DI TRUYEN - GIONG VAT NUOI
DA D A N G NUCLEOTIDE TREN \ T J N G D-LOOP T^ THE CUA MOT SO Q U A N THE TR.\U BAN DIA \ I E T NAM
\guven Sgoc TdiP; Nguyen Phgni Kmi Vy.in Ho,hig Tudn Tlidnli^.
Phgm Cong Viicu'i.l Sguyin Cong DiiiiP Ngay n h a n bai bao: 27/06/2019 - \ i : a y nhgn bai p h a n bipn: l^/iC 201^
Ngay bai bao d u u i c h a p n h a n d a n g 2S/0S/2li^' TOM T,\T
Myc tieu ciia nghien cuu nay nhSm buvK dau khao ^.M da ttong di truven nueleotidc thupc trinh t y viing D-Ump ty the cua hai quan the trau ban dia Viil Nam (L.inj;tM.inj; - Lam Dong va Bao 'i i n • Lao Cai) va trau Murrah (nh^p ve tu An Op). C i p moi ilupi Uuvl ko dv khuyccli dgi dogn gen myc tieu cd kich thudc khoang 760 bp, troiii: J o co ^(i5 dcn 572 bp viinj; D-ioop. Ket qua cho tha'v tmng binh cic lo^i nucleotide- Thymine (I i = 27,22".., C\iosiiiv (C) = 2?,(\1".., Adenine (A) = 33,02%, Guanine (G) = 14,13% \ i chi so' da dqng nucleotide (p) la 0,07. Cd 100 vj tri bien dSi da hinh nucleotide va 10 haplotypc da dupc quan sal voi chi so da dgng haplotype (Hd) la 0,9697. Co 3 bicn doi chung \ . n ra o tat ca cac ca the trau, c6 1 vj Iri bioh doi chung chi cd a trau Murrah va nhicu d u m bii'h Joi \ay ra o quan lhi- irau npi. Khoang each di Iniycn giira trau Murrah va trau Bao Yen la Ion nhai tronj; khi khacbii'l v e d i truven nhd nhat dupc tim thay giiia hai quan the Bao VOn \ .1 Langbiang. Ket qua cav di tru> vn da chiing mmh nguon goc him hyp cua quan the trau Bao Yen (Lao Cai) vi 1 .in[;bi.inj; (l-im Dong), khi haj quan the ph.il Irivn thanh nhanh neng va khac biC't so vni gidng trau Murr.ih. Gio lang d u n g lu^mg mau dc phan tich la dieu can thiet cho nghien cuu ticp theo.
Tir khoa: Tnin, ^ic gen ty the, vung D-loop, da dang dt truyen.
ABSTRACT
Genetic diversity of Vietnamese native buffalo populations based on D-loop sequence The aim nl this study was to investigate Ihe gc-netic diversity of Vietnamese native buffalo (Bubalus Inibalis] in two regions of Vietnam on mitochondrial D-loop region by nucleotide .sequencing, Tho primers was designed to amplified the Lirgol geno with 760 b p in size, in which contained the fragment of 5(1? to 572 b p of tho D-Ioop region. The sequence from 12 buffalo samples woa' used to analyze tho nucleotide diversity, genetic distance and reconstruct the phylogenetic treo. The results showed that the average number of nucleotide was Thymine (27.22%), Cytosine (25.63%), Adenine (33 02%), Guanine (14.13%) and the nucleotide diversity (p) was 0.07. There were too polymorphic sites and 10 haplotj'pes wen? found and the haplotype diversity (Hd) was 0,9697.
Three pt^lyniorphic silos occurred in ail buffalo individual and only one polymorphic site was obser\'ed in Nhirrah buffalo. There were several polymorphic sites were observed in Vietnamese native buftaloes The genetic di>lancc value t>elwoon Murrah and Bao Yen buffaloes \\ as Iho hiehest while this value was lowest between Bao Yen and Langbiang bufallo populations The results of phylogenetic tree demonstrated that the mixed origin between Bao Yen and Langbiane buffalo which were grouped in one d a d e then divided into sub-populalion.s, while the Murrah buffalo was separated in other clade- Thus, increasing sample size for further <nnaK sis is required.
Keywords: Bubatus bubalts, mtlochotirtal 0.V.4 D-loop, genetic diversity.
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• T.ic gia liOn W TS Ni^uyvn Nfi>v Tan. Ciang vien dunh B9 mon Cong nghe Smh hi^ - Dai hoc Nong lam Tp, i I,', C hi Mmh t">T i^H.s"-'""- •Vs Em.iil nnian'hcniuafedu.\-n
KHKT Chdn nudi UJ 254 • thdng 2 nam 2020
TNUOI
I.BATVANB^
Con trau thuc su dong vai tro c^uan trpng trong san xuat nong nghipp va gan bo mat thie't vdi ngudi ndng dan, la bieu tupng van hda Clia ngudi Viet Nam ndi rieng, van hda Ilia nudc ndi chung. Cung gid'ng nhu nhieu noi tren the gidi, khi tang muc dp co gidi hda canh tac ndng nghiep dan deh suy giam so lupng quan the trau (Nguyen Ngpc Tan va ctv, 2017). Khuynh hudng chung ve chan nudi trau khu vuc Ddng Nam A theo xu hudng giam (Yusuf va ctv, 2013). Tru'dc dd, Nanda va Nakao (2003) cdn cho rang nguyen nhan giam dan trau khu vuc Chau A la chan nuoi trau khdng nhan dupc su quan tam cua cac nha hoach dinh chinh sach ciing nhu cac nha nghien ciiii mpt each tuong xung vdi Hem nang cua nd. Trong thdi gian qua, da cd mpt so nghien cuu tim giai phap de tac dpng nham tao dieu kipn phtic hoi sp phat trien cua nghe nudi trail {Mai Van Sanh va ctv, 2008; Dinh Van Cai va ctv, 2012; Nguyen Ngpc Tan va Nguyen Thanh Thuc, 2018) nhung chua co nliieu ngliien ciiii ve linh vpc di truyen phan tir. Cac nghien cmi trudc day ve da dang di truyen cac gid'ng trau nudc chau A dua tren tr'mh tiv vimg gene D-loop ty the da cung cap cai nh'm td'ng quan ve nguon goc phat sinh va chieu hudng thuan hda ciia loai nay d khu vv'c (Lau va ctv, 1998). O Viet Nam, cac nghien CUTJ di truyen dua vao vimg D-loop ty the da dupc tie'n hanli tren mot so'doi tu'png nhu chd Phu Qudc (Tran Hoang Dung va ctv, 2016), ga (Ngo Thi Kim C u c v a ctv, 2011), de ban dia Ninh Thuan (Nguyen Ngpc Tan va ctv, 2018), nhung chua cd nghien CUTJ sir dung vimg D-loop ve quan the trau Viet Nam, ngoai trif sir dpng chi thi microsatelUte (Nguyen Ngpc Tan va ctv, 2019)- Vi the^ muc tieu ciia nghien cu\i nay la nhan bie't su khac biet di truyen giiia hai quan the trau ban dia Viet Nam la Bao Yen va Langbiang vdi gid'ng trau Murrah nhap tir An Dp di^a vao vimg D-loop gen ty the lam co sd dii lieu ban da'u cho viec nghien cuTi ngudn ^oc va quan hp di truyen trau ban dia Vipt Nam theo ddng mp.
2. VAT UEU VA PHUONG PHAP NGHIEN ClKU
2.1. Vat lipu, dia diem va thdi gian Mdu mo: Mau md tai trau ban dia Viet Nam duoc thu thap tai mpt so dia phuong:
Lang Biang (Lam Dong), Bao 'Yen (Lao Cai) va dupc cung ca'p bdi Bd mdn Di truyen Gid'ng Vat nuoi - Phan Vien Chan nuoi Nam Bp. Mau md tai trau Murrah (nhap td An Dp vao Viet Nam vao nhung nam 80 the ky XX) dirpc thu nhan tai Trung tam Nghien ciiii va Phat trien Chan nuoi Gia siic ldn (Lai Hung, Ben Cat, Binh Duong). Mau dupc ki hipu: trau Bao Yen (BY1-BY5), Langbiang (LB1-LB5) va Murrah (M1-M5).
Hda cha't: Phan irng khue'ch dai FCR duoc thuc hien bSng hoa chat la bp kit MyTaqTM Mix - Bioline. Hoa cha't dipn di bao gom Agarose 1,5% (Bioline), GelRed 0,6X (TBR), ladder lOObp (Thermo Scientific - My), dung dich dpm TBE 0,5X (Viet Nam).
Dia diem vd thdi gian: Bp mon Cpng nghe Sinh hpc - Trudng Dai hpc N6ng Lam Tp. Ho Chi Minh, tii thang 11/2018 de'n thang 6/2019.
2.3. Phuong phap
Tliiei ke" mdi: Cap mdi T760 dupc thie't ke' bang phan mem PrimerS dua tren mach khuon chinh co ma so truy cap AY488491.1.
Trinli hJ> (5'-3') moi xudi AATACCAACG- GCCAGCATAA va mdi ngupc GAGCATGG- GCTGATTAGACA, mdi xuoi bat dau tir vimg gen cytochrome B va moi ngupc nam tai vung bao ton tren D-loop cua mtDNA.
Khue'ch dgi dogn gen bdng PCR: Kich thudc san pham khue'ch dai la 760bp, trong do 576bp thupc vung D-loop, Phan ung PCR (25\il) chua cac thanh phan: 12,5pl MyTaqTM Mix, l,25pl moi primer, 2,5pl DNA khudn mau va 25pl H20. Chu trinh nhiet dupc thuc hien theo cac budc; 1) 94''C trong 3 phut; 2) 94''C trong 45 giay; 3) 55°C trong 30 giay; 4) 72°C trong 45 giay; 5) lap lai 35 chu ky tir budc 2 de'n 4; 6) 72''C trong 5 phut va 7) giir nhiet dp 4"C trong 10 phut bang may MasterCycler Pro S (Eppendorf, Diic). Cac san pham khue'ch dai duoc dien di tren gel agarose 1% (30 phiit,
KHKT Chdn nudi so 254 - thane 2 ndm 2020
90V), q u a n sat va c h y p h i n h a n h dipn di b a n g m a v G e l D o c lt2 (UVP, USA) vdi t h a n g c h u a n lOObp.
2.4. X u ly sd l i ? u
So s a n h t r i n h t y cua v i i n g D-Ioop ty t h e thi/c hipn b a n g p h a n m e m M E G A \ va BioE- d i t d e p h a n tich trinh t u n u l e o t i d e . S u d y n g p h a n m e m M L C A \ d e d a n h gia k h o a n g each di t r u y e n vdi m o h i n h T a m u r a - Xei, p h u c m g p h a p M a x i m u m likelihood p h a n h'ch boot- s t r a p 1.000 (liip l^i 1.000 Ian) d u p c s u d y n g d e
\ a y d u n g cay p h a t sinh loai. Dir lipu \ e cac ca the trail T r u n g Q u o c , t r a u .An D p d u p c thu I h a p tir n g a n h a n g g e n va d u p c s u d y n g la n g u i m t h a m k h a o .
3. K^T QUA VA THAO LUAN
3.1, K e t q u a k h u y e c h dai d o ^ n g e n m u c tieu CO k i c h t h u d c 7 6 0 b p
P h a n irng PCR k h u e c h d^i v i m g g e n ki'ch t h u d c 7 6 0 b p tren v i m g D-loop cho 15 m a u ca t h e cua 3 q u a n lhi- trau, ket q u a dipn di san p h a m PCR d u p c trinh b a y d H i n h 1.
Til ket qua o H i n h 1 cho tha'y chi xua't hipn 01 band san p h a m o m d i gieng, ro net vdi kich t h u d c phii h p p vdi kich t h u d c m o n g d p i (760bp) d a d u a ra ve ly t h u y e t khi thiet ke c^p mdi. Sau d o , san p h a m PCR d u p c gdi giai trinh t y cho cac b u o c p h a n tich ki' tie'p.
H i n h 1. San p h a m PCR tren gel agarose l^S-o LD lOObp, BY, LB, M 3.2. N h u n g b i e n d o i t r o n g t r i n h tir v i m g D - l o o p
Til 15 m a u s a n p h a m PCR d u p c giai trinh t y , t r o n g d o co 3 m a u bi ldi kv t h u a t (BY2, LB3.
LB5) d a d u p c loai b o . S u d u n g ket q u a giai t r i n h t u ciia 12 m a u ca t h e vdi kich t h u d c s a n p h a m k h o a n g 565-572bp t h u p c v i m g D-loop cho p h a n tich d a h i n h n u c l e o t i d e va d a n h gia d a d ^ n g di t r u y e n . Ket q u a cho tha'y: T r u n g
b'lnh cac loai n u c l e o t i d e la T h v n i m c (T) 27,22'\.; C y t o s i n e (C) = 25.63"..; A d e n i n e (A' = 33,02",., G u a n i n e (G) = 14,13'V. Cac nucleotide Io?i A va T c h i e m h ^ i n g b i n h 60,24'\. phii hpp v d i cac ly t h u y e t t r u d c d a \ ve t h a n h p h a n .A-T t r o n g v i m g D - l o o p t\' the. Chi so da d o n g n u c l e o h d e la p = 0,07. \ g h i e n cuxi cua Lau ya ct\- (1998) khi p h a n tich 25 c.i the trau d a m lav va 10 ca t h e t r a u >dng cho th.iy da d a n g n u c l e o t i d e h r o n g u n g 0.056) v.i 0,():il3, t r u n g b i n h c h u n g la 0,0526. T r o n g khi do. Lei v a ctv (2007) klii n g h i e n ciru tren trau b a n dia T r u n g Qud'c cho thay chi sd d a d g n g n u l e o t i d e la 0,L)06.S-
C a c vj tri da h i n h n u c l e o t i d e ciia 12 t r i n h t y d u p c chi ra d Bnng 2 va t o n g cpng 100 vj tri da h i n h n u c l e o t i d e d u p c p h a t hipn, T r o n g dd;
cd 1 vj tri da h i n h chi xua't hipn d cac ca t h e t r a u M u r r a h (401, C e T), n h d m t r a u npi (Bao Yen va L a n g b i a n g ) co 28 vi tri bie'n d o i c h u n g (5, 16, 47, 94, 132, 138, 139, 146, 147, 150, 162, 182, 184, 239, 256, 299, 300, 326, 338, 340, 344, 354, 416, 450, 465, 520, 528, 534) va co 3 vi tri d a h i n h xua't hipn d ta't ca cac ca t h e t r a u (273, 419, 449). N g o a i ra, xua't h i p n n h i e u vi tri d a h'mh d a c bipt chi xay ra d ca t h e n a y ( B a n g 2),
C o 34 vi tri c h u y e n d d i ( t r a n s i t i o n ) (T«C) va 37 vi tri c h u y e n d d i {A«G), t r o n g k h i chi cd 16 \'i tri c h u y e n vi ( t r a n s v e r s i o n ) , ti le c h u y e n d d i ; c h u y e n vi la 4,4:1. Ket q u a t r e n cd n h i e u n e t tu-ong d o n g v d i n g h i e n CIJU c u a H a s s a n va ct\' (2009) khi da q u a n sat d u p c 27 c h u y e n d o i (T<'C), 16 c h u y e n doi (A'.G), t r o n g khi chi co 9 c h u y e n vi tren 77 vi tri d a h i n h v i i n g D - l o o p cua trau. Ty le c h u y e n d o i : c h u y e n vi d n g h i e n ciiu tren la 4,8:1, C d t h e tha'y d u p c cac d o t bien c h u y e n doi co xu h u d n g xay ra t h u d n g x u y e n va p h o b i e n h o n s u c h u y e n vi ciia cac n u c l e o t i d e .
C a c vi tri da h i n h co t h e xay ra o riens n h d m t r a u npi, d rieng h-au M u r r a h h o a c o ta't ca cac ca t h e t r a u d u p c thi n g h i e m , Trung c u i v m p t g i d n g t r a u ciing xay ra cac vi tri buVp din k h a c n h a u , t h a m chi cd ca the d u p c q u a n •,dt cd ca n h u n g b i e n di rieng bipt.
KHKTChdn nudi so 254 - tbdng 2 nam 2021)
Ul I K U Y t N - G I O N G vAT N U O I
Bang 2. Vi tri da hinh nucleotide trong viing D-loop dirac khuech dai
VITRiDAHTNH ~ MAU 5 U 16 34 47 68 94 114 118132138139146 147 150 161162164168177162183184185186
C A G C A G C G G T C T T C T A C A C C T BV3 C A G C C C A C C T G G G G G
T G A T A A C C A C C T
9 190 192199 207210 239 C T C T C C
G G G G G C C C
244 247 256 262 264 265 268 269 271 273 274 284 2S6 293 299 300 301 305 SOS 319 320 321 326 338 340 342 343 344 345 349 354 361 362
BYJ CAA
T G G A T A G G C A T TTT AT CGT T
T C C C A T C G G A T C G G A
TGCGC C G C
T GCC CCC CC TGCGC
366 369 370 375 377 382 383 393 396 401 415 416 417 419 423 434 447 448 449 450 453 461 465 467 481 484 520 523 528 534 546 548 565 CG TG GA TG TCG GC TGG TTA TC CT A G G T T G C G
AY, Irinh tv dimg l i m mfldi kliuon (AY488491 1),
Them vao dd, bang viec nhdm cac da hinh dan nucleotide ciing cho phep quan sat haplotype rieng biet d cac mau (Bang 3)-
- Rang 3. Phan tirh haplnfypp haplntype
Ten ca the the'
Hap 1Hap 2 Hap 3 Hap 4 Hap 5 Hap 6 Hap 7 Hap 8 Hap 9 H:.p 111
BYI BY3 BY4 BV5 LBI LB2 LB4 M],M2
M3 M4 U S
Chi so da dang haplotype (Hd)
Bao Yen (BY), Langbiang (LB), Murrah (M)
Co tdng cpng 10 haplotype dugc nhan bie't tren 12 ca the thi nghiem. Cac ca the d quan the trau npi khong chia se chung mpt haplotype nao vi vay quan sat duac 7 haplotj'pe. O quan the trau Murrah, ca the Ml va M3 chia se chung 1 haplotype, ca the M4,va M5 co haploU'pe gio'ng nhau- Nhu vay, cd 3 haplotype hien dien tren cac ca the trau Murrah-
Lau va ctv (1998) khi phan h'ch dua vao vimg D-loop ty the tren 80 ca the trau dam lay va trau song vdi viing trinh tu chi 158 nucleotide da phat hien 55 vi tri da hinh va
dm 2020
L/i in.\j I O -
tao nen 33 D-Ioop haplot\'pe- Da so cac vi tri bie'n ddi cung -vav ra tren nhdm trau dam la\, tuong tu nhu ket qua nghien ciiu ciia chung toi. Ben canh dd, nghien cuu ciia Wang va ct\- (2017) aing chi ra rang, cd 2 haplogroup tren nhdm trau sdng la Rl va R2 va 5 haplogroup rieng bipt hinh thanh tren nhom trau dam lay lan lupt la SA, SB, SC, SD, SE. Vi so lupng mau phan tich o nghien ciiu nay it nen chua nhan dipn dupc cac ca the theo nhdm haplogroup phil hpp, vipc gia tang lupng mau phan tich d nghien ciiu tiep theo la dieu can thiet.
3.3. Khoang each va moi quan h? di truyen Khoang each di truyen giiia quan the trail Bao Yen va trau Langbiang la nhu nhat (d=0,0481±0,0061), dieu nav cho th.Vy mdi quan hp di truyen giiia hai nhom trau na\
la gan nha't trong ba nhdm trau. Trong khi dd, khac bipt vi* di truyen Idn nha't tim tha'y giira qua'n the trau Bao Yen va trau Murrah (d=O,1038±O,O129), va giua hai quan the trau Murrah va trau Langbiang la 0,0886±0,0122.
Mn tr^n khoang each di truyen dupc the hipn d Bang 4.
Bang 4. Khoang each di truyen giiia cac quan the trau
Trau Bao Yen langbiang Murrah Bao Yon 0 ±0,0061 ±0,0129 Langbiang 0,0481 0 +0,0122 Murrah 0,1038 0,0886 0
Pban dir lieu dudi dirdng cheo Id khodng cacb dl truyen, tren dirdng cheo Id chenh lich ciia khodng cach di truyen
Cay quan hp di truyen dupc xay dung dua tren mo h'mh Tamura - Nei bang phuong phap Maximum Likelihood vdi gia tri bootstrap lap lai 1.000 tan. DO- lipu ve cac ca the trau Trung Qudc va trau An Dp dupc thu thap hi ngan hang gene va dupc su dung la nguon tham khao. Dp lin civ cua cav dupc \ac dinh bang
\ ipc su dung nhom ngoai la trinh ty bo on ddi (^Bos TiiuriL-:) Su phan hoa di truyen giiia cac gidng trau dupc tho hicn o Hinh 5
Hinh 5. Cay quan h? di truyen giira cac quan the trau
' Sda cdc nhdnh Id gta tri bootstrap, tbang ti le bieu lhi klioang cacb di truyen
Dya vao cay quan hp di tmyen cd the thay dupc cdc ca the trau chia lam hai nhdm rieng bipt la nhom trau ban dia Vi^t Nam va trau Murrah. Trau Murrah thupc nhom trau song, quan the trau spng tach ra dpc lap vi^chung dupc thuan hda dpc lap so vdi trau dam lay.
Trau Murrah d Viet Nam cd nguon goc tix An Dp va vSn giu' dupc cac dac diem da dang ciia quan the trau song nay Cac ca the (M4, M5) va (Ml, M2) Cling chia se chung mpt haplotype nen sy khac bipt ve di truyen khong dang ke.
Khi phan tich 30 ca the thupc 6 quan the trau ban dia Trung Qud'c cho tha'y cd hai dong me (A va B) dupc phat hien, dong A chie'm un the ciia trau ban dja cdn ddng B duoc cho la nguon gen thupc vimg Dong Nam A (Lei va ctv, 2007), Ben canh d d Kierstein va ctv (2004) da chung minh trau sdng (river buffalo) va trau dam lay (swamp buffalo) co cung ngudn go'c thuan hoa tir trau nudc hoang da.
O quan the trau npi, cd su tach thanh hai nhdm phu the hipn tinh hon hpp cua hai quan the do co su dan xen cua mpt so' ca the giua hai nhom phu. Nguyen nhan cu the la do nguon gd'c hdn hpp giira hai quan the trau nay va ljch sir qua trinh di cu nguon gen cua chiing. Dieu nay can dupc lam rd hon d nghien ciiu tiep theo.
U. KET LUAN
La nghien ciiu dau tien sir dung trinh ty nucleohde tren vung D-loop cho hai quan the
KHKT Chdn nuoi .sd 254 - thdng 2 nam 202(j
V.:.T NUOI
trau npi cua Vipt Nam (trau Bao Yen va trau Langbiang) cho tha'y chiing ngudn gd'c gan giJi va cd khoang each di truyen kha ldn so vdi trau Murrah. Cac ca the thupc nhom trau Bao Yen va Langbiang cd su khac biet dang ke ve mat di truyen khi chung cd cac haplotype rieng bipt. Cay quan hp di truyen cho tha'y hai gio'ng trau ban dia cua Vipt Nam co ngudn gd'c phat sinh chung, sau do hinh thanh hai nhanh phy va khac bipt so vdi gid'ng trau Murrah.
LCII CAM 0 N
Ngudn vgt lieu di tniyen cua trdu Langbiang vd Bdo Yen dirge cung ca'p bdi Bg mon Di truyen Giong Vat nuoi - Phdn vien Chdn nuoi Nam Bg, Ihugc cbuong trinh bdo ton Quy gen 2018.
TAI LI|U THAM KHAO
1, D i n h V a n Cai, Lini C d n g H o a , D a u V a n H a l , N g u y e n H i h i Tra, H o a n g K h a c H a i , N g u y e n T h i T h u H o n g , N g u y i n D u e C h u y e n , H a n Q u o c V u o n g , P h a n V a n K i e m , Lini C o n g H o a , T h a i K h a c T h a n h va T o n g V a n G i a p (2012), N g h i e n OJTJ cac bien p h a p ky t h u a t n a n g kha n 3 n g sinh sim va s i n x u a t trau. Bao cao t o n g ke't de till c a p Bp (2009-2012), Vi?n K h o a hg.c Ky t h u a t N o n g n g h i e p m i e n N a m .
2 N g o T h i K i m Cue, S i m i a n e r H., G r o e n e v e l d L.F. a n d W e i g e n d S. (2011), Multiple Maternal Lineages of Vietnamese Local Chickens Inferred by M i t o c h o n d r i a l D N A D-loop S e q u e n c e s . A s i a n - A u s t J, A n i m . Sci,, 24(2): 155-6L
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P.S. a n d R o u s h d y K h . (2009). M i t o c h o n d r i a l D-loop nucleotide s e q u e n c e s of Egyptian n v e r buffalo V a n a h o n .ind p h y l o g e n y s t u d i e s . Liv- S d , 125(1)' 37-42 5 K i e r s t e i n G., Vallinoto M., Silva A., S c h n e i d e r
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