Journal of Biology, Agriculture and Healthcare ISSN 2224-32 ! "#a$er% ISSN 222&- '3( ")nline% *ol.3, No.

+,, 2 +3

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Identification of Badh2 Mutation Type among Indonesian Fragrant Rice Varieties
-.arot Sasong/o Ha0i Seno+1, I 2ade Arti/a+, 3aras Nurcholis+, 4ri Jo/o Santoso2, 5urniawan 6 4ri.at0i/o2, Ba07ang #ad0adi+, -ewi #ra$tiwi+, Ja$ 2ai 8hing+, 9ainal Ali0 2as:ud3 +.-e$art0ent of Bioche0istry, Bogor Agricultural ;ni<ersity, Bogor-3est Ja<a +==! , Indonesia 2.4he 8enter for Agricultural Biotechnology and >enetic 6esources 6esearch and -e<elo$0ent, -e$art0ent of Agriculture, 2inistry of Agriculture 6e$u7lic of Indonesia 3.-e$art0ent of 8he0istry, Bogor Agricultural ;ni<ersity, Bogor-3est Ja<a, Indonesia 1 ?-0ail of the corres$onding author@ ha0isenod.arotAg0ail.co0 Abstract 4he $re0iu0 $rice as well as the high and increasing world 0ar/et de0and for fragrant rice has triggered the de<elo$0ent of <arious fragrance 0ar/ers. 4he a$$lication of these 0ar/ers on genoty$ing of <arious $o$ular Indonesia rice <arieties is re$orted in this $a$er. #86 $rofiles of $o$ular Indonesia non-fragrant "8iherang, Bat0awati% and fragrant "#andan wangi, 6o.o Cele, 2enti/ 3angi, >unung #era/, #ulu 0andoti, #are 5e07ang, Sintanur% rice <arieties were co0$ared using aro0atic 0ar/ers of Brad7ury et al. "2 &7%, Cang and Buu "2 !%, Shi et al. "2 !%, and Sa/thi<el et al. "2 '%. Bor co0$arison, I6=4, Ni$$on7are and 4ai$ei 3 ' <arieties were included. 6ice geno0ic -NA was isolated fro0 young lea<es using the 0ethod as descri7ed 7y -oyle and -oyle "+'' %, #86 a0$lified using each of the a7o<e fragrance 0ar/ers and #86 $roducts were analysed 7y agarose-gel-electro$horesis. Bragrance 0ar/ers of Brad7ury et al. "2 &7%, Shi et al. "2 !%, and Sa/thi<el et al. "2 '% were only a7le to discri0inate fragrant 2enti/ 3angi and >unung #era/ fro0 non-frangrant rice <arieties, while other fragrant rice <arieties "#andan wangi, 6o.o Cele, #ulu 0andoti, #are 5e07ang, Sintanur% showed si0ilar 7and $atterns as those of non-frangrant rice <arieties. 4his suggests there are at least two grou$s of 7adh2-deletion $atterns a0ong Indonesia fragrant rice <arieties. >rou$ + include 2enti/ 3angi and >unung $era/, while grou$ 2 includes #andan wangi, 6o.o Cele, #ulu 0andoti, #are 5e07ang and Sintanur. )nly the 62 223 0ar/er of Cang and Buu "2 !% was a7le to distinguish 7oth fragrant grou$s fro0 non-fragrant 8iherang. 4he difference in 62233-a0$licon siDes 7etween fragrant grou$s + and 2, also su$$orts the <ariation of 7adh2 0utation $attern a0ong Indonesia fragrant rice. Keywords: Brad7ury, BadeE,-&, B2-?,, B2-?2A, 62 223, fragrance, #andan 3angi ! Introduction *arious fragrance analysis ha<e 7een de<elo$ed to assist selection in rice 7reeding for aro0a. 4hose 0ethods include@ organole$tic 7ased on taste "6ein/e et al. +''+, #etro<e et al. +''=% or odor "Sood and SidiF +',!, #aule and #owers +'!'%G BADH2 enDy0e-7ioassay "Sri<ong et al. 2 !%G 2-A# detection using 0ass s$ectro$hoto0etryHselecti<e ion 0onitoring "4anchoti/ul and Hsieh +''+%, >8 "CorieuE et al. +''=, 3id.a.a et al. +''=% or sta7le isoto$e "Ioshihashi 2 2% and the use of 6BC# "6estriction Brag0ent Cength #olyor$his0s%, SN#s "Single Nucleotide #oly0or$his0s% or SS6s "Si0$le SeFuence 6e$eats% 0ar/ers "Ahn et al. ''2, 8ordeiro et al. 2 2%. Howe<er, these 0ethods are considered difficult, la7our intensi<e, unrelia7le, ti0e consu0ing and reFuire large sa0$les, or are una7le to $redict the fragrace gene or allele statues "Brad7ury et al. 2 &7, Sa/thi<el et al. 2 '%. In addition, 0ost of these 0ethods are ina$$lica7le for fragrance analysis in heteroDygote $rogenies, since fragrance in rice is a recessi<e trait "CorieuE et al. +''=, -ong et al. 2 , Brad7ury et al. 2 &a, BorFuis et al. 2 !%. Bragrant-0ar/er-assisted #86, followed 7y agarose-gelelectro$horesis, has 7een considered as the 0ost $otential and a0ena7le a$$roach for routine fragrance genoty$ing in large scale 7reeding 0aterials, due to its low-cost, s$eed, si0$licity, and the sensiti<ity of this 0ethod "Brad7ury et al. 2 &7, Cang and Buu 2 !, Shi et al. 2 !, Sa/thi<el et al. 2 '%. 4he finding of an ! 7$ and 3 SN#s 0utation in the badh2 gene of fragrant rice has led to the de<elo$0ent of a s$ecific fragrant 0ar/er-assisted #86 0ethod "Brad7ury et al. 2 &a, BorFuis et al. 2 !% that can 7e used to facilitate early selection of non-fragrant and fragrant rice, as well as their cross and 7ac/cross $rogenies which are always heteroDygous. 4he siDe difference in the badh2 gene 7etween non-fragrant and fragrant rice results in a different a0$licon siDe "Brad7ury et al. 2 &7%. Cang and Buu "2 !% ha<e also re$orted fragrant 0ar/er 62 223 that $roduces ∼+= and ∼+2 7$ a0$licons for non-fragrant and fragrant rice, res$ecti<ely. Besides the eEon ,-deletion "badh2.7% that had 7een $re<iously re$orted "Brad7ury et al. 2 &a, BorFuis et al. 2 !%, Shi et al. "2 !% also re$orted another deletion in the eEon2 of badh2 gene " badh2.2%, and constructed fragrance 0ar/ers B2-?, and B2-?2 that flan/ed the ! 7$ or , 7$ deletion in badh2.7 or badh2.2, res$ecti<ely. Howe<er, the suggestion of using #A>? 0ethod to resol<e #86 $roducts were not Fuite $ractical for routine genoty$ing in large scale 7reeding 0aterialsHger0$las0s. &'

Journal of Biology, Agriculture and Healthcare ISSN 2224-32 ! "#a$er% ISSN 222&- '3( ")nline% *ol.3, No.+,, 2 +3

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Although <arious fragrance 0ar/ers "Brad7ury et al. 2 &7, Cang and Buu 2 !, Shi et al. 2 !, Sa/thi<el et al. 2 '% ha<e 7een re$orted which succesfully discri0inate 7etween <arious fragrant rice <arieties in Australia, India, 4hailand, and 8hinaG none of these ha<e 7een a$$lied to Indonesia rice <arieties. Bragrant genoty$ing was carried out on $o$ular Indonesia non-fragrant "8iherang, Bat0awati% and fragrant "#andan wangi, 6o.o Cele, 2enti/ 3angi, >unung #era/, #ulu 0andoti, #are 5e07ang, Sintanur% rice <arieties using <arious re$orted fragrance 0ar/ers "Brad7ury et al. 2 &7, Cang and Buu 2 !, Shi et al. 2 !, Sa/thi<el et al. 2 '%. Bor co0$arison, I6=4, Ni$$on7are and 4ai$ei 3 ' were included. 2! Materia"s and Methods 2.1 Plant materials and reagents 6ice sa0$les were su$$lied 7y the Indonesian 8enter for Agricultural Biotechnology and >enetic 6esources 6esearch and -e<elo$0ent as well as 7y the Indonesian 8enter for 6ice 6esearch. 4he Brad7ury 0ar/er was o7tained fro0 the 2idland 8ertified 6eagents 8o. 6NAse, #86 7uffer, fastart 4aF -NA $oly0erase and dN4# were su$$lied 7y 6oche -iagnostics Indiana$olis. )ne /7 standard -NA siDe 0ar/er and agarose low ?? were su$$lied 7y In<itrogen and A$$liche0 Bioche0ical 8he0ica Synthesis Ser<ice, res$ecti<ely. All other che0icals, of the highest $urity a<aila7le, were $urchased fro0 Sig0a. 2.2 DNA extraction >eno0ic -NA was eEtracted fro0 fresh young lea<es following the $rotocol descri7ed 7y -oyle and -oyle "+'' %. 2.3 PCR and electrophoresis 4he co0$osition of #86 0iE and cycling conditions, as well as the electro$horesis conditions were as descri7ed 7y the $re<ious authors "Brad7ury et al. 2 &7, Cang and Buu 2 !, Shi et al. 2 !, Sa/thi<el et al. 2 '%, eEce$t for the source of reagents which are descri7ed a7o<e. In addition, agarose gel was used for the B2-?, and B2-? 0ar/ers #86 $roducts, in $lace of the reco0ended $olyacryla0ide gel "Shi et al. 2 !%. #! Resu"ts and $iscussion 3.1 enot!ping "sing Bradb"r! mar#er Isolated -NA fro0 non-fragrant and fragrant rice sa0$les were #86 a0$lified using Brad7ury et al."2 &7% $ri0er, and analysed 7y agarose-gel-electro$horesis. 4he results "Bigure +% showed that only fragrant 2enti/ 3angi and >unung #era/ rice #86 $rofiles were as $re<iously re$orted "Brad7ury et al. 2 &7%, and these were different fro0 those of non-fragrant sa0$les "8iherang, Bat0awati, I6=4, Ni$on7are, 4ai$ei 3 =%. Howe<er, other frangrant "#andan wangi, 6o.o Cele, #ulu 0andoti, #are 5e07ang, Sintanur% were si0ilar. Based on these results, it was deduced that there were two grou$s of badh2-deletion $atterns a0ong Indonesia fragrant rice <arieties. >rou$ + includes 2enti/ 3angi and >unung $era/, while grou$ 2 includes #andan wangi, 6o.o Cele, #ulu 0andoti, #are 5e07ang, and Sintanur. Brad7ury 2ar/er has 7een $ro<ed to $ositi<ely a0$lify the badh2 gene of <arious rice <arieties "Brad7ury et al. 2 &7, A0arawathi et al. 2 !%. 4he eEistence of a &!& 7$ badh2 7and "Bigure l% indicates that the badh2 gene of all the eEa0ined Indonesia rice <ariety sa0$les were identical to those rice <arieties of other countries. 4he $resence of the fragrant 7and "2&, 7$% in grou$ l fragrant rice sa0$les "2enti/ 3angi and >unung #era/% indicated si0ilarity of the badh2.7 0utation $attern with those $re<iously re$orted fragrant rice sa0$les "Brad7ury et al. 2 &7, BorFuis et al. 2 !%. Howe<er, those of >rou$ 2 "#andan wangi, 6o.o Cele, #ulu 0andoti, #are 5e07ang, and Sintanur% which did not $ossess the fragrance 7and see0ed to ha<e a different badh2-0utation $attern. Si0ilar results ha<e also 7een re$orted $re<iously "A0arawathi et al. 2 !%. 4he badh2 0utation in grou$ 2 0ay 7e shorter than that of grou$ l "! 7$% as illustrated in Bigure 2. As a results, the internal fragrant $ri0er "IBA#% 7eco0es unsuita7le, and therefore no fragrance 7ands were o7ser<ed. )n the other hand, the non-fragrance internal $ri0er "INS#% 7eco0es suita7le, and therefore uneE$ected non-fragrance 7ands "3!& 7$% were o7ser<ed in the sa0$les of grou$ 2 fragrant rice sa0$les. 8onseFuently, the siDe of grou$ 2 badh2 gene should 7e larger than that of grou$ l fragrant rice <arieties.

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Journal of Biology, Agriculture and Healthcare ISSN 2224-32 ! "#a$er% ISSN 222&- '3( ")nline% *ol.3, No.+,, 2 +3

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Bigure +. Brad7ury-#86 $rofiles s J standart siDe 0ar/er, N J Ni$$on 7are, 8 J 8iherang, I6 J I6=4, B J Bat0awati, 4 J 4ai$ei 3 =, 2 J 2enti/ 3angi, > J >unung #era/, 6CJ 6o.o Cele, # J #andan 3angi, #2 J #ulu 2andoti, #5 J #are 5e07ang, Si J Sintanur, >i J >ilirang, and w J water "negati<e controll%. + and 2 indicate 0e07ers of grou$ + and 2, res$ecti<ely. 3.2 enot!ping "sing $%&'7 '7 or Badex mar#er 4he #86 $rofiles of non-fragrant fragrant and fragrant rice <arieties were further co0$ared using B2-?, B2 "Shi et al. 2 !% or BadeE,-& "Sa/thi<el et al. 2 '% 0ar/er. 0ar/er 4he results showed that 7oth B2-?, ?, "Bigure 3% and BadeE ,-& "Bigure 4% % 0ar/ers were a7le to distinguish distin 7etween non-frangrant and fragrant rice of grou$ +, 7ut not of grou$ 2. 4he #86 $rofile results using B2-?, "Shi et al. 2 !% "Bigure 3% or BadeE ,-& & "Sa/thi<el 2 '% "Bigure 4% 0ar/ers also su$$orted the <iew that there are 2 grou$s of eEon ,, badh2 deletion $atterns a0ong Indonesian Indonesia fragrant rice <arieties. 4he he grou$ 0e07ers are identical to those o7tained $re<iously $re<io sly using the Brad7ury et al. "2 &7% 0ar/er. 4he difference in 7and 0igration distance was w relati<ely s0all "Sa/thi<el et al. 2 '%. 4he use of $olyacryla0ide gel has 7een suggested to resol<e #86 a0$licons resulted using the Shi et al. "2 !% rnar/er. Howe<er, no further eE$eri0ents were w carried out since these 0ar/ers were una7le to discri0inate #andan 3angi and 6o.o Cele fro0 non-fragrant fragrant rice sa0$les. sa0$les

Bigure 2. Suggested 0utation-$attern 0utation differences 7etween grou$ + and 2-fragrant fragrant rice <arieties

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Journal of Biology, Agriculture and Healthcare ISSN 2224-32 ! "#a$er% ISSN 222&- '3( ")nline% *ol.3, No.+,, 2 +3

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Bigure 3. B2-?,-#86 $rofiles s J standard 0ar/er, N J Ni$$on 7are, 8 J 8iherang, I6 J I6=4, B J Bat0awati, 4 J 4ai$ei 3 =, 2 J 2enti/ 3angi, > J >unung #era/, 6CJ 6o.o Cele, # J #andan 3angi, #2 J #ulu 2andoti, #5 J #are 5e07ang, Si J Sintanur, >i J >ilirang, and w J water. + and 2 indicate 0e07ers of grou$ + and 2, res$ecti<ely.

Bigure 4. BadeE,-&-#86 $rofiles. s J standard 0ar/er, N J Ni$$on 7are, 8 J 8iherang, I6 J I6=4, B J Bat0awati, 4 J 4ai$ei 3 =, 2 J 2enti/ 3angi, > J >unung #era/, 6CJ 6o.o Cele, # J #andan 3angi, #2 J #ulu 2andoti, #5 J #are 5e07ang, Si J Sintanur, >i J >ilirang, and w J water. + and 2 indicate 0e07ers of grou$ + and 2, res$ecti<ely. 3.3 enot!ping "sing $%&'2A mar#er An earlier re$ort has found the badh2.2 0utation in fragrant rice "Shi et al. 2 !%. 4herefore, eE$eri0ents were done to co0$are non-fragrance and fragrance #86 $rofiles using the eEon 2 fragrance 0ar/er "B2-?2A%. Howe<er, the results "Bigure &% showed no differences in the #86 $rofiles of all sa0$les. 4he fact that no difference 7etween fragrant, as well as with those of non-fragrant rice sa0$les "Bigure &% in the #86 $rofiles was o7tained using the B2-?2A 0ar/er "Shi et al. 2 !% suggested that the $re<iously re$orted eEon 2-badh2 0utation is not a$$lica7le to 0ost of the eEa0ined Indonesian fragrant rice sa0$les.

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Journal of Biology, Agriculture and Healthcare ISSN 2224-32 ! "#a$er% ISSN 222&- '3( ")nline% *ol.3, No.+,, 2 +3

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Bigure &. B2-?2A-#86 $rofiles. s J standard siDe 0ar/er, N J Ni$$on 7are, 8 J 8iherang, I6 J I6=4, B J Bat0awati, 4 J 4ai$ei 3 =, 2 J 2enti/ 3angi, > J >unung #era/, 6CJ 6o.o Cele, # J #andan 3angi, #2 J #ulu 2andoti, #5 J #are 5e07ang, Si J Sintanur, >i J >ilirang, and w J water. + and 2 indicate 0e07ers of grou$ + and 2, res$ecti<ely. 3.( enot!ping "sing the R%223 mar#er In atte0$t to find suita7le fragrance 0ar/er for #andan 3angi or 6o.o Cele "grou$ 2%, their #86 $rofiles were further co0$ared with non-fragrant 8iherang and fragrant rice of grou$ + "2enti/ 3angi and >unung #era/% using the 62223 0ar/er of Cang and Buu "2 !% as $ri0er. 4he results "Bigure =% showed that 62223 was a7le to discri0inate #andan 3angi and 6o.o Cele fro0 non-fragrant 8iherang. 4he siDe of the grou$ 2 a0$licons was o7ser<ed to 7e s0aller than those of the grou$ + fragrant rice <ariety sa0$les. A$art fro0 the a7ility of the 62 223 SS6 0ar/er "Cang and Buu 2 !% to discri0inate fragrant fro0 nonfragrant rice sa0$les "Bigure =%, the results also suggest that there are differences 7etween grou$ + "2enti/ 3angi and >unung #era/% and grou$ 2 "#andan 3angi and 6o.o Cele% rice sa0$les, and that the siDe of grou$ +a0$licons were also found to 7e s0aller than those grou$ 2. 4his su$$orts the results o7tained $re<iously using the Brad7ury 0ar/er "Brad7ury et al 2 &7% in Bigure +.

Bigure &. 62233-#86 $rofiles. w J waterG > and 2J >unung #era/ and 2enti/ 3angi "∼+2 7$%G 6C and # J 6o.o Cele and #andan 3angi "∼+4 7$%G 8 J non-fragrant 8iherang "∼+= 7$%G and s J standard 0ar/er. + and 2 indicate 0e07ers of grou$ + and 2, res$ecti<ely. %! &onc"usion 4he results of fragrance genoty$ing using <arious fragrance "Brad7ury, B2-?,, BadeE ,-&, and 62 223% 0ar/ers su$$orted the <iew that there are at least 2 grou$s of eEon, badh2-0utation $atterns a0ong Indonesian fragrant rice <arieties. >rou$ + includes 0enti/ 3angi and >unung #era/G while grou$ 2 includes #andan wangi, 6o.o Cele, #ulu 0andoti, #are 5e07ang, and Sintanur. No badh2.2 0utation was found a0ong Indonesian rice

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Journal of Biology, Agriculture and Healthcare ISSN 2224-32 ! "#a$er% ISSN 222&- '3( ")nline% *ol.3, No.+,, 2 +3

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<arieties. Ac'now"edgments 4he authors than/ the Higher ?ducation -irectorate and Bogor Agricultural ;ni<ersity, 2inistry of ?ducation and 8ulture of the 6e$u7lic of Indonesia for funding the research wor/ "B)#4N 8ontract no@ +3,HI43.4+.2HC2HS#5H2 +3, +& 2ay 2 +3%G and C##2 I#B, B2I#A I#B, -e$art0ent of Bioche0istry B2I#A I#B, C4 I#B, BB Biogen, and CI#I for coo$eration, ad0inistration 0anage0ent, la7oratory, and hu0an resource su$$ortG as well as all research and technical assistance "-i0as Adrianto 2si, Ani<ersari A$riana 2Si, Joel 6i<andi Sinaga SSi, ?uis 2arlina SSi, 6udy 2unDirwan SSi, Sugihartati SSi, 4aufiF SSi, -ewi #ra$tiwi, SSi, Ba07ang #ad0adi SSi, 2uha00ad 4aufan Bataha.udin, Hel0y 6a0adhan Al Anshary% for coo$eration and hard wor/ during this research. References Ahn, S.N., Bollisch, 8.N., K 4an/sley S.-. "+''2%. 6BC# tagging of a gene for aro0a in rice. )heor Appl enet* !4@!2&L!2!. doi@+ .+ ,HBB 22,3'+. A0arawathi, I., Singh, 6., Singh, A.5., Singh, *.#., 2oha$atra, 4., Shar0a, 4.6., K Singh N.5. "2 !%. 2a$$ing of Fuantitati<e traitloci for 7as0ati Fuality traits in rice ")ryDa sati<a C.%. %ol Breed* 2+@4'L=&. doi@+ .+ ,Hs++ 32- ,-'+ !-! Bourgis, B.6., >uyot, H., >her7i, ?., 4aillieD, I., A0a7ile, J., Salse, 2., CorieuE, 2., -elseny, K >hesFuiMre A. "2 !%. 8haracteriDation of the 0a.or fragrance gene fro0 an aro0atic +aponica rice and analysis of its di<ersity in Asian culti<ated rice. )heor Appl enet* ++,"3%@ 3&3L3=!. Brad7ury, C.2., Bitgerald, 4.C., Henry, 6.J., Jin, N., K 3aters -.C.?. "2 &a%. 4he gene for fragrance in rice. Plant Biotech ,.* 3@3=3L3, . Brad7ury, C.2., Bitgerald, 4.C., Henry, 6.J., Jin, N., K 3aters -.C.?. "2 &7%. A $erfect 0ar/er for fragrance genoty$ing in rice. %ol Breed* +=@2,'L2!3. 8ordeiro, >.2., 8hristo$her, 2.J., Henry, 6.J., K 6ein/e 6.B. "2 2%. Identification of 0icrosatellite 0ar/ers for fragrance in rice 7y analysis of the rice geno0e seFuence. %ol Breed* '@ 24&L2& . -ong, J.I., SuDu/i, ?.4., K 4erao H. "2 %. Inheritance of aro0a in four rice culti<ars "-r!.a sati/a C.%. I66I. International 6ice -oyle, J. J., and -oyle, J.C. "+'' %. A ra$id total -NA $re$aration $rocedure for fresh $lant tissue. $oc"s* +2@+3-+&. BitDgerald, 2.A., Ha0ilton, N.6.S., 8alingacion, 2.N., *erhoe<en, H.A., Butardo, *.2. "2 !%. Is there a second fragrance gene in rice. Plant Biotech. ,.* =@4+=-423. Ha0i Seno, -.S., Santoso, 4.J., 2as:ud, 9.A. "2 + %. Brad7ury 0ar/er-assisted introgression of 2enti/ 3angi0utated badh2 gene. Proc Ann Res 0PB 2 + , ,,&-,!&. ISBN @ ',!-= 2-!!&3- 3-3, ',!-= 2-!!&3-+3-2. Ha0i Seno, -.S., #ad0adi, B., #ra$tiwi, -., Sugihartati, 4aufiF, Bataha.udin, 2.4., Al Anshary, H.6., Santoso, 4.J., K 2as:ud, 9.A. "2 ++%. 2enti/ 3angi-0utated badh2 gene transfer into non-fragrant rice c.<. 8iherang. ,0P0* <ol. +=, + @ =&-, . ISSN @ !&3-42+,. 5o<ach, 2.J., 8alingacion, 2.N., BitDgerald, 2.A., K 2c8ouch, S.6. "2 '%. 4he orign and e<olution of fragrance in rice "-ri.a sati/a 1.%. #NAS, + =@+4444-+444'. Cang, N.4., K Bu, B.8. "2 !%. -e<elo$0ent of $cr-7ased 0ar/ers for aro0a "2gr3 gene in rice "-r!.a sati/a0 C.%. -monrice* +=@ +=-23 CorieuE, 2., #etro<, 2., Huang, N., >uiderdoni, ?., K >hesFuiMre, A. "+''=%. Aro0a in rice@ genetic analysis of a Fuantitati<e trait. )heor App enet* '3@++4&L++&+. #aule, 8.2., K #owers, J.J. "+'!'%. Sensory and che0ical eEa0ination of aro0atic and non aro0atic rices. , $ood 4ci* &4@343L34=. #etro<, 2., -anDart, 2., >ia0$aoli, #., Baure, J., K 6ichard, H. "+''=%. 6ice aro0a analysis -iscri0ination 7etween a scented and a non scented rice. 4ci Aliments* +=@34,L3= . 6ein/e, 6.B., 3elsh, C.A., 6eece, J.?., Cewin, C.>., K Bla/eney, A.B. "+''+%. #rocedures for Fuality selection of aro0atic rice <arieties. 0nt. Rice Res. Ne5slett* +=@ + L++. Sa/thi<el, 5., 6ani, N.S., #andey, 2.5., Si<aran.ani, A.5.#., Neera.a, 8.N., Balachandran, S.2., 2adha<, 2.S., *ira/ta0ath, B.8., #rasad, S.*., K Sundara0, 6.2. "2 '%. -e<elo$0ent of a si0$le functional 0ar/er for fragrance in rice and its <alidation in Indian Bas0ati and non-Bas0ati fragrant rice <arieties. %ol. Breeding* -)I + .+ ,Hs++ 32- '-'2!3-E Shi, 3., Iang, I., 8hen, S., K (u, 2. "2 !%. -isco<ery of a new fragrance allele and the de<elo$0ent of functional 0ar/ers for the 7reeding of fragrant rice <arieties. %ol Breed* 22@ +!&-+'2. Sood, B.8., SidiF, ?.A. "+',!%. A ra$id techniFue for scent deter0ination in rice. 0ndian ,. enetic Plant Breed* 3!@ 2=!L2,+.

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Sri<ong, #., 3angso0nu/, #., K #ongdontri, #. "2 !%. 8haracteriDation of a fragrant gene and enDy0atic acti<ity of 7etaine aldehyde dehydrogenase in aro0atic and nonaro0atic thai rice culti<ars. 667 4ci ,* 3="4%@ 2' -3 +. 4anchoti/ul, ;., and Hsieh, 4.8.I. "+''+%. An i0$ro<ed 0ethod for Fuantification of 2-acetyl-+-$yrroline, a O$o$cornP-li/e aro0a, in aro0atic rice 7y high-resolution gas chro0atogra$hyH0ass s$ectro$hoto0etryHselecti<e ion 0onitoring. , Agric $ood Chem* 3'@ '44-'4,. 3id.a.a, 6., 8ras/e, J.-., K 3ootton, 2, "+''=%, 8o0$arati<e studies on <olatile co0$onents of non-fragrant and fragrant rices. , 4ci $ood Agric* , @ +&+L+=+. Ioshihashi, 4., Huong, N.4.4., K Inato0i, H. "2 2%. #recursors of 2-acetyl-+-$yrroline, a $otent fla<our co0$ound of an aro0atic rice <ariety. , Agric $ood Chem* & @2 +L2 4.

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