from disseminated edly were respon- not beenidentifi man, 1938; Richharia,etable fi 1962). fi The forlinen 1990). Rowland, grown Itand is center of origin of fl ax has lax ( in the world and is one of the oldest cultivated plants (Bhatty (Bhatty plants cultivated oldest oneofthe is world and the in sad2 ber domesticated by mankind, and as an oilseed (Dill- oilseed an as and bymankind, domesticated ber Linum usitatissimum , stearoyl-ACP desaturase II. desaturase , stearoyl-ACP AFLP, amplifi ed; ITS, internal transcribed spacer region; NCRPIS, NCRPIS, region; spacer transcribed internal ITS, ed; ed (Lay and Dybing, 1989), report- butit was Dybing, and (Lay ed L.) is the sixth largest oilseed crop crop oilseed largest sixth L.) the is ed fragment length polymorphism; REVIEW &INTERPRETATION cation of polymorphic ber, the earliest veg- ber, the earliest Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. of Canada, 2007a). of Canada, Council (Flax Korea South and U.S., the Japan, Europe, to mainly production, total ofthe 90% to 80 exported and offl2006) tonnes Canada, million (Statistics ax seed 1.041 produced Canada 1994. 2006, held since In it has offl export and aposition production the seed, ax in leader world flthe world’s currently is Canada output. ax seed total ofthe for64% responsible are together States United the and China, Canada, 2005). Jimmerson, and (Smith tonnes metric 1.903 was million production seed and tries, 47 in coun- grown was linseed 2004, In 2006). (Vromans, linen quality of high production forthe coast European northwestern the near and Egypt, Russia, China, in ily Today, times. fi ancient since France northern and Belgium in probably and erlands 1994). flMiller, Fiber the Neth- in cultivated been has ax and (Hammond westward settlers followed production as Risk assessment of transgenic fl of transgenic assessment Risk transgene including ax 1998). 1988; Alstad, and Andow (Ellstrand, evaluated be must resources genetic and 2003) etal., (Wilkinson sity onbiodiver- impact potential the addition, 2003). In al., 1993; et Conner (Dale, spread fortransgene potential the wild gene fl ofcrop-to- understanding Abetter 2004a). 1993; CFIA, Gray, and (Raybould populations wild to GMcrops from movement oftransgenes potential the of anovel is crop before release the beaddressed must that concerns critical ofthe 2003). One Napetal., 2003; (James, increase to continues production under area the and ofspecies ber 2005). etal., Millam also see 2004; Breithaupt, 2002; (Hricova, crops transgenic and bioproducts open to more being toward moving EUis the then, Since uct. fl and paint in use even crop, development forprimary of the genic modifi ofgenetically import the to 1998),et al., over market’s reaction the aconcern (Polyakov resistance 1989) (McHughen, ordisease control signifi to a solution fl transgenic havemay been 2007b). Although ax Canada, of Council fl ofthe request at the (Flax diately industry ax imme- almost deregistered butit 2004b), was (CFIA, dues for unconfi (McHughen et al., 1997), was released in Canada in 1998 fl1988). onetransgenic Only CDC Triffi cultivar, ax McHughen, and (Jordan resistance 1995), glyphosate and Holm, and (McHughen resistance fosinate-ammonium sulfuron methyl resistance (McSheff fland met- in expressed been chlorsulfuron including ax have novel traits Several 2002). (McHughen, value nomic agro- ofpotential genes with transformed and formation with engineered genetically 826 GM crops are now grown worldwide, and the num- the and worldwide, now grown are GM crops fi the among was Flax years many for flFiber America in North ax prospered ooring industries and animal feed as a co-prod- a as feed animal and industries ooring ned use in fi in use ned ow is essential for ecological risk assessment of of assessment risk ecological for essential is ow such weed as issues agronomic cant elds with persistent herbicide resi- herbicide persistent with elds both be to species crop rst ed material halted all trans- all halted ed material Agrobacterium flber primar- grown is ax rey et al., 1992), glu- mediated trans- mediated WWW d d . CROPS of approximately 70 species indicates that blue-fl that indicates 70 species of approximately analysis Their (ITS). region spacer transcribed internal encoded nuclear the and markers chloroplast multiple of study phylogenetic comprehensive amore conducted (2005) son fi of evidence is there 2003), although Richards, sen and 1931;Winkler, 1968; Walters, Ockendon Diederich- and (e.g., species ofthe sections traditional with compatible classifi researchers Many knowledge. increased (AFLP) based phylogeny of17 phylogeny of based species (AFLP) limited. An amplifi paper. this in followed be will evidence, interspecifi and cytological, morphological, ing of 41 classifi prevailing 1987). 1983; However, Joshi, nosingle Gill, and is there 1966; 1962; Ockendon, 1971; Gill, Lewis, Chennaveeraiah 1941; 1929; Nagao, Ray, and (Kikuchi, 1944; Osborne research- other Alternatively, grouped ers 1962). Dill- 1933; Richharia, 1953; Dillman, man, 1931; Winkler, 1926; Vavilov, 1925; 1857; 1904; Tammes, DeCandolle, (Linnaeus, oforigin ter orcen- characters ofmorphological basis onthe either cies taxonomy and classifi vardtia Linum usitatissimum (1968). Walters byOckendon fl and added Cultivated linum tions, and species) (eight progress (Hall et al., 2006). etal., (Hall progress fl in is relatives transgenic movement weedy from to its ax Flax is a TAXONOMY PHYLOGENY AND of 22 genera (Vromans, 2006) and and 2006) (Vromans, genera of 22 ily includes: includes: ily fam- the in genera 2003). Important II, (APG Malpighiales order the in placed been has family the recently but most 1981), (Cronquist, taxonomists bysome Linales order the cies (Hickey, 1988; Heywood, 1993). Linaceae is placed in wild or weedy species. orweedy wild fl transgenic between and its introgression ax and tion ofinterspecifi probability (iii) and ability, fl to cross- species and related closely relatedness (ii) ax, fl and on (i)biology, distribution, based testing, fl transgenic experimental from before ax fl ofgene risk potential the establish to is objective Our pairing. chromosome and genome, level, ploidy lar asimi- having species related closely with hybridize to ve species clusters (Vromans, 2006). McDill and Simp- and McDill 2006). (Vromans, clusters ve species . ORG Phylogenetic studies based on molecular markers are are markers onmolecular based studies Phylogenetic The genus genus The We hypothesize that (Winkler, 1931) with an additional section, section, 1931) additional an (Winkler, with

(two–four species), species), (two–four Linum, Linastrum, Cathartolinum, Dasylinum, Dasylinum, Cathartolinum, Linastrum, Linum, member Linum Linum Linum Linum Linum based on DNA sequence variation from from onDNA variation sequence based species proposed by Gill (1987), on byGill based proposed species cation scheme for this genus. The group- The genus. forthis scheme cation of the family Linaceae which is composed composed is which Linaceae family the of (230 species), species), (230 Radiola species based on chromosome number number onchromosome based species , is placed in the section section the in placed , is ed fragment length polymorphism is traditionally divided into fi into divided traditionally is cto of cation CROP Anisadenia (Heywood, 1993). (Heywood,

L. usitatissimum SCIENCE Hugonia Linum approximately owering phenology of phenology owering , (two species), species), (two VOL has changed with with changed has . (40 species), 48 , c compatibility compatibility c is more likely MAY ed c hybridiza- c Linum Linum – Linum JUNE 300 spe- 300 Roucheria Cliococca, owered owered and ve sec- ve is not . The . The

Rein- 2008 spe- Syl- ow ax,

Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. for the introduction of favorable traits to fi to traits offavorable introduction for the and a wild relative relative awild and fi et al., 2002). (Fu cultivars oil than base Vromans’ genetic homogenous more and (2006) fl oil AFLPand fi that indicated ax study supports fi comparing study Amolecular 2006). Fu, and ichsen this diff notshow marked did characters tive qualita- two and however, RAPD traits; quantitative in fl ofcultivated groups observed four the was cultivars ax than cultivars. Considerable diff landraces in existed variation more comparatively Overall, distinct. most were the Africa and subcontinent Indian the from accessions whereas diverse, were most regions European and Asian East the from accessions out that geographic patterns of on 1986). fl astudy In (Durrant, species inbreeding ax obligate variability, Fu (2005) pointed This fi 2000). etal., (Mansby believed earlier than outbreeding gosity found in in found gosity heterozy- large the that conclusion the to led accessions within diversity genetic high unexpectedly An groups. defi and fl in diversity genetic the ax study to markers isozyme used of variation species 1941; 1987).(Nagao, Gill, ofnine number karyotype haploid same the have they of a member specifi complicated. Neither Neither complicated. classifi cate that indi- (2006) ofVromans study 1968), molecular butthe from other 2002). ber (e.g., num- chromosome same the share that species other with with clustered is (2n =30) analysis based on RAPD data indicate that among relationships Sclerolinon digynum, classifi previously bers Linum VARIABILITY INCHROMOSOME NUMBERS CROP diff and ognized rec- be to species several allowed has which ago, century Karyotypic analysis of fl America. The subsequent diversifi to have appear North colonized independently lineages yellow-fl blue and the ofboth members and asia ered lineage. These diversifi lineages initially owered owered nding and he further speculated that linseed cultivars cultivars linseed that speculated he further and nding Additional molecular studies have focused onwithin- focused have studies molecular Additional Karyotype number is notrefl is number Karyotype

SCIENCE c group, even though though even group, c nding was unexpected as fl ax is reported to be an species were sister to a predominantly yellow-fl apredominantly to were sister species Linum perenne ned ned fi Linum L. angustifolium , Linum ve groups, but with low variation within the the within low variation butwith ve groups, VOL L. perenne cation among the the among cation species in North America includes mem- includes America North in species erentiated (Tutin et al., 1968). The genus L. usitatissimum . 48 species morphologically (Ockendon, morphologically species and and L. usitatissimum. L. bienne , group can be easily distinguishable distinguishable easily be can group MAY ed as separate genera: Linum L. perenne Linum Cliococca selaginoides group (Diederichsen, 2007) and and 2007) (Diederichsen, group – and JUNE ber cultivars have a narrower have anarrower cultivars ber L. grandifl orum species began more than a half ahalf than more began species could be important sources sources important be could species. For example, an L. austriacum

L. usitatissimum may be the result of more ofmore result the be may erence within and among among and within erence 2008 nor of phylogeneticective L. perenne yellow ofthe cation

Mansby et al. (2000) (2000) etal. Mansby L. austriacum erences (Dieder- erences ber fl ber ax. (2n =16), not . is considered group is still L. decumbens Hesperolinon Eur- ed in ; Fu et al., etal., ; Fu owered owered form a form WWW ow- ber . , CROPS

some number sifi Linum ofother numbers chromosome the regarding researchers 1927). (Martzenitzin, mitosis somatic the in segmentation 1966), accidental an and Gill, 1944; (Ray, stain some retain to fragments observed of the in chromosomes ofthe size small the confl 1983). forthe Joshi, reasons The were results icting and 1966; 1962; Chennaveeraiah Gill, 1944; Richharia, 1938; 1941; Nagao, Ray, 1929; Dillman, (Kikuchi, = 30 tion studies confi chromosome the interspecifi and However, cytogenetic later estimated flax 1927; Lutkov, 1939). =32 2n (Martzenitzin, be to number cultivated of number some chromo- ofthe studies 1983). Joshi, and Initial naveeraiah (Chen- aneuploidy and polyploidy dueto be may numbers chromosome in 1955; 1987). Wylie, and Diversity Gill, n a remarkable diversity in chromosome number including Linum derived from two or more ancestral forms (De Candolle, Candolle, (De forms ancestral ormore two from derived (Gill, 1987). 1904; 1982). Zeven, Candolle, (De area the in found offl form adiverse Many because oforigin center another as suggested been authors has India toward Mediterranean of the east area an 1904). Finally, Candolle, (De of dissemination reported acenter be could Egypt that believe tively, researchers other offl centers probable four these discussed cultivated (1962) have also (1987) Richharia and Center. Gill sinian Abys- the and Mediterranean, the Eastern, Near the atic, flwere reported toax have originated inis four,important the cultivated Central plants (Vavilov, Asi- 1926), world’s ofthe most oforigin centers independent eight the Among theories. existing many 1989) with Dybing, and The centerEVOLUTION OF of origin AND OFORIGIN CENTER of cultivated fl or 36 (Gill, 1966).(1929) (1941) Nagao and observed bonense results. the in variability forthe account could 1939), Chopinet, and (Simonet which tetraploid a Japanese be may (1929) chromosomes counted Kikuchi which from (Table 1). species The ofthis ber num- chromosome the in uncertainty (1966) indicated (1941) as species this have grouped (Seetaram, 1972), and (Seetaram, of count 1929). with qualifi = 8, 9, 10, 12, 14, 15, 16, 18, 30, and > 30 (Darlington 9, =8, 10, 14, 12, 15, 16, (Darlington >30 18, and 30, . ORG ed There were some disagreements among various various among disagreements were some There The progenitor of cultivated fl ax is also uncertain 827 L. alpinum species (Table 1). For example, Kikuchi (1929) (Table clas- 1). species Kikuchi Forexample, exhibit that species ofdiploid number alarge has Linum hirsutum Linum was grouped as as grouped was n = 8 (Ray, 1944), =8(Ray, cation, as as cation, n = 18, whereas, Ray (1944) Ray =18, Nagao whereas, and as a member of group III with chromo- rm the chromosome number to be 2n Linum monogynum Linum has a variable reported chromosome chromosome reported avariable has L. USITATISSIMUM L. n =16 1966; or18 Table (Gill, 1). n n = 43 and 2n = 86 (Kikuchi, (Kikuchi, =86 2n =43 and = 14 (Ray, 1944), but Kikuchi =14 1944), (Ray, butKikuchi n = 9 (Nagao, 1941), =9(Nagao, Linum alpinum Linum n =9, =18 2n and and/ n ax is uncertain (Lay Linum =9(Table 1). Gill has been reported, beenreported, has ax origin. Alterna- , the tendency tendency , the Linum c hybridiza- c Linum nar- specimen specimen species species n =15 ax is ax Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. 828 species for which cytological information is available. is information cytological which for species of groupings various of Table 1. Comparison L. usitatissimum L. fl avum L. corymbiferum L. angustifolium L. sibiricum L. americanum ( II. Group L. perenne L. narbonense L. lewisii L. hologynum L. altaicum L. muilleri L. extraaxillare L. austriacum I( Group L. monogynum ( IV Group L. alpinum ( III Group Kikuchi (1929) Ray (1944) Nagao (1941) Gill (1987) Gill (1941) Nagao (1944) Ray (1929) Kikuchi Pursh n L. Moris. n n =9) n Jacq. L. Fisch. = 18) Group III ( III Group =18) = 15) Group II ( II Group =15) = 43) Group IV ( IV Group =43) DC L. Reichb. Kit. L. L. Forst. Huds. L. Desf. L. hologynum L. collinum L. austriacum L. altaicum L. alpinum L. grandifl orum I( Group L. loreyi L. lewisii L. hirsutum L. medium L. capitatum L. gallicum L. perenne L. usitatissimum L. narbonense L. tommasinii L. strictum L. fl avum L. angustifolium V( Group Jord. Pursh. n L. n n n =8) n Jacq. = 9) Group II ( II Group =9) L. Britton. L. = 15) Group V( Group =15) L. Guss. Fisch. = 10) Group III ( III Group =10) L. = 14) Group IV ( IV Group =14) Kit. Nym. L. Reichenb. L. Desf. Huds. L. Linum WWW species based on chromosome numbers, including only those those only including numbers, chromosome on based species L. usitatissimum L. crepitans L. angustifolium L. grandifl orum I( Group L. tenuifolium L. sibiricum L. perenne L. narbonense L. muilleri L. montanum L. lewisii L. hologynum L. austriacum L. hirsutum L. fl avum L. campanulatum L. extraaxillare L. maritimum L. altaicum L. alpinum . CROPS Pursh. n L. n Moris. n n =8) n Jacq. ( Jacq. = 15,16) Group II (2n II =30) Group =15,16) L. = 9) Group VOthers Group =9) Ldb. . = 14) Group III (2n III =28) Group =14) L. = 9) Group IV (2n IV =16) Group =9) DC. Dum. ORG L. Schleich L. L. Reichb. Kit. L. Desf. Huds. L.

L. n =18) L. corymbiferum L. bienne L. angustifolium L. album L. africanum L. strictum L. perenne L. narbonense L. lewisii L. julicum L. hologynum L. austriacum L. decumbens L. tenuifolium L. grandifl orum L. anglicum L. alpinum I(2n =18)Group L. schiedeanum L. marginale L. altaicum L. dolomiticum L. capitatum L. campanulatum L. usitatissimum L. tenue L. sulcatum L. fl avum L. rupestra L. monogynum L. maritimum L. catharticum L. rigidum L. hispanicum L. viscorum L. hirsutum L. rigidum L. pallescens L. nervosum texanum L. medium L. humile L. gallicum CROP Desf. Pursh.

Kotschy L.(2n =28, 30) Mill Mill. (2n =30,32) Mill. Hayek. Pursh. var. Pursh. Pursh. var. Pursh.

Jacq. (2n =18,36) Jacq. L. Planch. (2n = 30,36) var. (2n =30,36) Planch. L. L.(2n =20) Engelm. (2n =36) Engelm. Ldb. SCIENCE L.(2n =16,18) Mill. L. Riddell. A. Cunn. (2n =80) Cunn. A. L.(2n =30,32) Waldst. L.(2n =24,28) L.(2n =20) Ldb. L. L. Reichb. Mill. Desf. L.(2n =18,28) L.(2n =16,57) Borb. Forst. (2n =86) Forst. Desf. (2n =16,18) Desf. Huds. (2n =30,32) Huds. S.&C. (2n =36) L. Desf. (2n =18,30) Desf. L. , VOL rigidum fi lifolium . 48 Rog. Rog. , MAY – JUNE

2008 Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. (Ockendon, 1968). Hand pollinations in (Ockendon, in 1968). pollinations Hand not forfi angustifolium locus in cultivated ( II desaturase stearoyl-ACP ofthe diversity genetic fl the that ax estimated it was study is1997). amolecular In low compared Jaworski, and (Ohlrogge acids fatty ofunsaturated lation to pale fl formanipu- used been has ax ( thus, ACP oleoyl-ACP to and, 2003). etal., (Muravenko cies ofsubspecies and genetically related although some consider to species sister the is al., 2002). A diff et (Fu group same the in clustered consistently species usitatissimum seven of 1967b). analysis Yermanos, 1967a; and ARAPD Gill 1929; Yermanos, Ray, and 1944; Gill (Kikuchi, studies cytological and 2006) Fu, and 1936; Diederichsen man, This hypothesis is supported by morphological (Dill- CROP fl of yellow species “thrum”. Several called is reverse “pin” the as and have fl fl contrasting (tristyly) or three 1941). (distyly) (Rogach, have two species Heterostylous selecting 1967a). Yermanos, and between fl gene potential predict to information provide and mum ofinterspecifi studies The 1966; 1982). 1962; Zeven, Gill, 1904; Richharia, dolle, offlCan- form (De where adiverse found been has Asia ax of of species related closely ofseveral 1967).Knowles, Hybridization and (Briggs parent either in not found of newcharacters expression forthe responsible also is 1997) and (Arnold, plants crop of many evolution the in arole played has relatives wild and species crop between Hybridization Interspecifi and ( species three ofthese markers 2000). However, Hopf, and 1968; (Tutin etal., Zohary species same the be genome comparisons with molecular species wild asingle from fl cultivated that tively, suggested it was ax originated from those diff be to were believed indigenous Egyptians byancient vated to1904; 1962). culti- species Vavilov, The 1926; Richharia, Russia and Siberia. Alterna- L. usitatissimum The Heterostyly must be taken into consideration when consideration into taken be must Heterostyly with its wild relatives enable estimates of crossability ofcrossability estimates enable relatives wild its with

L. usitatissimum SCIENCE owers with long styles and short stamens are known known are stamens short and styles long owers with Linum ber (Allaby et al., 2005). etal., (Allaby ber sad Linum Linum Linum owered owered ) suggesting fl) suggesting fi ax was gene is responsible for converting stearoyl- forconverting responsible is gene , have a high RAPD similarity and these two two these and similarity RAPD have ahigh species revealed that that revealed species c Hybridization in c Hybridization VOL L. usitatissimum, species (Kikuchi, 1929; Gill, 1966; Gill 1966; Gill 1929; Gill, (Kikuchi, species erent AFLP study indicates that that indicates study AFLP erent might have played a role in the evolution evolution the in arole played have might species forinterspecifi species . 48 in the Mediterranean and Southeast Southeast and Mediterranean the in Linum ) confi , MAY L. usitatissimum L. bienne L. angustifolium – were found to be heterostylous c hybridization of c hybridization JUNE thatrm they are very closely L. angustifolium ower types. The plants that that plants The types. ower rather than a separate spe-

2008 L. angustifolium can be considered as a rst domesticated for oil, foroil, domesticated rst L. angustifolium

Linum (Vromans, 2006), and L. grandifl orum c hybridization hybridization c (Heer, 1872). L. bienne L. usitatissi- , L. bienne, L. bienne and erent WWW sad2 ow ow to of L. L. . ) CROPS

when crossed with with when crossed crossed with with crossed 1). karyotype with relatives wild nine at least with fl hybridize to cultivated the potential that has cate ax 1972; 1). 1967a; Fig. Yermanos, and Seetaram, (Gill tively crepetans with low F with hirsutum, L. decumbens, of Hybridization chromosomes. nonhomologous two involving each translocations, two duced fertile F fertile duced L. angustifolium with hybridized it successfully parent, a male All these crosses produced fertile F fertile produced crosses these All 1972). 1970; Seetaram, Godward, 1967a; and manos, Bari Yer- 1929; and 1966; Ray, Gill 1928; 1944; Kikuchi, Gill, Seetaram, 1972; 1). Fig. Seetaram, successful (Kostopoulos, 1970). fl orthrum ofpin pollination but self (85–97%), fertile were highly ×pin orthrum ×thrum pin In a cytogenetic study Gill (1966) reported that (1966) that reported Gill study acytogenetic In cies ( angustifolium Linum bens species, related simum vated fl ax are also successful. When flvated successful. also are ax 1972; 1). Fig. 1970; Seetaram, Godward, biferum, L. fl L. biferum, pallescens, L. occosum, usitatissimum between hybridization ofsuccessful reports many with but later between by Kolreuter fi The Linum direction, presumably due to their similarity in ploidy lev- ploidy 1972). fi among in Crosses Seetaram, similarity 1969; Godward, and their (Bari to ofchromosomes size and els due presumably direction, tum, species: tile F fer- have produced species three 1). (Fig. All relatives wild other with introgression and byhybridization butalso ing, backcross- through notonly ecosystem, fl natural the in ax novel from transgenes retain and occur species three these between hybrids if determine to conducted be should ies F simum decumbens, L. corymbiferum, L. angustifolium, . 1 ORG progeny exhibiting 80 to 90% germination (Gill, 1966). (Gill, germination 90% to 80 exhibiting progeny Linum africanum, L. angustifolium, L. africanum, Linum , but did notdiff , butdid In summary, interspecifi Hybridization events among species other than culti- than other species among events Hybridization and L. africanum, L. corymbiferum, 1 rst interspecifi 829 L. angustifolium diff were highly successful in at least one direction with with onedirection atleast in successful were highly seeds in crosses with at least two of the following following ofthe two atleast with crosses in seeds Hybrids among Taxa among with Hybrids and L. decumbens, L. fl occosum, L. hirsutum, L. stric- L. narbonense L. tenue closely three from byonetranslocation ers 1 fertility (Sharma and Khanna, 1964; Bari and and 1964; Bari Khanna, and (Sharma fertility and L. usitatissimum L. humile , and , and 1 L. africanum plants (Fig. 1). Therefore, further stud- 1). (Fig. further Therefore, plants (Sharma and Khanna, 1964; Gill, 1966; 1964; Gill, Khanna, and (Sharma L. africanum, L. angustifolium, L. corym- diff er from c hybridization in in c hybridization L. hirsutum (Tammes, 1928). There have been L. fl occosum pollen have produced fertile plants plants have produced pollen fertile was considered to be synonymous ers from the other three wild spe- wild three other the from ers and L. usitatissimum , L. nervosum and all reciprocal crosses pro- crosses reciprocal all and L. corymbiferum L. angustifolium, c hybridization studies indi- studies c hybridization and (Seetaram, 1972). (Seetaram, ve taxa, ve taxa, L. usitatissimum and 1 and hybrids in at least one atleast in hybrids owers were only 3.0% 3.0% wereowers only and L. hispanicum, L. strictum Linum and L. tenue were but reported L. pallescens L. decumbens n L. africanum, L. and in this respect. respect. this in =15 and L. narbonense, was reported reported was n L. africanum was used as as used was = 15 (Fig. =15 (Fig. (Tammes, L. usitatis- L. usitatis- L. decum- with respec- Linum were were ) by L. L. ,

Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. cross (male to female). These are the related species with the greatest potential to hybridize with fl with to hybridize potential greatest the with species ax. related the are to These female). (male cross bonense that ofthese ofchromosomes pairing The L. austriacum, L. julicum oftaxa species between crosses genus the in group largest the tute taxa other than among hybridization ofsuccessful studies have been There ( Taxa than Other Linum 1.Figure Artifi interspecificial among c crosses 830 The diploid diploid The 1929; 2). Fig. (Laibach, culture by embryo butonly duced perenne 1966). When (Gill, chromosomes nonhomologous three discovered between was the of genomes However, occur. to ing adiff the that indicating were suffi species six bivalents, normal formed diff apparently also austriacum that speculated 1967b). Yermanos, and (Gill further They alpinum, L. austriacum, L. julicum, L. narbonense, duced fertile F fertile duced autotetraploid autotetraploid were observed at metaphase I (Nagao, 1941). I(Nagao, at metaphase were observed univalents and bivalents, trivalents, and studied was cross The chromosomes not involved in translocations translocations notinvolved in chromosomes The L. altaicum and and was crossed with with crossed was and Hybrids amongHybrids L. perenne L. perenne 1 L. narbonense, diff plants (Gill, 1966; Gill and Yermanos, 1967b). L. alpinum n = 15 (Fig. 2). The taxa with with 2). =15 taxa (Fig. The ers by one reciprocal translocation from from translocation by one reciprocal ers er by one translocation, whereas whereas er by one translocation, pair- fornormal homologous ciently diff , was successfully hybridized with n er by two translocations. er bytwo L. alpinum L. narbonense, (Kikuchi, 1929). Meiosis in this 1929). this in Meiosis (Kikuchi, =15) L. austriacum and erence of two translocations translocations erence of two n =9, L. julicum Linum and L. alpinum, L. altaicum, n and = 9 species revealed revealed =9species , hybrids were pro- were , hybrids L. perenne (Gill, 1966). Some Linum and L. perenne and n L. narbonense =9, consti- species ( species involving L. perenne L. nar- , pro- WWW n L. L. L. = 15) that resulted in fertile progeny. Arrows indicate the direction of the ofthe direction the indicate progeny. Arrows fertile =15) in resulted that . CROPS

of gene fl of gene diff with species that suggest studies greenhouse These plants. fertile produced has cross notasingle number, mosome diff flvated a having species with crossed is ax 1972; 2). culti- Fig. When 1970; Seetaram, Godward, and into nine areas: Africa, Antarctic, Asia-Temperate, Asia- Antarctic, Africa, areas: nine into world terrestrial (1992), the divides which Brummitt and publication of Hollis the standard accordance with in Linum GEOGRAPHIC DISTRIBUTION F fertile producing in successful was numbers chromosome 1970; 2). Fig. Godward, and 1966; Bari role in interspecifi important an plays karyotype suggest 2). results Fig. These 1972; 1970; Seetaram, Godward, and 1966; Bari 1964; Gill, F fertile produce to orfailed seeds, any notproduce did either with species other between vulgaricum between diff tum, L. strictum, L. usitatissimum . 1 1 ORG plants (Kikuchi, 1929; Ray, 1944; Sharma and Khanna, Khanna, and 1929; Ray, 1944; Sharma (Kikuchi, plants plants (Rogach, 1941; Richharia, 1962; Gill, 1966; Bari 1966; 1962; Bari 1941; Gill, (Rogach, Richharia, plants erent chromosome numbers was also studied. Crosses Crosses studied. also was numbers chromosome erent erent chromosome numbers have no or minimal risk risk have noorminimal numbers chromosome erent Thus, only hybridization between species with equal equal with species between hybridization only Thus, Interspecifi

species’ distribution records are grouped into regions regions into grouped are records distribution species’ ow them. to L. alpinum ( n =9), and c hybridization between between c hybridization c hybridization in in c hybridization ( n CROP =9,18), L. usitatissimum

n SCIENCE =15 (i.e., ) and ) and L. austriacum , VOL L. grandifl orum Linum L. crepetansL. . ; as well as crosses Linum 48 , MAY species (Gill, (Gill, species ( species with with species erent chro- erent – n =9), JUNE , L. hirsu- ( n =8),

2008 L. Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. tum, L. capitatum tum, L. extending up to Asia including Mediterranean the in found species perennial other many 1928). are (Tammes, There UK southern the and Ireland, area, subMediterranean and Mediterranean ofthe species lium oilseed. an as grown where it is Romania forfi ily primar- 1987). grown it is Europe, (Gill, In climates ate (male to female). Solid lines indicate fertile F fertile indicate lines to Solid female). (male CROP of41 distribution geographic the Here we discuss America. North in except were not itemized distributions for orprovincial State 2006). (USDA-NRCS, taxa widespread species ofthe presence the within records specimen) herbarium countries, (i.e., evidence orother citation aliterature that indicates but region, that in occurrence widespread imply essarily notnec- does region orpolitical ageographical in a taxon for report However, adistributional America. South and Pacifi America, North Europe, region), Australia, single a as Asia considered we have manuscript, this (in Tropical 2. InterspecifiFigure in c hybridization occurred, but F but occurred, and and so these species are frequently cultivated in European European in cultivated frequently are species these so and narbonense, tum, L. such as Huds, a putative wild progenitor offl progenitor aperennial is wild ax, aputative Huds, Flax is cultivated in almost all continents with temper- with continents all almost in cultivated is Flax L

SCIENCE . viscorum L. austriacum, L. fl avum, L. grandifl orum, L. hirsu- ber, except in Germany, Hungary, Poland, and , 1 VOL hybrids were not obtained with embryo rescue and/or treatments with colchicine. with treatments and/or rescue embryo with obtained not were hybrids (Tutin et al., 1980; (Tutin Table etal., species 2). Many , . L Linum 48 and . , dolomiticum, L. hologynum, L. julicum, L. hologynum, L. dolomiticum, MAY L. perenne species (Table 2). species – JUNE

2008 L. alpinum, L Linum have attractive fl attractive have

(species with different chromosome numbers). Arrows indicate the direction of the cross cross ofthe direction the indicate Arrows numbers). chromosome different with (species 1 hybrids were obtained with viable seed production. Dotted lines indicate hybridization Linum angustifo- . campanula- owers owers WWW c, . CROPS

Flax is grown primarily for seed in the Canadian prairies, prairies, Canadian the in forseed primarily grown is Flax 1907; 1987; 2007; Table Budd, (Small, 3). Diederichsen, Mexico and U.S., the Canada, New World throughout 63 than more are There Linum tifolium, L. fl L. tifolium, pallescense L. nervosum, L. avum, ria, 1962). Many species including BC; Richha- 2000–1500 (around period Dravidian the during India in cultivation in have been might species perenne Linum 1962). (Richharia, plants ornamental as India in duced strictum (1875) reported two additionalrecorded species, in many states1982). Two species, of India (Cooke, 1903). Hooker plants. ornamental as nurseries in available and gardens botanical Canadian and graphical ranges of individual species is given in Table in given 2. is species ofindividual ranges graphical ongeo- 1990; Table information 3).Hnatiuk, Detailed 1972; (Willis, NewZealand and Australia in distributed 1984;al., Table 2). (Greuter et region that within extensively distributed and and . ORG There are many many are There L. tenuifolium 831 . Species in the New World the in Species Linum angustifolium Linum was reported in escaped clusters and this this and clusters escaped in reported was are native to the Russian Federation Federation Russian the to native are Linum marginale Linum L. mysorense Linum Linum and species native to Asia (Zeven, L. grandifl orum species distributed in the the in distributed species and and L. altaicum, L. angus- L. usitatissimum L. monogynum L. perenne , were intro- were L. perenne, and were were are L.

Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. Group II (2n II =30) Group I(2n =18)Group of distribution Table Geographical 2. 832 humile L. humile L. hispanicum L. fl avum L. decumbens L. corymbiferum (2n 32) =30, Mill) L. angustifolium L. album L. africanum L. tenuifolium L. strictum L. perenne L. narbonense L. lewisii alpinum L. julicum L. hologynum L. grandifl orum L. austriacum anglicum L. anglicum L. altaicum (2n =18, 36) alpinum L. alpinum Mill Pers.) (Jacq.). &Stef.) Stoj (Jacq.) Stoj.& Stef.) Pursh North America: Canada: AB, BC, MB, ON, QC, SK, YT Hitchcock et al. (1969); al. et Cody, (1996) Hitchcock YT QC, SK, ON, BC, MB, AB, Canada: America: North Pursh Kotschy Asia: Iran, Syria Guest et al. (1966) al. et Guest Syria Iran, Asia: Kotschy L. (2n = 28, 30) Asia: Russian Federation, Turkey Davis et al. (1988); al. (1992) et Davis Huxley Turkey Federation, Russian Asia: L.(2n =28, 30) Mill ( (Mill.) Ockendon Hayek ( Hayek Jacq. ( Jacq. L. Africa: Algeria, Egypt, Ethiopia, Libya, Morocco, Tunisia Rechinger (1963); Tutin (1980); al. et Rechinger Tunisia Morocco, Libya, Ethiopia, Egypt, Algeria, Africa: L. L. Asia: India, Russian Federation Richharia (1962); (1969); Komarov Richharia Tutin Federation Russian India, Asia: L. Ldb. Asia: Kazakhstan, Mongolia, Russian Federation Czerepanov (1995) Czerepanov Federation Russian Mongolia, Kazakhstan, Asia: Ldb. Mill. ( Mill. L. (2n = 30, 32) Africa: Cape Province Bond and Goldblatt (1984); Arnold and (1984); and Arnold Goldblatt and Bond Province Cape Africa: L.(2n 32) =30, L. Asia: Armenia, Azerbaijan, Georgia, Iran, Russian Federation, Syria, Syria, Federation, Russian Iran, Georgia, Azerbaijan, Armenia, Asia: L. L fia lei,MrcoTutin (1984) (1980); al. et al. et Greuter Morocco Algeria, Africa: L. Rih.Erp:Abna ugra ree oai,Ygsai Tutin (1984) al. et (1980); al. et Greuter Yugoslavia Romania, Greece, Bulgaria, Albania, Europe: Reichb. Mill. Europe: Albania, Bulgaria, France, Greece, Ireland, Italy, Portugal, Portugal, Italy, Ireland, Greece, France, Bulgaria, Albania, Europe: Mill. L. usitatissimum Taxon Desf. Europe: Germany PGRC, personal communication (2006) communication personal PGRC, Germany Europe: Desf. L 2 8 8 fia lei,MrcoTutin (1984) al. (1980); et al. et Greuter Morocco Algeria, Africa: L.(2n =18, 28) Desf. Africa: Algeria Greuter et al. (1984); al. et (1992) Greuter Huxley Algeria Africa: Desf. Huds. ( Huds. Desf. Africa: Algeria, Tunisia Greuter et al. (1984); al. et NCRPIS, Greuter USDA, Tunisia Algeria, Africa: Desf. L. perenne L. perenne L. perenne L. bienne subsp. subsp. subsp. var.

Asia: Armenia, Azerbaijan, Cyprus, Georgia, India, Iran, Iraq, Israel, Israel, Iraq, Iran, India, Georgia, Cyprus, Azerbaijan, Armenia, Asia: Turkey Yugoslavia Spain, Italy, Portugal, Greece, France, Bulgaria, Albania, Europe: Turkey Syria, Pakistan, Jordan, Israel, Iraq, Iran, Asia: AK U.S.: Mexico YT Island, Victoria SK, ON, MB, Canada: America: North Kingdom, Yugoslavia United Ukraine, Switzerland, Spain, Federation, Russian Romania, Poland, Italy, Moldova, Hungary, Greece, Germany, France, Czechoslovakia, Bulgaria, Belarus, Austria, Albania, Europe: Yugoslavia Spain, Italy, Portugal, France, Albania, Europe: U.S.: CO, ID, MT, CA, NV, ND, SD, UT, OR, WV, WA, WY Ockendon (1968) Switzerland, Yugoslavia Italy, Spain, Greece, France, Bulgaria, Austria, Europe: Yugoslavia Ukraine, Switzerland, Spain Romania, Poland, Hungary, Germany, France, Czechoslovakia, Austria, Europe: Turkey, Siberia Federation, Western Italy, Russian Iran, Greece, Bulgaria, Azerbaijan,, Armenia, Albania, Asia: Europe: England, Scotland Switzerland, Yugoslavia Italy, Spain, Greece, France, Bulgaria, Austria, Europe: Africa: Mediterranean Komarov (1969); Komarov Tutin (1980) al. et Mediterranean Africa: Yugoslavia Kingdom, United Ukraine, Spain, Yugoslavia Italy, Bulgaria, Romania, Albania, Moldova, Ukraine, Belarus, Poland, Hungary, Germany, Czechoslovakia, Austria, Europe: U.S.: IA America: North Yugoslavia Ukraine, UK, Spain, Italy, Portugal, Ireland, Greece, France, Bulgaria, Albania, Europe: Tunisia Portugal, Morocco, Libya, Algeria, Africa: Turkey Syria, Federation, Russian Lebanon, Linum species classifi species (endemic and/or naturalized) WWW Location ed by Gill, 1987. byGill, ed . CROPS . ORG

CROP Tutin (1984) al. (1980); et al. et Greuter Huxley (1992) Tutin (1984); al. (1980); et al. et Greuter Komarov (1969); (1988) al. et Komarov Davis personal communication (2006) Meikle (1985) (1966); al. et TutinGuest (1980); al. et DeWet (1993) (1969);Komarov Tutin (1980) al. et (1985); Hooker,Meikle (1875) (1980) al. et

SCIENCE , VOL Reference . 48 , MAY – JUNE

2008 Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. of Hayley, and species 1967). eight (Mosquin are There Canada western to Mexico northern from extending Plains Great the in range avast have also and Florida southern in tributed dis- are species These America. North in primitive most the that CROP distribution of (1963; classifi 1968), extensive an whodeveloped 1993; Rogers USDA, 2007). (Scoggan, Wisconsin and Dakota, North Nebraska, Montana, Minnesota, Kansas, (2n = 16, 57) Asia: Azerbaijan, Georgia, Iran, Russian Federation, Turkey Federation, Russian Iran, Georgia, Azerbaijan, Asia: (2n =16, 57) Table Continued. 2. L. tenue L. sulcatum var. var. L. rigidum L. pallescens L. nervosum L. medium L. usitatissimum L. schiedeanum L. rupestre L. monogynum L. maritimum L. marginale L. gallicum VOthers Group L. viscorum L. hirsutum L. catharticum (2n IV =16) Group L. dolomiticum L. capitatum L. campanulatum (2n =28) III Group Linum L. rigidum

rigidum fi lifolium SCIENCE Ds.Arc:Agra ooc Tutin (1980) al. et Morocco Algeria, Africa: Desf. distributed in Canada (Scoggan, 1993). Gene Plant (Scoggan, Canada in distributed Pursh. North America: Canada: AB, MB, ON, SK Rogers (1963); Rogers (1968); (1963); Rogers Rogers SK ON, MB, AB, Canada: America: North Pursh. Planch. (2n = 30, 36) North America: Canada: ON Rogers (1963); Scoggan (1993); (1963); Magee Scoggan Rogers ON Canada: America: North (2n 36) =30, Planch. L.( (Gray) Engelm. Ex Gray North America: U.S.: NM, TX USDA-NRCS (2006) USDA-NRCS U.S.: TX NM, America: North Gray Ex Engelm. (Gray) (2n = 16, 18) Europe: Bulgaria, Czechoslovakia, Hungary, Yugoslavia Greuter et al. (1984); al. et Greuter Tutin (1980) al. et Yugoslavia Hungary, Czechoslovakia, Bulgaria, Europe: (2n =16, 18) L uoe uti,Fac,Gray tl,San uolvaTutin (1980) al. et Yugoslavia Italy, Spain, Germany, France, Austria, Europe: L. Rg U.S.: TX FL, Rog. Riddel. North America: Canada: MB, ON, QC Rogers (1963); Scoggan (1993) (1963); Scoggan Rogers QC ON, MB, Canada: America: North Riddel. Rg Mexico Rog. A. Cunn. (2n = 80) Australia: New South Wales, Queensland, South Australia, Tasma- Australia, South Queensland, Wales, South New Australia: (2n =80) Cunn. A. Wls.Ai:Amna zrajn eri,Ia,RsinFdrto,Tre Tutin (1980); al. (1988) et al. et Davis Turkey Federation, Russian Iran, Georgia, Azerbaijan, Armenia, Asia: Waldst. L 2 4 8 uoe lai,Blai,Gec,Iay uolvaTutin (1980) al. et Italy, Yugoslavia Greece, Bulgaria, Albania, Europe: L.(2n =24, 28) L fia lei,MrcoTutin (1984) (1980); al. et al. et Greuter Morocco Algeria, Africa: L. Ldb. Asia: Kazakhstan, Kyrgyzstan, Russian Federation, Tajikistan, Komarov (1969) Komarov Tajikistan, Federation, Russian Kyrgyzstan, Kazakhstan, Asia: Ldb. , ao Location Taxon L. Africa: Morocco Rogers (1963); Scoggan (1993) (1963); Scoggan Rogers Morocco Africa: L. L. trigynum Br.Erp:HnayTutin (1980) al. et Hungary Europe: Borb. Forst. (2n = 86) Australia: Australia and New Zealand Allan (1961); Allan (1990) Hnatiuk Zealand New and Australia Australia: (2n =86) Forst. VOL and closely related species are believed to be be to believed are species related closely and S. & C. Mexico Rogers (1969) Rogers Mexico S.&C. L. Cultivated species of species Cultivated L. Linum . 48 , species in North America, reported reported America, North in species MAY L.) Cyprus, Lebanon, Syria, Turkey Davis et al. (1988); al. et (1969) Davis Komarov Turkey Syria, Lebanon, Cyprus, L.) – JUNE

ND, OK, PA, TN, TX, VA, WI PA, TX, TN, ND, OK, MO, NC, NE, MN, MD, MS, KS,KY, IA, LA, IL, U.S.: GA, AR, AL, Yugoslavia, Romania, Ukraine, Bulgaria, Europe: VA MO, NC, SC, TX, OK, KS,LA, IA, U.S.: GA, FL, AR, AL, Europe: Portugal, Spain Portugal, Europe: nia, Victoria, Western Australia Western Victoria, nia, Tutin (1980) al. et U.S.: PA MI, NF, QC Canada: ON, NS, America: North Portugal, Romania, Spain, Yugoslavia Ukraine, Federation, Russian Switzerland, Poland, Netherlands, Kingdom, United Sweden, Norway, Lithuania, Italy, Latvia, Ireland, Hungary, Greece, Germany, France, Finland, Estonia, Denmark, kia, Czechoslova- Bulgaria, Belgium, Belarus, Austria, Albania, Europe: Spain Italy, France, Europe: U.S.: TX FL, 2008

(endemic and/or naturalized) cation and Linum WWW , grown in almost all continents Richharia (1962); (1966) Gill Richharia continents all almost in , grown . CROPS can species, Ameri- North the in each karyotypes basic three are There representing groups). three these among 1969Rogers, for relationships an invasion fl yellow and blue, white, groups: three from the Old (Table 3). ofCanada provinces and States United ofthe states several in distributed extensively is wild relatives (Diederichsen, 2007). of of 5296 accessions collection agermplasm has (PGRC) ofCanada Resources . ORG Linum 833 species in North America can be divided into into divided be can America North in species L. usitatissimum Scoggan (1993) (1999) Ahles and Willis (1972); Hnatiuk (1990) Linum sulcatum and 76 identifi Reference owered species (seeowered species ( ed fl ed n =15) ax Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. ered species), and (iii) species), (iii) ered and raploids ( with ofinterspecifi number A large species many as well as 1968) (Harris, (i) World, of Table Distribution 3. 834 15) of the Great Plains, Plains, 15) Great ofthe var. tives with The rela- geographic of distribution wild North American Hoff and strand 1988; Ell- 1970; Govindaraju, (Kostopoulos, compatibility sexual and heterostyly, of hybridization, direction edness, fl of chronicity availability of pollinators, duration of pollen viability, syn- species, weedy and ofcrop sympatry including factors other infl also is Hybridization 2006). al., et Hall 1999; etal., (Ellstrand variables spatial and temporal, tal, infl be can complexes crop-weed transgenic fl from introgression fortransgene potential the be may there ( number chromosome same have the species these fl cultivated with studies of hybridization reports in included 3have been Fig. in presented species tion of between is an important factor in hybridization and introgression tion experiments, we conclude thatvar. chromosome Plains, number Great of the diploids the with hybridization successful in resulted have tetraploids ley, 1967). Florida the ofboth However, crosses failed to develop seeds orthe seed produce to into failed either crosses of the mature plants (Mosquin and Hay- L. alatum L. sulcatum var. var. var. var. var. L. rigidum L. puberulum L. imbricatum L. hudsonioides L. corymbiferum L. australe L. aristatum On the basis of the results ofinterspecifi results ofthe basis the On rigidum, carteri carteri fi lifolium compactum berlandieri rigidum L. corymbiferum (Small) Winkler (Small) Linum n Pursh Heller , were attempted with two diploid species ( species diploid two with , were attempted n = 30) =30) Riddell pce itiuinReference Distribution Species Engelm. n

ax to them (Fig. 3). However, hybridization of of 3). (Fig. However, them to ax hybridization = 15 is given in Table in =15 3. given is (Engelm.) Heller (Engelm.) =8( and and (Raf.) Shinners Planch. Desf. oeig fl owering, man, 1990; Rieseberg and Wendel, and 1993). 1990; Rieseberg man, species (Fig. 1 and 2). oneexcep- the 1and With (Fig. species L. elongatum Linum rigidum Linum Linum catharticum Linum L. rigidum , none of the native North American American North native , noneofthe Linum n L. alatum =18 fl (yellow oral morphology, genetic relat- genetic morphology, oral species ( species . c crosses of two Florida tet- Florida oftwo c crosses PA, TN, TX, VT, WV, WI PA, TX, TN, MO, NJ, NE, NY, MN, MS, MI, PA, NC, ND, OH, U.S.: Canada: U.S.: Canada: U.S.: U.S.: U.S.: U.S.: U.S.: U.S.: U.S.: var. var. uenced byenvironmen- and TX Osborne and Lewis (1962) Lewis and Osborne (1967) Hayley and Mosquin (2006) communication personal NCRPIS, USDA, MD, KS,KY, CT, IA, LA, IN, AR, IL, AL, GA, (1963) Rogers (1970) Johnston and (1968); Correl Harris (1966) Harris and Rogers TX NM, FL, (1966) Harris and Rogers TX OK, FL, TX TX OK, KS,NM, IA NM AZ, UT, CO, NM, AZ, TX TX ), (ii) (ii) ), rigidum MB, ON, QC, SK. Rogers (1963); Rogers and Harris (1966); Scoggan (1993) (1966); Scoggan Harris and (1963); Rogers Rogers (1993) (1967); (1968); Hayley Scoggan and Rogers Mosquin SK. QC, ON, MB, SK ON, MB, AB, berlandieri n = 15) in North America. =15) North in L. aristatum n uenced by many bymany uenced owered species) species) owered = 9 (blue fl =9(blue ax. Because all all Because ax. and n = 15 (Fig. 3). =15 (Fig. c hybridiza- c , L. rigidum L . n , but all , butall rigidum =15), WWW ow- n = . CROPS

lowing is information onspecifi information is lowing during most content moisture high relatively with green stay ofstems the fi fi are species ofthe Most fl potential the increases butalso 2007) (USDA, appealing aesthetically Limited information is available on the biology and ecol- and biology onthe available is information Limited BIOLOGY ECOLOGY AND and in unmanaged ecosystems (CFIA, 1994; Thomas 1994; Thomas (CFIA, ecosystems unmanaged in and habitats, land 1962), disturbed in (Richharia, roadsides usitatissimum cultivated, being to addition In 2006). (Anonymous, d 40 to of30 period maturation afruit and period, 15 dfl 25 to period, d vegetative a 45 60 to ofafl 1945). cycle Gubin, life The of consists plant ax butterfl and bees, bumble bees, 1938; (Dillman, ies honey- being pollinators 1987), important with Gill, 1938; 1916; 1937; Smith, and Robinson, (Eyre Dillman, 5% of1to range the in reported have been rates tion pollina- butcross species pollinated aself dominantly isfruit a capsule, containing 8 to 10 seeds. is Flax pre- 1938). The (Dillman, arrangement symmetrical ally offi pistil a compound and stamens, 1916) Smith, fi and with (Eyre protandrous slightly and hypogynous, hermaphroditic, fi the to pared com- branches secondary more many produces which fi and oil Linum usitatissimum Linum usitatissimum fi beauti- in and control forerosion mixes in value for their of ogy cation. A long period of fl period Along cation. . ORG

Linum ber. Linseed type fl type plant short ber. Linseed relatively a is ax is found as an escape in waste places, along along places, waste in escape an as found is species. Almost all owering synchronicity with cultivated fl cultivated with ax. synchronicity owering 1987). fl The (Gill, type ber (cultivated fl (cultivated forseed grown is ax) re season (USDA, 2007). The fol- CROP re resistant because leaves and and leaves because resistant re L. (2n =30) (2n L.

owering makes the plant more more plant the makes owering SCIENCE ve sepals, fi sepals, ve c Linum Linum , VOL ve carpels in aradi- in ve carpels . species are noted noted are species 48 species. , ve petals, fi petals, ve MAY owers are are owers – JUNE owering owering

2008 ve L. Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. CROP from male to female. with fl ax offl hybridization ( 3. Potential Figure species related with ax flvey onvolunteer it varied that indicated ax emergence compared to conventionalmany fi fl volunteer as that (1997)twice in reported tillage present ax was systems. etal. Thomas 2006). (Anonymous, time atharvest culty A recent sur- napus sica ( canola and cereals like crops in losses yield in result 1988; 1994), Wall, fl volunteer and usually not ax does (Friesen, apoorcompetitor is 2003). Flax etal., (Leeson resulting in volunteer weed problems in succeeding crops This can result harvest. during soil the to returned being ofseeds inbers an increase in the fl ax 1999).seed Rowland, and (Saeidi seed bank and vigour tests, yellow seed had lower fl seeded brown than seed concentration oil and weight seed vigour than brown yellow revealed that seeded fl (2006) Raney fl several in content oil seed and color, weight, onseed seed 1997). study et al., arecent In fl delay can which development, and growth inhibit may spring the below 10°C in temperature crop, air fl Although 2006). mous, (Anony- available is ofmoisture supply alarge unless soils onsandy notthrive It does fertility. inherent good and ity study. further warrants habitats disturbed offl spread and 1997).et al., establishment The in ax Flax grows best on soils with high water holding capac- holding water high with onsoils best grows Flax Poor management practices may result in large num- large in result may practices Poor management

SCIENCE elds under zero tillage, but at lower densities when butatlower densities tillage, zero under elds L. usitatissimum L.). However, it can cause considerable diffi , VOL . 48 , MAY , but no evidence of hybridization has been reported, while a solid arrow indicates a successful hybridization hybridization asuccessful indicates arrow asolid while reported, been has ofhybridization evidence no , but ax is considered to be a cool season beacoolseason to considered is ax – JUNE ax accessions, Diederichsen and

2008

ax had a higher ahigher had ax owering (Gusta (Gusta owering ax. In WWW n Bras- = 15) in the New World. A dotted line indicates species that may hybridize hybridize may that species indicates line =15) Adotted World. New the in - . CROPS in moist, well-drained, calcareous soils (Scoggan, 1993). (Scoggan, soils calcareous well-drained, moist, in openfi in and prairies, plains, over northern the banks eroded and hills in found 1993). commonly It is (Scoggan, It is known as blue 1966). (Gill, group species ofits many fl in as heterostylous, ax (USDA, 2007) blue fl to white containing orcymes perennial Infl leaves. fl lanceolate-linear to ax linear and node, cotyledonary the from arising stems with height perenne Linum Linum perenne in the direct m 189 0to plants from seeded season growing the throughout plots to 1 to 1510 plants m single dispersed seeds (unpublished data, 2006). data, (unpublished seeds dispersed single many seeds germinate within seed bolls, rather than as fl ofvolunteer clumps in afi ax seedlings tial gene fl fl volunteer that infer substan- to contribute also can ax data These 2006). etal., (Dexter plots seeded conventional season. Birds use seeds and capsules in the fall and winter. It is It is winter. and fall the in capsules and seeds use Birds season. growing the throughout green stay plants because wildlife but also in some provinces of Canada (Scoggan, 1993). (Scoggan, ofCanada provinces some in but also fl Blue 2007), U.S. (USDA, the in notonly distributed been 2007). has and (USDA, seed or herbage ginianus ( fordeer desirable considered also . ORG Blue fl 835 ), antelope ( antelope ), ax is noted to have forage value for livestock and and forlivestock value have forage to noted is ax ow if not controlled. The distribution of small ofsmall distribution The notcontrolled. ow if is a perennial that grows 20 to 80 cm in Antilocapra americana Antilocapra (2n =18) (2n Odocoileus hemionus fl owers. The ax is a native to Eurasia Eurasia to native a is ax orescences are loose ), and birds, either as eld indicates that that eld indicates owers are are owers –2 , in the O. vir- O. elds elds –2

Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. lous, so so lous, under 15% moisture conditions (USDA-NRCS, 2006). (USDA-NRCS, conditions 15%under moisture years forseveral viability retain Seeds open. once capsules some fl of possibility the is there and indeterminate is Flowering 2007). (USDA, conditions land dry under peracre pounds 300 to 200 and conditions irrigated under expected be ofbluefl acre per pounds 700 to of600 can ax yields Seed Dakota. North and Iowa, Oregon, in purposes etation 1987; Fu et al., 2002) it can be artifi be 1987; it can 2002) etal., Fu 1975; Bakker, 1962; and Gill, Zeist van 1938; Richharia, 1904; Dillman, Candolle, (De years for>5000 cultivated occurred (Ellstrand progenitors—has to species crop from alleles domesticated wild their et fl the from al., isolation—halting complete that unlikely is 1999). diverged plants it and ago, Whereas generations crop afewthousand than less relatives fl cultivated the of ax Most has been CONCLUSIONS reported that many species (1971) 2003). Robertson (Zaremba, of pollinated insect be genus to believed butit is mechanisms, onpollination available information nodetailed is There easily. newsites colonize fl produce (4 can cm) plants even small that and fall, early the through continuing June fi are bank (Blake, 1935). Zaremba seed soil the in (2003) persist to 1932). known are seeds The observed that plants unsuccessful (Harris, 1968). were chromosome similarities to assess relatives near its (1968).of Harris to hybridize Attempts study (1970) Rogers and byacytogenetic and by Giannasi genus the in complexes two classifi has fl with hybridize to (1963) ax. Rogers potential have may cultivated flax, Like NewHampshire. and Georgia to Texasto east and L. sulcatum (1963) divided has 2003). Rogers (Zaremba, self-compatible probably and Grooved apex. fl atthe rounded are insect-pollinated is ax 1966). have fi (Gill, Flowers racemes to 70 cm and infl wide, with a single midrib. Plants in vary height from 20 2mm and long about 2cm sessile, alternate, are Leaves as a biennial recorded been also buthas species, plant annual an sidered in North sulcatum, Linum CarolinaLinum sulcatum (Radford et al., 1968). 836 (Gill progeny fertile produce and relatives wild several var. Linum perenne Linum Grooved fl (Stevens, a weed considered be can ax harperi rst evident in early June and begin to fl to begin and June early in evident rst owers present at harvest. Some seeds will shatter shatter will seeds Some atharvest. owers present L. sulcatum ed var. (Small), distributed in Southern America; and L. sulcatum sulcatum known as grooved fl grooved as known con- generally is ax, L. sulcatum orescences loose are panicles axillary or is cultivated for horticultural orreveg- forhorticultural cultivated is L. sulcatum may be self-compatible. be may Riddell (2n =30) (2n Riddell , widely distributed from Manitoba Manitoba from distributed , widely as an intermediate form between form intermediate an as is owers and seeds. It can also also It can seeds. owers and Linum n in two cultivars, cultivars, two in = 15 (Dillman, 1933), it so =15 (Dillman, , which was supported supported was , which ve yellow petals that that petals yellow ve Linum cially crossed with with crossed cially L. sulcatum are homosty- ower by late late by ower L. sulcatum ow of of ow with WWW . CROPS

uted in this area. Two area. ofthem, this in uted distrib- are species wild three only world and the in Science, and Alberta andFood. Agriculture forthisworksupport was provided by InnovationAlberta and also thankDebby Topinka forhertechnicalassistance. Financial for thevaluable commentsandsuggestionson themanuscript. We We thankDr. G. G. Rowland, Dr. MelissaJ. Hills, and Keith Topinka tum fl volunteer and plants wild of size population the reduce also would practices control weed and populations ofwild tion distribu- Alimited occur. eventually will cross a successful that chance the increases environment the in ofplants ber lower than that of greenhouse studies. However, the num- be to predicted is rate hybridization the 1999). nature, In etal., (Ellstrand species those between compatibility cross potential establish it can ecosystem, natural the in ization ofhybrid- success the notpredict does conditions controlled locations. several in sympatry in flgrow may cultivated since relatives worthy wild and ax note- particularly is This species. wild these to transfer to simum between crosses from produced be can hybrids 1972; 2). 1and fertile If Fig. 1970; Seetaram, Godward, 1967; 1967a; and Yermanos, and Gill, and Bari Yermanos in the United States, States, United the in cultivatedLinum lewisii, fl n ax (Gill, offl 3), (Fig. hybridization species these other with ax 1987).interspecifi Among the a transgene can occur. can a transgene of introgression fl whether determine to is warranted ax cultivated with potential outcrossing quantify to study agreenhouse including onspecies, research Further cies. spe- ofthese size orpopulation habitat, preferred time, fl distribution, the about known is 3).Fig. Little fl with outcrossing which (see reported been not has ax for species other several are butthere documented been fl cultivated with hybridization has ax where successful Acknowledgments occur. introgression should species, wild on impact have an could transgene the whether mining deter- before required are expression ofits consequence specifi onthe Details biodiversity. to harm fl to conventional crops, pollen seed via movement and the oftransgenes are concerns volunteerOther mediated gene crop. ofanewtransgenic release the before assessment risk relatives is one of many components of an environmental ow, infl ow, ax and thus minimize gene fl gene minimize thus and ax = 15 species suggests outcrossing may occur (Fig. 1). (Fig. occur may outcrossing suggests =15 species . ORG , have the same karyotype as cultivated fl cultivated as While karyotype ax. same , have the Western Canada is the largest fl largest the is Western Canada region growing ax Although artifi The assessment of the potential for gene fl forgene potential ofthe assessment The

and closely related species, a transgene may be able able be may atransgene species, related closely and uences on non-target species, and the potential potential the and species, onnon-target uences c outcrossing has not been documented for documented notbeen has c outcrossing

n = 9, seems less likely to outcross with with outcross to likely =9, less seems cial hybridization of CROP L. corymbiferum

SCIENCE ow. L. rigidum , VOL is the only species Linum . 48 , c trait and the the and c trait Linum MAY and species under under species ow to wild wild to ow – L. usitatis- JUNE owering owering L. sulca- species

2008 Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. CROP F 1998. D.N. Alstad. and D.A., Andow, Volume 1, Government 1961. H.H. Zealand. ofNew Flora Allan, Y.B. R.G., and G.W. Fu. Allaby, Merriwether, D.A. Peterson, References Cody, W.J. 1996. Flora of the Yukon Territory. NRC Press, Cody, W.J. Yukon Press, NRC of the Territory. Flora 1996. cultivated in 1983. Karyotypes Joshi. K.K. and M.S., Chennaveeraiah, of biology The Agency. Inspection Food 1994. Canadian CFIA. CFIA. 2004b. Canadian Food Inspection Agency. Determina- Directive Agency. Inspection Food Canadian 2004a. CFIA. provinces. prairie Canadian 1987. A.C. ofthe Budd, Flora Budd’s breed- F.N., P.F. plant to and 1967. Knowles. Briggs, Introduction Rep. EMBO health. foryour plants GM 2004. H. Breithaupt, 1970. Interspecifi Godward. M.B.E. G., and Bari, Africa: T.H., B.C. DeWet. and ofSouthern Arnold, 1993. Plants Oxford evolution. and hybridization 1997. M.L. Natural Arnold, group phylogeny Angiosperm ofthe update An 2003. II. APG fl and Growing management 2006. Production, Anonymous. ax: Bond, P., and P. Goldblatt. 1984. Plants of the Cape Flora. J. S. J.S. Flora. Cape ofthe P.,Bond, 1984. Plants P. and Goldblatt. life early and ofseeds germination and 1935. Viability A.K. Blake, ofalpha- 1990. Measurement G.G. Rowland. and R.S., Bhatty, 1969. Infl Godward. M.B.E. G., and Bari, Printer, Wellington, New Zealand. New Wellington, Printer, 112:58–65. Genet. Appl. Theor. usitatissimum offl history domestication ofthe ax ( Evidence 2005. Ottawa, ON, Canada. ON, Ottawa, of wild-species and Canada. ON, Ottawa, Agency, verifi Feb. 2007; tion.gc.ca/english/plaveg/bio/dir/dir9410e.shtml; usitatissimum Linum accessed15 Canada. ON, Feb. 2008. 15 verifi Feb. 2007; accessed bio/dd/dd9824e.shtml; CanadianAvailable at http://www.inspection.gc.ca/english/plaveg/ Foodto soil residues of triasulfuron and metsulfuron-methyl. Inspection Triffi ‘CDC Centre’s Development Crop ofthe safety ofthe tion Agency, Ottawa, Canada. ON, Ottawa, CFIA, ate, 20 Feb. 2005; verifi tion.gc.ca/english/plaveg/bio/dir/dir0007e.shtml; http://www.inspec- at: Available Canada. in novel traits with accessed 2000–2007: Conducting confi Ontario. Ottawa, Canada, Branch,Agriculture Research York. New Corp., Publishing Reinhold ing. 5:1031–1034. ria, South Africa. Preto- Institute, Botanical National distribution. and Names York. New Univ. Press, 141:399–436. Soc. J.Linnean Bot. II. APG classifi Canada. Commission, Development Flax diagnostic guide. Flax 91:572–578. Entomol. J.Econ. alleles. Council of Canada & Saskatchewan African Bot. 13:1–455 Bot. African 5:405–460. Monogr. Ecol. plants. ofprairie history 67:364–367. Soc. Chem. fl oil Oil ofedible Am. development the J. ax. in acid linolenic 12:799–802. Cytol. size on of the radiosensitivity Linum

SCIENCE d’, aflax ( cation for the orders and families offl families and orders forthe cation . Euphytica 19:443–446. . Euphytica , VOL L.) from genetic diversity of the ofthe diversity genetic L.) from . ed 15 Feb. 2008. Canadian Food Inspection 48 Linum usitatissimum , L. (Flax). Available at: http://www.inspec- at: Available (Flax). L. ed 15 Feb. 2008. Plant Products Director- Products Plant 15ed Feb. 2008. Linum MAY . – . Cytologia (Tokyo) 48:833–841. . Cytologia JUNE ned research fi research ned

Linum 2008 2

screen for rare resistance screen for resistance rare species. Can. J. species. Can. Genet. ofchromosome uence L.) cultivar tolerant eld trials of plants owering plants: plants: owering in c crosses sad2 locus. locus. ed 20 ed WWW Linum . CROPS Diederichsen, A. 2007. Ex situ collections of cultivated fl of cultivated ax ( collections situ 2007. Ex A. Diederichsen, Dexter, J., K. Topinka, and L.M. Hall. 2006. Emergence peri- Emergence 2006. Hall. L.M. and Topinka, J., K. Dexter, Paul, Kegan plants. ofcultivated 1904. Origin A. Candolle, De the ofTurkey and Tan. Flora 1988. K. and P.H., Mill, R.R. Davis, Darlington, C.D., and A.P. Wylie. 1955. Chromosome atlas of atlas A.P. C.D., 1955. and Wylie. Chromosome Darlington, J. agriculture. into plants P.J. oftransgenic Dale, 1993. release The Czerepanov, S.K. 1995. Vascular plants of Russia and adjacent states classifi of system 1981. integrated A. An Cronquist, vascular ofthe 1970. Manual Johnston. M.C. D.S., and Correl, Lion ofBombay. Red Presidency T. ofthe Cooke, 1903. Flora ofgenet- release J.P. The and 2003. Nap. T.R. A.J., Glare, Conner, Dillman, A.C. 1953. A.C. Classifi Dillman, fl Cultivated 2003. Richards. K. and A., Diederichsen, weight seed colour, Seed J.P. and A., 2006. Raney. Diederichsen, Durrant, A. 1986. Flax and linseed. p. 190–193. 190–193. p. linseed. and Flax 1986. A. Durrant, fl in numbers 1933. A.C. Chromosome ax ( Dillman, fl in 1936. A.C. Improvement Agricul- ax. of Book Year Dillman, fl in Agron. Soc. crossing Am. J. ax. 1938. Natural A.C. Dillman, molecular and Phenotypic Y.B. 2006. and A., Fu. Diederichsen, Ellstrand, N.C. 1988. Pollen as a vehicle for the escape ofengi- escape forthe avehicle as Pollen N.C. 1988. Ellstrand, Ellstrand, N.C., H.C. Prentice, and J.F. Hancock. 1999. fl J.F. Gene and Hancock. N.C., Prentice, H.C. Ellstrand, Hoff C.A. N.C., and Ellstrand, Eyre, J.V., and G. Smith. 1916. Some notes on the Linaceae—The J.V., Linaceae—The Eyre, 1916. onthe notes Some G.Smith. and . ORG Institute of the U.S., Fargo, ND. U.S., Fargo, ofthe Institute Flax 2006 22–26, March Dakota. North Fargo, States. United seeding. direct and tional Resour. Crop Evol. 54:661–679. Crop Resour. usitatissimum odicity of volunteer fl ofvolunteer odicity ax ( UK. London, Ltd., Co. and Trench, Trulener UK. Univ. Press, Edinburgh Islands. Aegean East fl Agric. Sci. 120:1–5. UK. Cambridge, Univ. Press, Cambridge USSR). former (The York. New Univ. Press, Columbia plants. ering TX. Renner, Foundation, Texas ofTexas. Research plants London. Street, Fleet Court, J.33:19–46. Plant assessment. risk ofecological view modifi ically wood Academic Publishers, Amsterdam, The Netherlands. The Amsterdam, Publishers, Academic wood Evol. 53:77–90. Crop Resour. in content oil seed and ence 78:409. DC. Washington, USDA, ture. 30:279–286. DC. Washington, USDA, 1064. genus The (ed.) Flax: N.Westscott and Muir A. Linum flvated ax ( (RAPD) diff 125:372–377. Breed. Plant ofCanada. Resources Gene Plant neered genes? p. s30-s32. p. s30-s32. genes? neered UK. London, Technical, Scientifi Longman plants. (ed.) ofcrop monda Evolution reduction. Bioscience 40:438–442. Bioscience reduction. forrisk Strategies genes: forengineered avenue ofescape UK. Cambridge, Publications, Elsevier Publication. Special Evolution and Ecology in Biotechnology/Trends in Trends organisms. engineered ofgenetically release (ed.) Planned cross pollination offl 5:189–197.Genet. J. pollination ax. cross 30:539–563. Rev. Syst. Ecol. Annu. relatives. wild their into plants domesticated from introgression and owering plants. George Allen and Unwin, London. Unwin, and Allen George plants. owering 837 L.: taxonomy and germplasm conservation. p. 22–54. p. 22–54. conservation. germplasm and taxonomy L.: Linum usitatissimum Linum L.) and other species of the genus genus ofthe species other L.) and ed crops into the environment- Part II. Over- II. Part environment- the into crops ed erentiation of four infraspecifi offour erentiation Linum usitatissimum Linum offlcation Rep. 1946. varieties ax In Linum usitatissimum Linum In man. 1990. Hybridization as an an as Hybridization 1990. man. Proc. of 61st Flax Institute of the ofthe Institute of61st Proc. Flax J. Hodgson and A.M. Sugden Sugden A.M. and J.Hodgson L. subsp usitatissimum accessions heldby accessions ofculti- c groups ax and the genus genus the and ax Linum L.) in conven- L.) in cation of fl In Linum N.W. Sim- Linum L. Genet. ). Genet. Genet. ). . Hard- ). Sci- ). Linum c and c and ow- ow ow In

Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. Gubin, A.F. 1945. Cross pollination offi A.F. pollination 1945.Gubin, Cross (ed.). G.Long 1984. Med-Checklist. and W.,Greuter, Burdur, H.M. out between relationship onthe Anote D.R. 1988. Govindaraju, among relationship Evolutionary 1966. K.S. Gill, Research, ofAgricultural Council Indian 1987. K.S. Linseed. Gill, 1970. signifi Taxonomic Rogers. C.M. and D.E., Giannasi, Gusta, L.V., B.J. O’Connor, and R.S. Bhatty. 1997. Flax ( 1997. L.V., Bhatty. Flax Gusta, R.S. and B.J. O’Connor, A.Z.W.A. and Wizarat. Wizarat, A.Z. Townsend, C.C. E., Guest, on Studies 1967b. Cytogenetic Yermanos. D.M. and K.S., Gill, on Studies 1967a. Yermanos. Cytogenetic D.M. and K.S., Gill, 2002. K.W. and Richards. Diederichsen, A. Y.B.,Fu, G.Peterson, culti- in variation ofRAPD patterns Geographic Y.B.Fu, 2005. fl Canada—A 2007a. ofCanada. Council Flax 838 Friesen, G.H. 1988. Annual Grass Control in Flax ( Flax in Control Grass Annual 1988. G.H. Friesen, Can- in development Varietal 2007b. ofCanada. Council Flax Hammond, J.J., and J.F. Miller. 1994. Flax breeding in the United United the in breeding 1994. Flax J.F. J.J., and Miller. Hammond, Intraspe- 2006. Nielson. R.L. and Topinka, A.K.C. L.M., Hall, Hnatiuk, R.J. 1990. Census of Australian vascular plants. Australia Australia plants. vascular Australian of Census 1990. R.J. Hnatiuk, J.W. and Ownbey, M. Thomp- Cronquist, A. C.L., Hitchcock, offl families 100 1988. M. Hickey, Oxford world. ofthe plants V.H. 1993. Flowering Heywood, Altertum. im Flachskultur die und Flachs den O.Heer, 1872. Uber North in evolution and numbers B.D. 1968. Chromosome Harris, Conservatoire Botaniques de Geneve, Geneva, Switzerland. Geneva, Geneve, de Botaniques Conservatoire fl gene and rate crossing 7:627–631. Sci. Crop number. mosome India. Delhi, New of fl Iraq, Baghdad, Iraq. of ofRepublic ofAgriculture Ministry ofIraq. Flora 1966. 26:30–31. Genus 7:623–626. Sci. Crop number. mosome Genus CA. Riverside, ofCalifornia, University Ph.D. diss. genus the fl ofseven in species relationship ax ofgenetic analysis RAPD flvated 45:1084–1091.Sci. Crop ax. Canada. fi veri- 2007; 15 June accessed php?p=growing11&mp=growing; http://www.fl at: Available ada. Canada. MB, Winnipeg, of Canada, verifi 2007; 4 Apr. accessed what; http://www.flaxcouncil.ca/english/index.php?p=what2&mp= States. p. 53–40. p. 53–40. States. Canada. Quebec, Anne-de-Bellevue, Sainte- Society, Weed Science Canadian Canada. in crops tant Canadian Weed Science: Thecifi fi 77:97–99. Sci. J.Plant Can. yield. seed and ing usitatissimum simum fl WA. Seattle, Press, Washington son. 1969. Vascular plants of Pacifi UK. Cambridge, Univ. Press, UK. Oxford, Univ. Press, Zurich. Gesellschaft, Naturforschenden der Neujahrsblatt of species American CT. Haven, New Station, Experiment Agricultural Connecticut The Symposium. Flax World ofthe Fiber Proc. ora and fauna series no.11. Australian Biological Resources Resources Biological no.11. series Australian fauna and ora ed 15 Feb. 2008. Flax Council of Canada, Winnipeg, MB, MB, Winnipeg, ofCanada, Council Flax 15ed Feb. 2008. fl c gene oral pigments in in pigments oral ) with quizalofop. Weed Technol. 2:144–146. quizalofop. ) with Linum Linum Linum ow: Infl ow: L) responses to chilling and heat stress on fl on stress heat and chilling to responses L) . I. Hybrids among taxa with 15 as haploid chro- haploid 15 as with taxa among Hybrids . I. . II. Hybrids among taxa with 9 as haploid chro- haploid 9as with taxa among Hybrids . II. . Genet. Resour. Crop Evol. 49:253–259. Crop Resour. . Genet. In uencing factors. p. 87–100. factors. uencing J.F. Anderson and M.A. Schiavoni (ed.) Schiavoni M.A. and J.F. Anderson Linum Linum ow in plants. Heredity 61:401–404. Heredity plants. ow in . Am. J. Bot. 55:1197–1204. J.Bot. . Am. (Linaceae). Brittonia 22:163–174. Brittonia (Linaceae). ed 15 Feb. 2008. Flax Council Council ed 15 Flax Feb. 2008. decaderst of herbicide resis- owering plants. Cambridge Cambridge plants. owering axcouncil.ca/english/index. c Northwest. University of University cNorthwest. flber Bee ax. World ax leader. Available at: at: Available leader. ax Linum Linum usitatis- Linum In Topics in in Topics species. species. ower- cance WWW Linum . CROPS Kostopoulos, S.S. 1970. Genetics of heterostyly and incompatibil- and ofheterostyly 1970. Genetics S.S. Kostopoulos, ofthe Science Academic V.L.Komarov, USSR. 1969. ofthe Flora genus ofthe studies 1929. M. Cytological Kikuchi, fl tolerant Glyphosate 1988. ax McHughen. A. M.C., and Jordan, James, C. 2003. Preview: Global status of of dictionary commercialized society horticultural royal new The 1992. A. Huxley, trans- Martzenitzin, K.K. 1927. The chromosomes of some species species ofsome 1927. K.K. chromosomes The Martzenitzin, Preliminary 2000. von Bothmer. R. and O. E., Diaz, Mansby, A Northeast: D.W., ofthe 1999. Flora Magee, Ahles. H.E. and Lutkov, A.N. 1939. ofLon- Society Mass Royal The 1857. C. plantarum. Species productionLinnaeus, of tetraploid fl ax plants by Saskatchewan 2003. J.Y., Bremzil. C. and Leeson, Thomas, A.G. p. 416–430. 1989. Linseed. C.D. Dybing. and Lay, C.L., pos- genetic and plants—Methods in F. 1929.Laibach, Ectogenesis McHughen, A. 1989. Agrobacterium mediated transfer of transfer mediated 1989. A. Agrobacterium McHughen, Flax. Linseed 2002. A. McHughen, and relationships Evolutionary 2005. Simpson. B.B. J., and McDill, fi textile The 1908. J.M. Matthews, Hricova, A. 2002. Transformation and characterisation of transgenic J.D.Hooker, 1875.Co., fl & L. Reeve The India. British of ora scheme World 1992. geographical Brummit. R.K. and S., Hollis, . ORG fornia, Riverside, CA. Riverside, fornia, in ity Leningrad. and Moscow USSR, Genet. 4:201–210. Cell Rep. 7:281–284. plants from York. New ISAAA, No. 30. crops:2003. genic Agrobacterium UK. Co., London, McMillan gardening. mediated gene-transfer. Plant of the genus genus of the mum fl Swedish in diversity ofgenetic study ax ( MA. Amherst, Press, ofMassachusetts York.New University manual of the vascular 22:175–179. LUrss. De Sciences fl Des LAcademie De Rendus Comptes treatment. colchicine don, UK. Canada. SK, Saskatoon, Canada, Agri-Food and Agriculture Station, Research 03–1. Saskatoon Rep. 2003. in crops pulse and oilseed ofcereal, survey weed York.New McGraw-Hill, world. ofthe crops (ed.) Oil al. et Robbelen 20:201–208. J. Hered. seed. the in dying otherwise embryos ofpropagating sibilities New York.New Dekker, Marcel Plants. ofTransgenic (ed.). Handbook The Texas, USA. onomists, Tax- ofPlant Society American the and ofAmerica Society Botanical 2005. Texas, Aug., Austin, Taxonomists. of Plant Society American the and ofAmerica Society Botanical of the in biogeography UK. London, Ltd., Hall and Chapman properties. chemical and scopical 17:273–288. Comenius University in Bratislava. Bratislava, Slovakia. Bratislava, Bratislava. in University Comenius fl London. PA. Pittsburgh, University, Mellon Carnegie Documentation, Botanical for Institute Hunt science, forplant databases on taxonomic group working International distributions. plant for recording 2601, Australia. Act Canberra (ABRS), Study ax plants, transgene expression and its variability. Ph.D. diss. Ph.D. diss. variability. its and expression transgene plants, ax

). Genet. Resour. Crop Evol. 47:417–424. Crop Resour. ). Genet. Linum grandifl orum Linum Linum L. Bul. Appl. Bul. L. Bot. Genet. Plant Breed. CROP (Linaceae). Desf. M.Sc. diss. University ofCali- University diss. M.Sc. Desf. ora of New England and adjacent adjacent and England ofNew ora

SCIENCE bers, their physical, micro- physical, their bers, In , G. Khachatourians et al. In VOL Joint National Meeting . 48 , MAY Linum usitatissi- Linum Linum – JUNE . Jpn. J. . Jpn.

In 2008 G. Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. Mosquin, T., and D.E. Hayley. 1967. Evolutionary relationships of T., relationships Hayley. 1967. D.E. and Mosquin, Evolutionary biotech- and cell Plant 2005. P. and O. S., Anna. Bohus, Millam, CA. Sabbott, Rudolf. William ofCyprus. R.D. 1985. Flora Meikle, CROP Nagao, S. 1941. Cytogenetics in the genus Amosova, A.V. Samatadze, T.E. Lemesh, V.A. O.V., Muravenko, McSheff E.O. and Bhatty, R.S. F.A. Holm, G.G. A., Rowland, McHughen, envi- with Flax 1995. Transgenic F.A. Holm. and A., McHughen, Robinson, B.B. 1937. Natural cross pollination studies in fi in studies pollination cross 1937. B.B. Robinson, Natural J. United-States. Southeastern in 1971. K.R. Linaceae Robertson, the and J.F. zones and Wendel. 1993. L.H., Hybrid Rieseberg, Oiseeds Central Indian The Linseed. 1962. R.H. Richharia, Ver- Drunund Academische Iranica. 1963. Flora K.H. Rechinger, modifi Gray. 1993. A.J. Genetically and A., Raybould, fl the in studies genus ax Ray, 1944. Cytological C.J. ofthe 1968. Manual Bell. C.R. and Ahles, H.E. A.E., Radford, Zakharch- N.S. Kalyaeva, Polyakov, M.A. A.V., O.F. Chikrizova, of numbers W.P., Chromosome 1962. Osborne, W.H. and Lewis. syn- acid offatty 1997. J.G. Regulation Jaworski. and J.B., Ohlrogge, 1968. Walters. D.J., S.M. Ockendon, and the in studies D.J. 1968. Biosystemic Ockendon, ofnatu- morphology pollen D.J. and 1971.Ockendon, Cytology The 2003. Conner. A.J. and J.P.,Nap, Escaler, M. P.L.J. Metz, the southern Florida populations of populations Florida southern the 82:93–103. Cult. Organ Tissue in studies nology f l ax species and their hybrids. Russ. J. Genet. 39:414–421.Genet. J. Russ. fl hybrids. their and species ax related closely forthree markers molecular and chromosomal tyuleva, and A.V. Zelenin. 2003. Genome comparisionsZ.E. with Grushetskaya, V.K. Popov, O.Y. J.Can. Bot. 45:1277–1283. Semenova, V.L. Kho- usitatissimum acterization of transgenicSci. 77:641–643. 1997. Triffi CDC Kenaschuk. sulfonylurea-resistant 4:3–11. Res. genic Trans- attributes. sustainable agronomically and ronmentally fl ax ( Cell Rep. 8:445–449. chlorsulfuron resistance to commercial flax cultivars. Plant Committee, Hyderabad,Committee, India. Iran. Tehran, Graz., laganstalt 30:199–219.Ecol. hybridization with wild relatives: A UK perspective. J. Appl. 31:441–448. J. Bot. NC. Hill, Chapel fl vascular mation of fi Y.I. and transfor- Bur’yanov. The N.V. 1998. enko, Balokhina, Linum 48:109–136. Biol. Mol. Plant Physiol. Rev. Plant Ann. thesis. UK. Cambridge, Univ. Press, bridge Cam- T.G. Europaea. (ed.) V.H. Tutin Flora and Heywood 67:787–813. Phyto. New group. 70:599–605. Phytol. artifi and ral J.33:1–18. Plant regulations. and status ofcurrent Overview modifi ofgenetically release 17:109–116. Arnold Arbor. 52:649–665. UK. Oxford, Univ. Press, Oxford plants. in consequences its and gression 70–109. p. process. evolutionary

SCIENCE rey, S.A., A. McHughen, and M.D. Devine. 1992. Char- 1992. Devine. M.D. and McHughen, A. rey, S.A., from southern United States and Mexico. SIDA 1:63–68. Mexico. and States United southern from ora of Carolina. University of North Carolina Press, Press, Carolina ofNorth University ofCarolina. ora , ber fl VOL cial tetraploids in the the in tetraploids cial ). Theor. Appl. Genet. 84:480–486. Genet. ). Appl. Theor. . ax plants. Russ. J. Plant Physiol. 45:764–769. Physiol. J.Plant Russ. plants. ax 48 , Linum usitatissimum- MAY – JUNE ed crops into the environment. I. I. environment. the into crops ed d transgenic fl J.Plant d transgenic Can. ax.

2008 In R.G. (ed.) Harrison Intro- Linum perenne Linum

Linum rigidum Linum Linum A review. Plant Cell Cell Plant Areview. Linum L. p. L. 206–211. . Jpn. J. Genet. J.Genet. . Jpn. Linum perenne Linum group. New ed crops and and crops ed Linum (Linaceae). WWW Linum . Am. . Am. ber In . CROPS

Tutin, T.G., V.H. (ed.). D.H. Valentine and Tutin, T.G., Burges, N.A. V.H. Heywood, Heywood, N.A. Burges, D.M. Murre, 1997. Frick. Man- B.L. and A.G., R.F. D.J. Wise, Kelner, Thomas, D.H. Abstamm. Induct. Z. evolution. T. and 1925.Tammes, Mutation genus ofthe T. genetics The 1928. Tammes, by produced ofseeds weight and number 1932. The O.A. Stevens, Sharma, R.R., and A.N. Khanna. 1964. Development of fruit in in offruit 1964. Development Khanna. A.N. and R.R., Sharma, 1972. Interspecifi A. Seetaram, of Museum National ofCanada. 1993. H.J. Flora The Scoggan, 1999. eff G.G. The G., Rowland. and Saeidi, Stephens, G.R. 1997. A manual forfi 1997. G.R. Amanual Stephens, offi data Production 2006. Canada. Statistics Briefi 2005. H.V.,Smith, J.Jimmerson. and p. 67–87. 4Linaceae. 1907. J.K. Family Small, muta- ofgiant 1939. Chopinet. R. Appearance and M., Simonet, Rosberg, and R.J. 1996. Underexploited industrial temperate, chromosome new Some 1966. B.D. Harris. and C.M., Rogers, Rogers, C.M. 1968. Yellow 1968. fl C.M. Rogers, of species owered Rogers, C.M. 1963. Yellow C.M. fl Rogers, of species American ofNorth 1969. C.M. Relationships Rogers, 1941. A.R. Interspecifi Rogach, . ORG Europaea: Rosaceae to Umbelliferae. Cambridge Univ. Press, Univ. Press, Cambridge Umbelliferae. to Rosaceae Europaea: Valentine, S.M. Walters, UK. Cambridge, Univ. Press, Cambridge and Europaea. 1980. Flora D.M. Webb (ed.). 1968. Canada. MB, Winnipeg, Flora crop production tillage conventional and zero Comparing systems. survey: weed itoba Rep. 97–1. Manitoba Agriculture, 36:417–426. Vererb.u. Lehre 19:784–794. J.Bot. Am. weeds. CT. Haven, New Station, Experimental Agricultural necticut ica 21:489–495. ica Canada. ON, Ottawa, Sciences, Natural fl 79:315–319. in Sci. J.Plant vigour Can. seed ax. and ongermination concentration acid linolenic and colour seed VA. Alexandria, Press, 1995. ASHS Oct., 22–25 Indiana. Indianapolis, gies. Technolo- New Opportunities, New Crops: onNew posium nus Nijhoff nus Marti- Genetica. (ed.) Bibliographia W.A. and Goddijn Lotsy Canada. ON, Ottawa, Agency, Statistical National Canada’s verifi 2007; 17 Feb. accessed prim11b.htm; http://www40.statcan.ca/l01/cst01/ at: Available crops. ity MT.Bozeman, verifi University, State Montana Center, 2006; Policy Marketing June 18 cultural accessed ing56.pdf; http://www.ampc.montana.edu/briefi at: Available York.New Garden, Botanical Flora. American (ed.) North Garden ical 209:238–240. Sciences Des LAcademie De Seances Des Hebdomadaires fl fl and big with ax periwinkle rapeseed, in polyploids and tions J. 8:34–80. some interspecifi fi in numbers North America. Brittonia 15:97–122. Brittonia America. North 20:107–135. Brittonia America. North Western and America Central Linum (Hybridz) 2:84–88. 29:644–649. fl Agron. Soc. Am. J. ax. owers after the application of colchicine. Comptes Rendus Rendus Comptes ofcolchicine. application the after owers ber crops. p. 60–84. p. 60–84. crops. ber 839 (fl96:176–190. Club Bot. ax).Torry Bull. , The Netherlands. , The Linum c crosses of the genus genus ofthe c crosses . Madrono 18:179–180.. Madrono In owered species of species owered Proc. of the Third National Sym- National Third ofthe Proc. c hybridization in in c hybridization in c hybrids ber fl ax production. The Con- The production. ax ed 15 Feb. 2008. Agri- 2008. Feb. 15 ed Linum ng No. 56: Flaxseed. No. Flaxseed. 56: ng Linum In ect of temperature, oftemperature, ect Linum New York New Botan- ed 15 Feb. 2008. special- and eld Linum . Indian Oilseeds Oilseeds . Indian Linum . p. 1–34. . Vest Gibrid . Vest Gibrid in Eastern ngs/brief- Linum . Euphyt- In J.P. in Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved. 840 Wall, D.A. 1994. Fluazifop-P tank-mixtures with clethodim for clethodim with tank-mixtures 1994. D.A. Fluazifop-P Wall, fl in studies ax ( genetic Molecular J.2006. Vromans, Len- plants. ofcultivated origin the in Vavilov, Studies 1926. N.I. cul- V.W., Zeist, van forlinseed 1975. Evidence Bakker. H.H. and USDA-NRCS. 2006. United States Department of Agriculture- USDA. 2007. United States Department of Agriculture. National nol. 8:673–678. nol. fl in control grass ax ( annual The Netherlands. simum Moscow, Russia. Priklad, Inst. Vsesoiuz. ingrad, 2:215–219. Sci. J.Archaeol. bc. 6000 before tivation verifi 2007; 22, March accessed usda.gov; http://plants. at: Available database. PLANTS The NRCS. MD. Beltsville, Laboratory, 2007; verifi gov/cgi-bin/npgs/html/taxon.pl?22326; accessed February/8, http://www.ars-grin. at: Available Network—(GRIN). tion Informa- Resources Germplasm Program. Resources Genetic Cambridge, UK. L.). Ph.D. diss. Wageningen University, Wageningen, Wageningen, University, Wageningen L.). Ph.D. diss. ed 20 Feb. 2008. National Germplasm Resources Linum usitatissimum Linum ed 15 ed Feb. 2008. ). Weed Tech- Weed ). Linum usitatis- Linum WWW . CROPS Zohary, D., and M. Hopf. 2000. Domestication of plants in the the in ofplants Domestication 2000. Hopf. D., M. and Zohary, Zeven, A.C. 1982. Dictionary of cultivated plants and their cen- Zaremba, R.E. 2003. Yermanos, D.M., and K.S. Gill. 1967. Induction of polyploidy in in 1967. ofpolyploidy Gill. Induction K.S. D.M., and Yermanos, p. 111–120. linoideae-eulineae. I. trib 1931. H. Linaceae, Winkler, Melborne Victoria. in plants to 1972. Ahandbook J.H. Willis, of assessment Risk 2003. Poppy. G.M. and Sweet, J. M.J., Wilkinson, . ORG Oxford, UK. Oxford, Univ. Press, Valley. Oxford Nile the and Europe West Asia, Old World: The Netherlands. The Wageningen, origin Docmentation, and Publishing forAgricultural Center plants. and lower and spread trees forest ornamentals, excluding ofdiversity, ters of cultivated MA. Framingham, Society, Flower Wild plants England New England. forNew plan research and servation in Linum Germany. Leipzig, Nutzpfl ed. 2nd Engelmann, W. anzen. den insbesondere Arten, wichtigeren und Gattungen ihren In Australia. Victoria, Carlton, Univ. Press, 8:208–212. Sci. Plant Trends gridlock? Avoiding plants: GM

A. Engler (ed.) Die nature-lichen Pfanzenfamilien nebst nebst Pfanzenfamilien (ed.) nature-lichen Die Engler A. species. Crop Sci. 7:423–427. Linum sulcatum CROP

SCIENCE riddell (grooved flax): Con- , VOL . 48 , MAY – JUNE

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