I T R U Y E N - G I O N . \ T N L J » - ^ '
DAC DIEM DA HINH VUNG D-LOOP CUA GIONC; CHO DANG SOI U'NG DUNG CHO CONG TAC CHON ^'^^^^^^-..i^
Pimm VuinhHdi'', Bid Xudn PlurangK Trdn Hiru CdP. ^ ^'\^!""%,„,i]P Hd Vli Loan-vil Sgi'V^'"^""
N g a y n h a n bai bao: 01/03/2019 - Ng.i>' nhgn bai p h a n bipn: 30/03/201'^
Ngay bai b a o d u p c c h a p nhgn d a n g : 1 4 W ' 2 0 I ' ' TOMTAT
Nghien cuu dupc thpc hi?n tren 300 m i u mo Kr nhimg ca the cho dgng soi, bao gom ^^^
va 160 con cai. Ke't qua nghien cuu d i o tha\ vimg D-loop ciia giong cho dgng soi ai tinh a ^ ^'^..
voi 36 vj tri bien doi nucleohde trong trinh ty ciia cac man giong dpc va 30 a i a giong *^^';^ " ' i j - ^ ^ ^ p da s o l a bien doi dong hoan T-C hoSc G-.-\. rieng chi co mpt v.ii bion doi di hoan G-C (HapLi , F D13); G-T (Hap Dl 1), A-C (HapD28). Khoang each di hoiyon giir.i cac haplot)'pe cua cac ""^^ " T ^ I - ' cuu la 0,0-33%. Trong 59 Hap dupc tgo thanh khong co Hap nao mang bien dj don le cua mpt ca -^
Khoang each di tmyen giua HapD20 vdi 12 ca the d p c va HapC3, HapClS \ .Vi 24 aicc.i the cai la c o nha't (3,3%). Ke't qua nay chiing to giong ch6 dang soi nay co linh da dgng di tmyen rat cao, tu' d o tgo dieu kien thuan Ipi cho viec chpn lpc, phal tnen va nhan giong dgng cho nay.
Tir khoa: Cho bdn dia Viet Nam, cbo dgng .--oi. vimg D-loop.
ABSTRACT
Characlcristics polymorphic D-loop area of woIf-Iike dog breed applications for variety selection
Tho study wa.s carried out on 300 tissue samples from wolf-type individuals, including 140 males and 160 females. The ro-^uHs show that the high-polygon wolf-shaped D-loop region has 36 nucleotide modification positions in the sequence of male models and 30 nucleotide conversion po- sitions in the sequence of female models. The majority of changes are T-C or G-A conversion, only a few G-C transfiguraHon changes (HapD22, Hap D13); G-T (Hap D l l ) ; A-C (HapD28), Genetic distance between haplotypes of study samples ranged from 0.0% to 3,3%. In 59 H a p is formed there is no H a p that carries a single vanahon of an individual. Genetic distance between HapD20 and 12 males and HapC3, H a p C l S with 24 females is the highest (3 3%). The results show that this breed has a verj' high genetic diversity, thereby fadiitating the seleclion, development and breed this d o g .
Keywords: Dags Vietnamese indtgenotis dog, u>otf-like dog, D-loop region.
I.SATVAND^ ga'p 5-10 lan so vdi cac gen nhan (Brown va ctv, 1979). Sp nghiem ngat cua viee thira ke'dong Trong vai Ihap ky qua, Hnh da hinh eiia
gen me aia mtDNA (Kaneda va ctv, 1995) va DNA t,- the (mtDNA) da dupc ap dung thiinh j „ ^ . ^ ' , . ^ ^ ^ . ^ ^ j . ; j - ^„p^ j ; ,^„y-„ ^ ^^^g cong trong vice xac dinh nguon goc ben hoa, ,,„„ ,,§; (^^^^ ^^^^ ^ ^ ^ n g lao ra d,, hinh nhan dang, moi quan he di tniyen giiia cac mtDNA, dac biet la su bien doi nudeoHde, quan the ding loai \ a giira cac loai co Uen quan rnot cong cu hiru ich de tie't 16 mirc do tieh hoa chat che voi nhau (Sacks va rtv, 2013; Freedman phan hr trong cac dong doi ciia cac luai dpng va ch', 2014: Wang va ctv, 2015; Caims va cl\', vat (SindiHc va ctv, 2011).
2017;Marinovvactv,2018),nmtD\Atiehh6a Ben canh cac cong trinh sir dung toan
•T™g,amNh.ad»v«.Ns„ bo genome ty the (mtDNA) nhiej. nghi&n - Vien Sinh Ihai va Tai nguyen pinhv^t ciiii da sir dung viing dieu khien hi,,It dpng 'Hpc vien Nong nghiep Viet Nam nhien ma ciia genome ty the (contrcl region
• Tac gia Uen he Phen, Thanh Hai, Vien Smh Ihai Xhiel ' ^ ^ „ . . !,„ „ni v i i n g D - l o o p CO k r h IhiirV,- JAi, Tmng ,am Nhiel d,n Viel-Xgartii qnoc Phong. DT " ' " ' * ' h a y COn g V g J > HUoc 0%66953m Email- phamhai.vun.gma.l.com 1.270bp n a m O V) h i n u c l e o h d e 15.4 •• I 6 7 2 7
28 KHKTChdn nuoi so 246 - Ihdng 7 ;i,;.„ 2gi,j
D l TRUYEN - G I O N G VAT N U O I
trong phan h'ch nguon go'^c, quan he hp hang va tieh hoa gan a muc loai va quan the cita loai cho (Wayne va ctv, 1990; Okumura va ctv, 1996). Thong hn thu duoc tir su da dang trinh tu vimg D-loop eiing nhu vimg ma hoa ciia mtDNA ra't co ich lrong viec phan hch xac djnh cae nhom kieu dem bpi (haplotype) va nhom don bpi (haplogroup) va xae lap nguon go'c ciia cho (Gundry va ctv, 2007; Imes va ctv, 2012; Oskarsson va etv, 2012; Frantz va ctv, 2016; Ren va ctv, 2017; Marinov va ctv, 2018),
Tlieo ke't qua nghien cuu ciia Trung tam nhipt doi Vipt Nga hong giai doan tir 2006- 2012daxacdinh 13 dang eho khae nhau, trong do CO giong cho dang soi co the thue hien to't cac kha nang nghipp vp (Tran HiJii Coi va ctv, 2011). Vipc ling dung sinh hpc phan tii' hong nghien cuu ede gio'ng cho ban dia hien nay la mpl linh vpc moi dang phal trien, chi eo nghien ciiu ciia Pham Thanh Hai va ctv (2014) ve giong cho H'mong cpe duoi va Tran Hoang Diing va ctv (2016) ve giong cho Phii Quo'c cho loi nay. Do do vipe tie'n hanh nghien cihi ve da dang di truyen ciia giohg cho dang soi la ra't can thiet. De ke'thira va phat trien cac nghien eiiu truoc day, chung loi thue hien nghien eiru da dang di Iruyen vimg D-loop cua 300 ea the dio dgng soi de xae dinh mo'i quan he di truyen nguon gen giiia ehiing va lam co so khoa hpc cho de xua't cac giai phap ehpn lpc, nhan giohg va phal trien ben vimg. Ngoai ra, giup cac nha quan ly xay dung chien luoc quan ly va dinh hudng khai thac su' dung ben vimg hipu qua eho loai eho nay 6 Viet Nam.
2. VAT LI£U VA PHUONG PH/iP NGHllN COU
2.1. Vat lipu
Tong so'300 mau mo tai va mau cua gio'ng eho dang soi do Trung lam Nhiet dpi Viet - Nga cung cap duoc dimg cho phan rich bang phucmg phap sinh hpc phan tir. Trong do, 140 mau cho due (ky hipu CS1-CS140) va 160 m i u cho cai (ky hieu'cS141-CS300).
2.2. Phuang phap
DNA long so dupc laeh chiet bang bp hoa chat QIAGEN blood and hssue Kit (QIAGEN
hic-, Diic). Nhan ban gen dich bang ky thuat PCR: thanh phan moi phan iing PCR eo the h'ch 25[il vcri cae thanh phan: 7[i\ H^O deion; 12,5pl PCR Master mix kil (2X); l,25pl moi xuoi (10 pmoI/[al); 1,25 pl moi ngugc (10 pmol/|.il); 3\ii ADN (10-20ng). Cap moi dimg trong nghien cmi CO hinh hr lan lual la 15412F-CCAC- TATCAGCACCCAAAG va 42R-CCTGAAA- CCATTGACTGAATAG (Gundry va eW, 2007). Chu trinh nhiel ciia phan img PCR gom: 95''C trong 3 phiit; tie'p sau la 40 chu ky no'i tiep nhau voi cac buoc: 95''C lrong 45 giay, 56''C lrong 45 giay, 72''C lrong 45 giay; ket thlic phan ling nhan gen 6 72"C trong 10 phiil, giii' san pham 6 4''C. San pham PCR dupe kiem tra tren gel agarose 1,5%.
Trinh tir nucleotide duac xac dinh bang ky thuat xac dinh trinh tu' Irirc tie'p tii' san pham PCR theo hai chieu (moi xuoi va moi ngugc) hen may dpc trinh hr ABl PRIMS®
3100 Avant Genetic Analyzer (Applied Biosystems, USA). Trinh tir DNA sau khi dpc dugc hieu chinh voi su trg giiip ciia phan mem ChromasPro 2,1.6 (Teehnelysium, 2013) de loai bo eae viing lin hieu nhieu, Sau do so sanh, xac dinh vi tri bie'n doi cac nucleotide, so Haplotype, dac diem di truyen cac Haplotype hong eae gio'ng cho nghien euu va miie dp da dang di truyen eiia cac gio'ng cho sir dung phan mem Bioedit v7.0-5,2 (Hall, 1999) va DnaSP (Pablo va ctv, 2009).
3.K£TQUAVATHAOUJAN
3.1. Giai trinh tu doan gen vimg D-loop gen tythe
3.1.1. Tdch chiet ADN tong so'
DNA long so cua 300 mau thupc giong cho dang soi da dugc tach ehie't thanh cong voi chat lupng DNA cao (Hinh 1). Ket qua dien di kiem ha DNA hen gel agarose 1% cho tha'y ADN khong bi gay va tuong do'i sach.
Ke't qua do OD eho thay chi so OD260/OD280
oia cae mau luon nam hong khoang 1,8 de'n
2,0 (chi so cho bie't ham luong DNA co dp tinh
kliie't cao). Ke't qua hen khang dinh eac mau
DNA laeh chiet dugc hoan loan dii tieu chuan
DI TRUYtN-Gl6N vATNLiy'
cho nhimg phan tich tiep theo. De dam bao ket qua giai trinh tv dirge chinh xac, tieh hanh
rinh san pham D \ A bing Ge"^^""' Purihcation kit cua Fernieni'^
DNA
Hinh 1. Ket qua dien di DNA long so oia mpt so mau thupc giong cho dang soi tren ge agarose l°o (cac gieng 1-50 la thu tu cac mau thupc giong cho dgng soil-
vol Irinh h^' thupc loai cho nha Canis lupus fnmiliaris co ma hieu genbank KM061586.1- 3.2. Dgc diem di huyen cua giong cho dang 3.1.2. Nlidn bdn ADN bdng ky thudt PCR
Viing gen D-Loop (mtDNA) da nhan ban lhanh cong vdi cap moi ciia Gundr\- va ctv (2007) voi nhipt dp gSn moi la 56 C cho mau nghien ciru. San pham PCR co kich thuoc khoang 600 bp cho viing gen D-Loop (mtDNA) (dung nhu kich Ihuoc ly thuyet dp doan). Chat Iugng ciia san pham PCR dupc the hipn khi dipn di tren gel agarose l,5"<i ehi eo mpt bang duv nha't, sang dam. dii tieu chuan cho tach doan gen nhan de'giai ma trinh tu.
3.3.3. Ke't qud gidi trinh tif
Doi chieu kiem tra trinh tu thu duoc vdi eac trinh ttr luong dong tren Genbank bang cong cp BLAST, da \ac nhan cac trinh tu thu dupe la cac trinh hi' dich tuong dong 99%
B«ng 1. So mJu thu^ cic Hap cua ch6 dvrc 3.2.1. Ddn cho dtrc
Co 36 vi tri bien doi nucleotide trong 140 trinh ht' eiia cac mSu cho due. Cac bieh doi da so'la bieh doi dong hoan do la su thay the'tii'T thanh C, va ngupe lai hoac cd su thay the tu' G thanh A va ngupc lgi- Cd mgt vai su bien doi di hoan hr G thanh C d HapD22, HapD13, G-T d HapDll, A-C d HapD28. Co 28 Haplotype (Hap) hong 140 hinh hi' (ki hieu hi' HapDl den HapD28)- Dac diem ciia cac Hap va so Irinh tiJ' Ihugc eae Hap dupc hinh bay d Bang 1 va Hinh 2.
K\ hieu Hap Sg trinh tu
Ky hi^u trinh hr HapD!I lapD2 H.5pD3 HapD4 HapD5 HapD6 HapD7 HapDS HapD9
CSI CS38 CS88 CS97 CSI33 CSI34 CS2 CS9 CS39 CS46 CS89 CS96 CS98
CS4 CS41 CS9! CSIOO CSI36 C5106 CS109 CS12 CS28 CS78 CS123
CSI 05
CSIO CSI 3 CS34 CS47 CS50 CS61 C564 CS&4 CS7 CSS C544 CS45 CS3CS40CS90CS99CS135
CS26CS76CSI21
30 KHKTChdn nuoi ,vo 246 - thdng 7 >u,in ytm.
PI TRUYEN - G I 6 N G VAT NUOI
Ky hi^u Hap HapDlO HapDll HapD12 HapD13 HapD14 HapDlS HapD16 HapD17 HapDlS HapD19 HapD20 HapD21 HapD22 HapD23 HapD24 HapD25 HapD26 HapD27 HapD28
So trinh t\f 3 4 6 5 7 8 4 7 5 1 10 4 7 3 7 2 5 2 5
Ky hieu trinh tu CS27 CS77 CS122 CS5 CS17 C542 CS54 CS94 CS95 C5I03 CSIIU CSI39 CS140
CS68 CS92 C5101 CS113 CS137 CS29 CS79 CS124 CS30 CS80 CSI25 CS32 CS33 CS83 CS128 CS35 CS85 CS130 CS82 CS127
CS12CS49CS63C5108 CS36 CS86 CS131 C537 CSS7 CS132 CSll
CS48 CS62 CS107 CS144 CS14 CSI 20
CS22 CS23 CS24 CS59 CS60 CS73 CS74 CS75 CSllS CS119 CS15CS52CS66CS111
CS18 CS31 CS55 CS69 CSSl CS114 CS126 CS51CS65CS110 CS19CS56CS70CSn5 CS71 CS131 C537
CS16 CS53 CS67 CS21 CS58 CS72 CS117 CS116]
CS20 CS57 CS83 CSI 28 CS35 CS85 CS8!
Hinh 2. Dac diem cac Hap dan cho due cua giong cho dang soi
D l TRUYEN - GIO-- VAT NUI-''
I • long hoan 3.2.2. Ban cho cdi soi. Cac bien doi da so la hic; Joi u » j^j
do la sir thav the tir T tlianh i va "h . Co 30 vi tn bien doi nucleotide trong 160 ^ ^ j ^ ^^ ^^ ^^^. j , , ^ (if G tlianh \ > a nguyc trinh tu' ciia giong cai thupc giohg chd dang Ket qua dupc trinli bay a Ban'_
Bang 2, So m a u thupc cac H a p d a n cho
• ,--^ Hinh 3.
Ky hi?u Hap So trinh lu Ky hi?u Irinh ty
HapCl HapC2 H a p O HapC4 HapC5 HapC6 HapC7 HapCS HapC9 HapClO HapCll HapC12 HapClS HapC14 HapClS HapC16 HapC17 HapClS HapC19 HapC20 HapC21 HapC22 HapC23 HapC24 HapC25 HapC26 HapC27 HapGS HapC29 HapC30 HapC31
CS188 CS197 CS201 CS238 CS2S8 CS297 CSUl CS:89 CS196 CSI96 CS:02 C-^il'"
^c-yAi r<i29l CS300 CS203 CS239 CS246 CS289 CS296 CS298 CS191 CS200 CS204 Cbi4J >-=•
CS240 CS290 C5299 CS194 CS195 CS207 CS20S CS244 CS245
CSI 90 CS199 CSI 93 CS149 CS168 CS191 C5205 CS2I7 CS242 CS254
CS294CS295CSI84CS284 CS210 CS213 CS234 CS247 CS250
CS268 CS292 CS2fr4 CS206CS243CS293 CSI42CS145CS161CS164 CS176 CS226 CS276 CS154 CS191 CS205
CS187 CS237 CS287 CS186 CS236 CS2S6 CS185 CS235 CS285 CS183 CS233 CS283
CS181CS232CS282 CS180CS230CS280CS279 CSI 79 CS229 CS178 CS228 CS278 CS227 CS277
CS177 CS176 CS226 CS276 CS154 CS155 CS159 CS160 CS173 CS174 CS175 CS222 CS223
CS150 CS169 CS181 CS218 CS231 CS255 CS269 CS281 CS224 CS259 CS260 CS273 CS153 CSI58 CS171 CS221 CS258 CS272 CS275 CS261 C5143 CS161 CS211 CS248 CS262 CS112 CS148 CS167
CS15I CS156 CS170 CS219 CS256 CS270 CS274 CS271 CS263
CS152 CS157 CS171 CS220 C5257 C5216 CS253 CS267 CS147 CS166 C5215 CS252 CS266
CS265 CS163 CS212 CS249 CS146CS165C^14CS251
KHKT Chdn nuoi so 246 - thdng ,,„ 21,
Dl TRUYEN - G I O N G VAT N U O I
Hinh 3. Dgc diem cac Hap dan cho cai thupc giong cho dang soi 3.3. Ung dung ky thuat di truyen trong chpn
lpc nhan giong cho dang soi
Cae gidng chd ban dia Viet Nam chii ye'u dupe nuoi tlia tir do, don le trong cac hp gia dinh, vipe quan Iy nhan gid'ng tu phat, nguon gde ciing nliu pha he ciia chung khdng dugc theo ddi va luu trii', do dd khdng chi nguon gen ciia gid'ng cho dgng sdi ma eiia eac gid'ng eho khac eo kha nang bi lap giao va de bi giao pho'i can huye't nhieu ddi gay hien tupng thoai hoa gidng, Chiing tdi tham khao khoang each di truyen (KCDT) giira cac Hap cho due va cho cai va mdi quan he di truyen cua chimg de lao ra cae the'hp sau thuan chiing va co su da dang di truyen cao.
Ma Iran khoang each di truyen va cay phat sinh chiing logi cua eac Hap dugc kiem tra cae gia thuyet va thir md hinh hen hda bang chucmg trinh phan mem MEGA 7.0 (Tamura va et\', 2016), eho tha'y md hinh thay the'nucle- otide phil hgp nhat la T92+G, Tamura 3 tiiam so (Tamura 3-parameter) vdi eac chi so' (Baye-
sian Information Criterion) BIC=3146,189, -lnL=1118,189, Base frequencies: freqA=0,291, heqC=0,209, freqG-0,209, freqT=0,291, Substi- tution model: Ti/h' ratio=62,46) (Hasegawa va ctv, 1985). Kel qua tinh loan khoang each di truyen the hien d Bang 3.
28 Hap ehd due va 31 Hap eho cai lap dugc hem 17.000 each tinh KCDT. Kel qua cho tha'y KCDT giiia eac haplotype ciia cac mau tir 0,0% de'n 3,3%. Trong nghien a i u ciia Studa va ctv (1997), KCDT giiia cac gid'ng cho lrong khoang 0-3,19%- N'hu vay, KCDT giua eae haplotype gid'ng ehd dang sdi tuong dong vdi eae giong cho khac tren the'gidi,
Theo bang 3 cho thay KCDT giiia cae haplotype cua gid'ng chd dang sdi ldn nha't la 3,3% va hong mgt Hap ed tir 1 de'n 12 ca the chd dang sdi, HapD20 chua nhieu chd due va HapC3 va HapClS ehua nhieu ca the nha't chiing to cac Hap nay bie'n dd'i ra'l dn dinh, Nhu vay, hong 59 Hap ciia giong ehd dang sdi khong ed Hap nao mang bie'n di dan le cua
33
Dl TRUYEN-GION • , V.\TNUi'-'
mpl ca the. Do dd tren nguyen tac phoi gidng giua cae ea the cd KCDT xa, ed the chgn lgc cac ca the due lir eae HapD20 cho ghep doi theo cap vdi cae HapC3 va C18 cua ca the cai. Trong mau nghien ciiu, cd 10 ca the due (ki hieu CS22 CS23 CS24 CS59 CS60 CS73 CS74 CS75 CSI 18 CSI 19) va 24 ca the cai (ki hieu: CS177 CS176 CS226 CS276 CS154 CS155 CS159 CS160 C5173 CS174 CS175 CS222 CS239 CS246 CS289 CS296 CS298 CS191 CS200 CS204 CS241 CS291 CS300 CS203 CS240 CS290 CS299) co the ghep doi nhan giong vdi nhau. Kel hpp vdi cac dac diem hinh thai de va ung dpng vipe nghien ciiu dac diem vimg gen D-loop hong cdng tac dipn gidng nham tang tinh da dgng, tang hieu qua ehpn lpc, Ininh liien lu(;mg giao phdi can huyel va giam thdi gian lrong qua trinh chpn Igc.
Bang 3. Khoang each di truyen giira cic haplotype
«,K^LUAN
, , , , „ gen CO Giai trinh tv * a n h coni, « ,^ 6^^ ^ ^ , ^ chieu dai 562bp thuoc ™ng bien •.
C g t l HapC4 HapClS HapD20 HapD20 HapD3 HapD2 HapD2 HapD14 HapD 14 HapD3 HapD16 HapD 16 HapDlO HapDlO HapD8 HapD3 HapD21 HapD21 HapD3 HapD3 HapD2 HapD2 HapDlS HapD17 HapC27 HapC2 HapC2
Cgl 2 HapC25 HapC25 HapC4 HapClS HapC25 HapC4 HapClS HapC4 HapClS HapD20 HapC4 HapClS HapC25 HapD20 HapC25 HapDU HapC4 HapClS HapC4 HapClS HapD21 HapC29 HapD23 HapDlS HapC2S HapC7 HapC19
KCDT 0,033 0,033 0,033 0,033 0,031 0,031 0,031 0,031 0,031 0,031 0,031 0.031 0.031 0.031 0,029 0,029 0,029 0.029 0.002 0.002 0,002 0,002 0,002 0,002 0,002 0,002 0,002
Cpl 1 Cgt 2 HapDlS HapC4 HapDlS HapClS HapC4 HapC26 HapC4 HapC29 HapC3 HapC25 HapClS HapC26 HapClS HapC29 HapDS HapD20 HapD3 HapD16 HapD20 HapC3
HapD2 HapDlO HapDlO HapDU HapC13 HapC25 HapD20 HapC13 HapDlO HapD36 HapC4 HapC23 HapClS HapC23 HapCl HapC4 HapD16 HapD21 HapDlD HapC3 HapC22 HapC28 HapC2 HapC20 HapDS HapC3 HapD6 HapC19 HapDS HapC21 HapD4 HapC30 HapDl HapC6
KCDT 0,029 0,029 0,029 0,029 0,029 0,029 0,029 0,029 0,029 0,029 0,029 0,029 0,029 0,029 0,029 0,029 0,029 0,029 0.002 0,002 0.002 0,002 0,000 0,000 0,000 0,000 O.OOO
gen D-loop ciia -WO mau gic '.. cho dang soi-
tir ciia cac jotide 1,; il.nlotvpe ''™''^
Da xdc dinh duoc 2 M l . i p l o t ^ P ^ ^ ^ . j ^ , tinh due ^•;l 31 Hap thuoc l'".^ • i'- ^ bieh doi nucleotide trong I mau cho dvrc va ^0 \ ! tn I' trong 160 trinh tu cua cho cJi.
Klioang cach di truyen giira cac Hap hr 0 O-y d&i 3 T o cho thay moi quan he d. truyen 0,0 /„ den 3,3 -o 1 i^i^j ^ ^ J
ciia cac giong dio ngnien LUU o D.a Kia chon duoc 10 ci the due (HapD20) v.a 24 . ^ t^e cii (HapC3 va CIS) co KCDT xanh d e c o t h e e h o g h e p d o i .
T A I L I ^ U TKAM KHAO
1 Brown W.M., G e o r g e M,J. a n d W i l s o n AC (1979), Rapid c w l u t i o n of a m m a l m i t o c h o n d r i a l D N A . Proc.
Nat Aca Sd, USA, 76:1967-71
Z C a i m s K.M., B r o w n S.K., S a c k s B,N. a n d Ballard I.W.O (2017). C o n s e r v a h o n i m p l i c a h o n s for dingoes from the niaremal a n d paternal g e n o m e , multiple populahons, d o g inlrogression, a n d d e m o g r a p h y . Ecology and Evolution, 7: 9787-07.
3 Tran H&u Coi, Bui X u a n P h u o n g , D m h T h e D u n g va N g u y e n Tien Tiing (2011), Kha n a n g h u a n luyen ch6 nghi?p v v giong H ' m o n g cpc d u o i va c h o d a n g s<5i, Tgp chi KHKT C h a n n u o i , 143, 27-31,
4 Tran H o a n g D u n g , T h a i K e Q u a n , N g u y e n T h a n h Cong, H u y n h V a n H i e u v a C h u n g A n h D u n g (2016).
Xac d m h n g u o n goc cho Phii Q u o c b a n g trinh h^ viing D-Loop trong g e n o m e ty the T a p chi Sinh hgc, 38(2):
269-78,
5 F r a n t z L-A-, M u l l i n V-E., F i o n n i e r - C a p i t a n M . , Lebras- s e u r O., O l l i v i e r M . , Perri A. a n d T r e s s e t A, (2016) G e n o m i c a n d archaeological e v i d e n c e s u g g e s t a dual origin of d o m e s t i c d o g s S a e n c e , 352 1228-31- 6 F r e e d m a n A.H,, G r o n a u I., S c h w e i z e r R,M-, Ortega-
Del Vecchyo D . , H a n E., Silva P.M. a n d N o v e m b r e J. (2014) G e n o m e s e q u e n c i n g highlights t h e d y n a m i c early history of d o g s . PLoS Generics. 10 eIG04016 7 G u n d r y R-L., M a r c W-, Allard M.W,, M o r e t t i T.R.,
H o n e y c u H R-L„ M a r k R- W i l s o n M.R., M o n s o n K . L a n d Foran D.R- (2007), Mitochondrial D N A analysis of the d o m e s t i c Dog control region vanafiun w i t h i n a n d a m o n g breeds J, Forensic Sci., 52(3)- 562-72, P h a m T h a n h H a i , D i n h T h e D i i n g . P h u c m g (2014) Dac diem g e n D-Loop , dia H ' m o n g cpc d u o i : d a d a n g , n g u o n ./
iJTig d u n g trong chpn giong. T a p chi Kl i, 189 8-14
Hall T A - (1999) BioEdit; a user-ffr ^ sequence a l i g n m e n l editor a n d anai
KHKTChdn nuoi .so 246 - thdng 'fig c h 6 ban
D! TRUYEN - GIONG VAT NUOI
W i n d o w s 9 5 / 9 K / \ l N u d A d d . Sym-Ser., 41:95-98, I m e s D . L , W i c t u m E.J., A l l a r d M.W. a n d S a c k s B.N (2012), IdentificaHon of single n u d e o t i d e p o l y m o r p h i s m s within the m t D N A g e n o m e of the domestic d o g lo discriminate i n d i v i d u a l s w i t h c o m m o n HVl haplotypes. Forensic S d hil Genel, 6(5): 630-39.
Kaneda H., H a y a s h i ] . , T a k a h a m a S., Taya C , L t n d a h l K.F. a n d Yonekawa H . (1995) Elimination of p a t e r n a l mitochondrial D N A in intraspecific crosses d u r i n g e a d y mouse embrj'ogenesis Proc, Nat. Aca, S a , USA.
92: 4542-46,
M a r i n o v M., Teofanova D., G a d j e v D., R a d o s l a v o v G. a n d H r i s t o v P. (2018) Mitochondrial diversity of Bu Iga n a n native d o g s suggests d u a l p h y l o g e n e h c origin Peer J . 6 e5060,
O k u m u r a N., I s h i g u r o N., N a k a n o M., M a t s u i A.
and Sahara M . (1996), Intra- and interbreed genetic variations of mitochondrial D N A major n o n - c o d i n g regions in Japanese native d o g b r e e d s (Canis famiharis).
Anim. C e n o l . 27:397-05,
O s k a r s s o n M . C , Klutsch C.F., B o o n y a p r a k o b U., Willon A., T a n a b e V. a n d S a v o l a i n e n P. (2012) Mitochondrial D N A d a l a indicate an i n t r o d u c h o n through Mainland Southeast Asia for Australian dingoes and Polynesian d o m e s t i c d o g s Proc Biol Sci,, 279(1730): 967-74,
Pablo L i b r a d o a n d J u l i o R o z a s (2009). DnaSP v5' A software for c o m p r e h e n s i v e analysis of D N A polymorphism data Bio inform a tics. 25,1451-52, Ren 2 , C h e n H., Yang X. a n d Z h a n g C. (2017), Phylogenetic analysis of Tibetan mastiffs b a s e d on
m i t o c h o n d r i a l hyperi^ariable region I-], Genet-, 96-119- 25-
17. S a c k s B.N., B r o w n S-K-, S t e p h e n s D., P e d e r s e n N . C , Wu J.T. a n d Berry O . (2013). Y c h r o m o s o m e analysis of d i n g o e s a n d Southeast .Asian village d o g s suggests a n e o h t h i c continental e-\pansion from Southeast Asia followed hv multiple A u s t r o n e s i a n dispersals. Moi, Bio Evo-, 13 1265-75-
18. Sindicic M., G o m e r c i c .\., C a l o v A., A r b a n a s i c H., K u s a k J., Slavica A. a n d H u b e r D . (2011), M i t o d i o n d r i a l D N A control region as a tool for species idenhfi cation a n d d i s h n c h o n b e h \ e e n \volves and d o g s from Croatia Vel, Arhiv,, 81(2) 249-58,
19. T s u d a K., K i k k a w a Y., Y o n e k a w a H . a n d T a n a b e Y- (1997) Extensive interbreeding occurred a m o n g m u l h p l e m a t n a r c h a l ancestors d u n n g the d o m e s t i c a h o n of d o g s : e v i d e n c e from inter- a n d i n t r a s p e d e s p o l y m o r p h i s m s in Uie D-loop region of m i t o c h o n d r i a l D N A between d o g s and wolves. Genes Genet, Syst,, 72(4) 229-3S
20. W a n g G.D., Z h a i W.. Yang H . C , W a n g L., Z h o n g L., Liu Y.H., Fan R.X., Yin T.T., Z h u C L , P o y a r k o v A-D., I r w i n D.M., H y l o n e n M.K,, Lohi R , Wu C.l„
S a v o l a i n e n P. a n d Z h a n g Y,P. (2015) Oul of s o u t h e r n east Asia: the natural history of d o m e s t i c d o g s acro.ss the world. Cell Res , 26 21-33.
21. W a y n e R.K,, M e y e r A-, L e h m a n N., Van V a l k e n b u r g h B„ Kal RW., Fuller T.K-, G i r m a n D . a n d O ' B r i e n S.J. (1990)- Large sequence divergence a m o n g m i t o c h o n d n a i D N A genot^'pes w i t h m p o p u l a t i o n s of eastern African black-backed jackals, Proc, Nat Aca- S d - U S A . 8 7 1772-76