» 1, · cluster anrlysis resolved tim:" main clnsters wilh gcm:tic relationships of 53 %,...

8
Slid . ./. Metru/., Vo/. 1(1), 2007 . Estimation of genetic diversity ill four Sudanese indigenous cattle breeds By l\Jakkawi, A. A. I. Sid-Ahmed, S. A. I, Ahmed , M-K. A. 2. AdaW) , S. , . '. EI-Itriby. Hanlya A. ' and Mckki, lblisam 1. 4 J. DepamllclIi of Animal ProduCiioli. Faculry ofAgril'Ulrure. Sudan Unil'ersirv o{Science alld Tec hn ology, Shall1bar, KhartoulJI No rth. 2 r'ICLIln of Allillwl ProdllCiioll, Ullive rsirY of KhanoulIl, 5;lwlJ1bm, P.O. Bo.\ .. 32. Kh arr olll1l Norrh, 3 AgriUllrural Generic Ellgilleering Research Insrilllre, Agricufrural Research CO lln cil, Egvpr, 4. N'ltiO I1<11 Cen te r. i'or Researcil, PO. Box. 2404, SUd,lO, 4 \..ly.JI t"') t y:;ll ;...",y,il AFLP C..A ......J ,l1 . , :iwlhJI ) csb.ll ' (.s --' "J' Lu) ;; --,liJl .::...L..,J......JI 0-<> .% 9 1.5 Y' tly\ U!-! .::...,llill 0LS C:-" %SO,3 L.,"" ob 0 LS -5.:0\1--, ..)LSl:>- j.ob i"1.:..:.:;.. w ",,-!\..,;::il l UPGMi\ j,b:i 0-" c.?.JI.9:iJ \ i\FLP i""""--' .%636 h..,ji.<> jS J (...s J ': l lu) .). ..iJ\ U!-!j (..cl .J-?-4) » 1 , ( ... u ':i4i) .j i,J\ U!-! %66 j %57 ,%53 '-,-lL:;j\ J %80 ,% 67 , %61 ..l.lW tl".;i U!-! :G).iJI ..::..wLS . .,JI . \ :iwL;S.llj 0-" .. '\ "\ 4...iL;S 4...ilb.., ,( , 'hi);; l.i, ) \. ,\1 % 72 .s' J-" s-- --' . ...,j .. --', Abstract 1\11 1' molec nlar markers were used to lll easure gene ti c di v<.: rsi ty of four Sudanese indi geno us ca tlk I",mely naggar,l (N i,liaw y) , Nil o ti c (Majock), Butana anel Kcnana cattle. l he level of p ol 'morphism revealed among tb e four typ es was 9 1. 5%. The da ta matrices \vcrc utili Lc 'd 10 cSLi Ill<l te til e genetic similarit y II s in g Jacca rd 's coefficient. It rangcd from -'l6,<) '7r to XO,::Wo witb an average of G:U) o/c , Tilc I\FLP dendrogram constructed i'rolll U PC; I\![. cluster anRlysis resolved tIm:" main clnsters wilh gcm:t ic relationships of 53 %, 57 ' i{ and ()(i';( be lwee ll l1 ag gara (Nia la wy) and Nil o tic ( Ma ioe k): between Haggara (l' I,1Iawy) and Bur ,l na and Kenan<1 cattie, re spectively, Vithin th e four 10«11 types the g ClIl' tic lTlatio nsitip s wc re 61%, 67?u, 80'l and T21'Jr, for Nilo lic (Majock), Baggara (N ialawy), Butallaand Kenana cattle respectively. 1ndigenous Molecular markn s Gcnetle divcrsity :53

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Page 1: » 1, · cluster anRlysis resolved tIm:" main clnsters wilh gcm:tic relationships of 53 %, 57'i{ and ()(i';( belweell l1 aggara (Niala wy) and Nilotic (Maioek): between Haggara (l'I,1Iawy)

Slid Stltf~ Metru Vo 1(1) 2007

Estimation of genetic diversity ill four Sudanese indigenous cattle breeds

By lJakkawi A A I Sid-Ahmed S A I Ahmed M-K A 2

AdaW) S EI-Itriby Hanlya A and Mckki lblisam 14

J DepamllclIi ofAnimal ProduCiioli Faculry ofAgrilUlrure Sudan Unilersirv oScience alld Technology Shall1bar KhartoulJI No rth

2 rICLIln of Allillwl ProdllCiioll Ullive rsirY of KhanoulIl 5lwlJ1bm PO Bo 32 Kharrolll1l Norrh 3 AgriUllrural Generic Ellgilleering Research Insrilllre Agricufrural Research CO llncil Egvpr

4 NltiOI1lt11 Cen ter ior Researcil PO Box 2404 Kh~rtoum SUdlO

~~

-s J~)1 4 lyJI --~) I0gt t) ~~I t yll yil AFLP ~jgtJi CAJ l1 c~1

~LSl iwlhJI +~J--4 ) csbll (s --J Lu) --liJl

~~ LJJI 0-ltgt ~ 9 15 Y tly ~J~I U- ~I ~I llill r5~ 0LS

C- SO3 ~4G9 L ob 0LS -501-- )LSlgt- job i1w ~I -ill y~ us~1

4-l~J ~Wc~ -5 ~6l 1 UPGMi jbi 0- cJI9iJ iFLP i-- ~ 636 ~ hjiltgt

jS J (s Jl lu) )iJ U-j (cl J--4) raquo 1 ( u i4i) jiJ U- 66 j 57 53 ~ cgt~ --lLj

J 80 67 61 llW tli ~J~I U- ~I G)iJI wLS JI ~ iwLSllj 4~ 0-

~ 4iLS 4ilb ( hi) li (~ ) 1 72s J- s-- -- j -- ~ ~

Abstract 111 1 molec nlar markers were used to llleasure genetic di vltrsi ty of four Sudanese

indigenous catlk Imely naggarl (N iliawy) Nilotic (Majock) Butana anel Kcnana cattle

l he level of pol morphism revealed among tbe four types was 9 1 5 The da ta matrices vcrc utili Lcd 10 cSLi Illltl te tile genetic similarit y IIsin g Jacca rd s coefficient It rangcd from -l6lt) 7r to XOWo witb an average of GU) oc Tilc IFLP dendrogram constructed irolll UPC I[ cluster anRlysis resolved tIm main clnsters wilh gcmtic relationships of 53 57 i and ()(i( belwee ll l1 aggara (Nia la wy) and Nilotic (Maioek) between Haggara ( l I1Iawy) and Bur l na and Kenanlt1 cattie re spectively Vithin th e four 10laquo11 types the gClIl tic lTlationsitips wcre 61 67u 80l and T21Jr for Nilo lic (Majock) Baggara (N ialawy) Butallaand Kenana cattle respectively

Kc~w(lrdo 1ndigenous catLl ~ Molecular markn s Gcnetle divcrsity

53

Sud ] Studs Metro VIII I ( 1)20117

Introduction There are muny ecotypes of calLIe ill the SuJan which are aJaplcd to Ihe wide

va riety of environments Many rcpulcd authors ha ve propocd varI OU S

classificaLions of Suclane~c calllc J epelldillg 011 morphlllogiul phenotypic mark rs in U1C assessmenl of oenelic diversity of nalivc middotaUh Or lhose authors 13CllIlCtl (

classified the Sudalle~e local caLtle into three maill groups Northern or Arab aud Southern or NiloLic which together CO[~LilUtc the greal hull- of Sudancse CaLtle alld

lhe relalively small callIe of the Nuba mOUlltaili

Four lypes of markers have been generally used in the assessment or g netic diversity in animals Morphological conditional uutolJOphic and molecular makers The laller encompasses elliymalic antigenic and DNA hased mark rs

The deV~()plllenl of molecular biology lechniques Jurillg the lwt dc aues creatcu new means f or stuuying livestock gel) gtlies and breeding DNA GUl now he targeted directly to sLUdy genotypes 01 animal and these techniqucs have hecome all important tool in modem day sehction and hrceding of farm animals Rtcent developri1cnts in DNA technologies ha e mauc il r ossihle to uncover a large numher of gcnetic polymoq)hisms at the DNA sequellce level and to usc them as

marker for e aluation of Lhe gencti( basis ror the observed rhenotypic variability in quantita tive uaiL5 The markers revealing variaLiou~ at DNA level arc referred to as molecular markers( I )

Molecular markers ha e played a major role it the genetic lharactcritaLiofl and variation in genomic DNA A number of ledUliquc~ of cla~sical finger rrinling haw heen particulmly usefu l for genetic analysis Of tllest lCchniques Amplified fragrnen l ltuglh pol ymorphi~m (AFT P) which is I powerful DNA marker techn ique based on tile detecljoll or DNA restriction fragmellLlt by PCR amp I iticaLion(l9 nl)

This ludy was initiated i ll order to quantify thl level of polymllrphhm among the foUl local ecol ypes or cattle ge nel ic similarilY and g(n( tic relationships using AfLP lechuiqucs

Materials and Methods Blood samples from 60 allimaL~ represen ti ng Ilalive cattle tyre ill the SUUU1

were collected from their home-land Nyala K osLi Athara allli Ulllbcnaill for lhl Baggara Nilolic13u lana amJ Kenana callie rcspecl iv Iy

Genomic DNA was iso[ated and purified acconllng to lrnl1lcga C(lrpO ralion l~J by wizard Genomic DNA purification Kit AFL Igt ana l ys i~ was performed ugting Ihc Gibcobrl AFLP analysis syslcm and AI- Ji slarter primer kit (CaL 0 10544-01 3 and 10483-0 14 respectively wi lli minor IlI()uificali(lIl)

An marker analysi s was cmTied oul by J igC~ lillg lbe lolal genomic DNA (250 ng) of each of Ille ni ll e ralldom g ro ups of local callie wi lh 1~(oRI (Rare cullcr) alld Ms e1 (FreLJucll l cutler) an d then doubl - srrandd altIapcrs were ligal d to the fragment ends Elccn fc oRl and M~e clecti e prim r comhinatiolls wen ucd ro r fragmell t a I11rll ficalio n

Sud J Stlll- Metro Vol l (l ) 2007

Data analysis Scoring o[ the data was performecl on the bu is of the pres lice or a bsence of

cadl band for all genotypes by assign ing one ( I) to vis ible bands and zero (0) to absent bands Only clear and UI ltlmbiuous bandgt were visually scored the scoring did nol consider di ffc rences in intensity of the bauds among profile [rom di fferent samples

Each band was interpreted as one allele Bands with the same mo bili ty were assumed to be I omologous (17) The pol morphiltIl1 information content (PIC) was calculated as repOft d by Morgante (11) The percen tage of pol ymorphic loci illl d mean numb r of alleles were estimated by Nei and Li (1 3) The genetic similariti es (USs) and similarity matrices were calculated using the Jaccards coefficient (6)

Clus ler illlalys is based on similali ly matri x walt obtained with unwcighled pair gro up method of ari thmetic amiddotvcrage (lJPGM Rohlf 1990) and rela tionship

WiUlin and between types were vi sualized as d ndrogram displaying the hierarchical associaLiol1s among all genotypes Computations were performed using the SAS software for data analysis (SAS Institute Inc) by B io -RAD computer system

Results and Discussion 1 Level of polymorphism

FLP technique permitted the production of (726) selecti vely amp lified DN A fragments rallging fro m 30 bp to 1600 bp in s ize and the identification of 664 (9 15) polymOll)hic markers as shown in Table 1 illld Figurel Figure 2 shows example of two of the eleven combination used in the study

Table] Number of ampl1cons generalcd and percentage polymorphism among local types or callie revealed by AFLPs marker

Primer combination Primer

code

Total No of

amplicons

Pulymolllhic

IImplimns

Polymml)hism

1 E - AA G M middotCAG 2113 82 SO 97 6

2 E - M G M - CAe 2112 7S 62 795

J E- AGe x M -CAe 7112 lt)) 82 863

E - ACC 111 middotCAG 4113 9 1 76 81 5

5 E- AAG 1gt 1- CAT 2114 60 S9 10 3

h E - AG M -CIA 2115 75 69 lt)2 0

7 E - ACT M - CAT 6114 66 64 17 0

S o IC - AG emiddot gtl- CAG 8fJ 42 4 1 97 6

Y j - AGC VI- CAA 711

Sill

74 72 97J

[0 E - AGG M -CAi 49 4 1) ~ JY

11 E - Arc M middot- CAl 4112 14 13 929

TOlar 726 (64 9 l5

55

Slid I Simi M elro Vol 1(1) 2007

DNA finger-printing of Korean cattle using AFLP markers was st udied by Chung 1lt)

and they reported that in the IIanw()o breed the numbel of N~LP bmlds per primer combination varied from 37 to 102 bands with an averag of 66 1 hallds using a 13 primer combination A tota l of 859 markers were generated ami 568 bands were polymorphic (66)- In Italian Holstei n dairy cows AFLP finger-p rin ting was used by Ajmone (3) They employed 16 primer com bi nations and generate d 1098 hands out of which 248 were polyrilOrphic averaging 155 bands pcr primcr combinat ion and ranging between 6 and 27 bmds Ajmone (2) ill vestigated variation aDOIlg Italian Friesian Italian Brown and Maremmana cattle and reported that 3J6 fragments were produced when they used AFLP to sludy lile polymorp l1isD md I J 1 of these fragment were polymorphic shy

l()()shy

90

80

7)

Amplicons 60

50

40

30

20

10

-r-

-- - -~I

r- I

F i r ~

Ill ~ i-

I ~

I

I

~

2111 2112 712 4 J 21M

-

----- 2115 6114 13m 7m

Primer combination

D Total No of llmplicolls

Polymorphic amplico[J

Fig 1 AFLP histogram showing the level of pnlymorphlsm as revealed

by each AFLP primer w mbination

56

Slid I Stnds Metro Vol 1(1) 2007

M 1 -----~ 9 9 1----shy

A B

Fig 2 Polyachrylamide gel showing AfLP profiles of four ocallypes Lane (1 2) Hagganl- Nia lawy (34) Nilulic - Majock (5 6) Bulana and (789) Kenana callie M

molecular weight slandard (lOU bp ladder) A primer- combination 23 B primer combination 24

2 Genetic similarity The breeder can use genetic similarity (Genetic distance = 100 - genetic

similarity) informaLion to make informed dec isions regarding the choice of genotypes to cross for the development of populations in order to maximize the expression of heterosis The genetic similariLies between pairs of type lines were calculiltcd using Jaccard s coefficient Pair wise similarities ranging from 46 9 to 803 with an average of 636 (Table 2) Maximum similarity of 8()Yc) was observed within Nilotic type Jines both share the same genetic background Nilotic (Majock) while the lowest genetic similarity of 46 9 was observed between Raggara and Kenana cattle which are different types from different locations

The mean genetic distance within Italian Friesian Italian Brown and Maremmana cattle studied by Aj mone (l J was 8YXi Jiang (7 ) showed that the genetic similarity within six autochthonus goat population s US ilig AFLP ranged from 0745 to 0 758 and 0951 to 0 970 between populations In Ital y Ajmone (2) investigated

57

Slid I Sinds Metro bull Iol 1([) 2007

the genelic variability in 4 goat breeds and r porld that with in hreed genetic diversity was 87 for the majori ty or breeds alld genetic similaritis hdweclI pltlirs of individuals of the same breeds averaged 070

Table 2 Genetic similarity matrix computed according to Jaccards cllcfficienl bmed on AFLP data between 9 lines of 4 local types lfiaggara - Nialawy Lines 1 and Nilutlcshy

Majock lines 3 and 4 Butana lines 5 and 6 Kenana callie breed lines 7 Sand 9)

Types Nialawy M aj ock I3 utana Kena na

1 2 3 4 5 6 7 8 9

Nialawy 1 I00 0 2 668 1000

Majock 3 651 598 1000 4 62 8 567 80 3 1000

Butana 5 603 56lt) 746 760 1000 6 54 1 520 62 1 65 1 76~ 1000

Kenana 7 527 544 559 5~ 5 5~ 2 52 6 1000 H 517 -469 _569 597 68 4 765 51 1 J 000 9 495 537 550 554 540 4lt) 2 63 470 100 0

3 GtIletic relationship

The dendrogram constructed from UPGMA cluster analysis betwe n [he local breeds based on AFLP dala resolved the 9 breed Iille~ into Ulree mail dusters a shown in Figure 3 The first cl uster with 053 genetic relationships is bctwecli Western I3aggara-N ialawy and NiloLic-Majock breeu Then the genctic relatiollship heLween Western Baggara and Nj jotic li nes are 067 amI 06 1 respectively The second cluster with 057 genetic re JaLi onship is between Western Haggara UUIula and Kenana breeus The genetic rela tionship between BUlltUla li lles is nRO The third cluster group wi th 066 genetic relaLionships is between DulanH emu KCIlltlnu catlle breed The genetic relationship betweclI line 9 8 and 7 is 072 wh ile between inc 8 and 7 of Kenana caltle breed the rd aLionship is 077 as lt110 vn in Figure 3

Fig3 Dendrogram of genelic relationship gcoclntcd om AFLP data constructed h) IJ1lGllA among 9 lines of 4 local types (Klnana nulana Nilotic - MaJock (N I) and Bag~ara - NialawyWN)

------------------~

1 ~ - t __~ I ~

M 1 Mshy

BN nN i3UI ltlIlltl

13 l1Lana

K nanltl

Kcnltlnltl

Kell am

Slid J ~11l1A Metrol Vlll I( 1) 2007

The genet ic relationltlbip~ he tween and witbin these loeal lypcs )f callIe wert IJIvesligated using duster analysis based )f) silllilarily JIlalricc~ of UPGMA and tllt dendrogram constructed resolved the 9 groups lines llllO three main cJuslcn as shown in Figure 3 The firs t with a )~ genetic relation hip hetween Baggara (Nialawy) am Nil otic (Mujock) Iype while Ihe genetic relationship iL)l1 Baggafa and Nilol ic types were 067 ltLillI 061 respecllvely The second cluster vith 0_7 genetic relationship was between Baggara (Nialawy) and Butana type whik tlle genetic relationship wi thin the Bulalla type was ORO The third luster group willI 066 gcn~tic rclaLioml1ip be tween I3utana and KClIana cat lie while [JIC range of genetilt relationship witllin KCllana caule was 072 10 077 hom t11i~ resull it is oh vious that the genetic relationship betweell KCllana and I3utana is the closest then foll oweu hy the genetic relationship hetween Baggara Kenalla I3utana Niloljc (Majo k) anu Buggma (Nialawy) me lllQre ui~tan t This result j ltonsislcllt Willl their geographic locallon and disuibutioll aou coinciues to some degree willl Illcir historical dassification mentiOlleu by Ma~oJl and Maule (iIll Rouse ( I ~ Payne (I)

anu Joshi ~) Joshi (8) esLimated phylogenetic relatjon~hip among [lanwou hincsc Yanbian unu foreign breeds They reponed thal the Hanwoo breed was Illlt~ most closely related to tbe Chinese Yanhian atlle followed by the lIereford and lIolstein hreeds

Ajmone (2) presented a d endrogram lmd concluded t11at t1le Italian Friesian and Italian Brown catlle were slightl y more closely related to each olher than tJley were to MaremmU1a caule Cluster analysis in six autochthonous goal populations ludieu hy Jiang et II (2003) using AfLP mdicaled t11al Chengdu Grey goal (eGG) is the most uislant popu lation while Chuunuong White goaL (CWO) and Yangtze Riwr Oel t~l White goat (ROWG) were IllC dosest popu lations followed hy Boujio gnat ( B(J ) Hui goat (HG)ltUld Malou goat (MCi)

References I AjmonemiddotMarsan II Negrinl R Milancsi E Bozzi R Nijman I 1 bullbull Ilunljcr J fl

Valentin A and LenstJ3 J A (211110) G ne ti c di stances wi tiJin and )Cross cat ile hrecus as illuitlt1 led b biallcl ic rLP lllatkcr Ani ma] Gelle(ics 33 2~O-2R6

2 Ajll1one-l1lttJsan P bull Otsen M bullbull NegJini R Vecchiulli-Anlald Cbull KUiper Mbull Len~lla JA and Piva C 0(99) Genetic diycrsily iLlIin an d betwlcn cIU le bneds a ~ measured wi th AFLP nlm)crs Recent 1 rore in limal production Science 1 Procttdi l1 ~ oj lbl AS]A XlU Congress PinCCII 7lt1 tt aly 21 - 2-l Jllllt

t 000 1 55 - 1 57

3 Ajmonc-lfanIII P Valentini A Ca~sandro M VccchioltjmiddotAntll ldl C Bcrtoni G and Kuiper M (1 997 )IFLP mlJ-kcrs Ill DNA fingerpri nl ing ill Cltlu it Anima l Genetics 2~ 4 18 - 420

4 Bennett S C J John J R and nwi~Hn J W (1954) nillwl Iusbdildry III At-ric nll lm in the S lId ll1 [o ll1ill l D (td ) pr 6n - 667 ()x ro rd l Jni erStt) Prcs~ UK

i)

SlIn ] SCnn Metrol Vol 1() 2J07

5 Chung ER Kim WT Kim YS and nan SK (2000) DN middot fingerprintin g of Korenn cattle llsiug ArLP markers Korean JOllrnal of Animal Science 42 19 1 shy406

6 Jaccard P (l9()8) ~ NOllve lles recherches sur la dislribution flora Ie Buil -SOc Vauu Sci Nat 44 223 - 270

7 Jiang XP Liu GO Ding JT Yang LG Cau Sx and Cheng S() (2003) Diversity in six gont populatiolls in the middle and lower Yangtze River Vallcy Asian Australian Journal of Animal Sciences 16 277 - 2~ I

8 Joshi C G Rank D N Brahmakshtrl It P Palel A V Vataliya P II Muraleed haran P Khoda V K and Soiankii 1 V (1998) RJ PO analsys by PCR llsing arbitrary primers in Jifferent animal species Indian Vet 1 75 1029-1031

9 Joshi N R McLaughlin E A and Phillips R W (l1S7) Types Hnd breeds of African Cattle Agric Stud No 37 FAO Rome

10 Mason 1 L and Maule J P (]960) T he indi ge nous li vestock of Eastern and Southern Africa Conunon Bnr An ima l Breed Genet Tech Com-mun No 14 Commonwealth Agricultural Bureaux Farnham Royal UK

] 1 Mitra A Schlee P Krause I Blusch bull J Werner T Balakrishnan C R and Pirchner r (1998) Molec ular marker and their applications in livestock improvement Anim B iotech 9 81- 87

12 Morganle M Raralski J A lliddlc P Tingey S and Olivieri A M (1994) Genetic mapping and variability of seven soybean simple sequenee repea r loci Genome 37 763 - 769

]3 Net M and Lt W H (1979) Mathematical model for studying genetic variation in tenus of restriction endonnceases Proceedings of the al io nal Academy of Sciences US A 76 5269 - 5273

14 Payne W J A (1970) Caltle Prodllction in the Tropics Vo l I Breeds and Breeding Longman Gronp Ltd Lo ndon P 65

15 Promega Corporation (1999) WiLltlld GeIli)mic DNA purificltioll Kit Technica l Manual 2800 Woods Ho llow Road Matli so n US A

16 Ruhlf F J (1990) NTS S-P Numerical Taxonomy and Multivariate Analysis System Version 17 Owners Manual

17 Rouppe Van der VoorlJNAM Van Zandvoort P Van Eck if 01 Fulkerlsma R T Hutten R n c Draainrabull1 Gummers (1 bull 1 Jaco h~en E Helder bullJ and Baker J (1997) Use of allele specificity of comi grating AFLP markers to align genetic maps from different potato ge notypes Mol (l cn l icnet 255 438 shy477

Ill Rouse J (1972) World Cattle Il Cattle of Afriea and AsiR Okhlhoma Uni versity Press OkLlhoma USA

19 Vos Igt Rogers R Bleeker M Rcijms M Van de Lee T Uorncs M Fijlers A Pot 1 Peleman J Kuiper M and Z3ueau Me ( 1I95) AfLP a Hew

technique for DNA fingerprinting Nuc Acids Res 23 4407 - 4414

20 Zabeml M and Vos P (1993) Selective restrictioll fragment ampliiicalion a eeneral method for DNA fingcrprinllng EuropcMl Patent Application of 92402G 2SJ Pub

o 053485~A 1 bull

60

Page 2: » 1, · cluster anRlysis resolved tIm:" main clnsters wilh gcm:tic relationships of 53 %, 57'i{ and ()(i';( belweell l1 aggara (Niala wy) and Nilotic (Maioek): between Haggara (l'I,1Iawy)

Sud ] Studs Metro VIII I ( 1)20117

Introduction There are muny ecotypes of calLIe ill the SuJan which are aJaplcd to Ihe wide

va riety of environments Many rcpulcd authors ha ve propocd varI OU S

classificaLions of Suclane~c calllc J epelldillg 011 morphlllogiul phenotypic mark rs in U1C assessmenl of oenelic diversity of nalivc middotaUh Or lhose authors 13CllIlCtl (

classified the Sudalle~e local caLtle into three maill groups Northern or Arab aud Southern or NiloLic which together CO[~LilUtc the greal hull- of Sudancse CaLtle alld

lhe relalively small callIe of the Nuba mOUlltaili

Four lypes of markers have been generally used in the assessment or g netic diversity in animals Morphological conditional uutolJOphic and molecular makers The laller encompasses elliymalic antigenic and DNA hased mark rs

The deV~()plllenl of molecular biology lechniques Jurillg the lwt dc aues creatcu new means f or stuuying livestock gel) gtlies and breeding DNA GUl now he targeted directly to sLUdy genotypes 01 animal and these techniqucs have hecome all important tool in modem day sehction and hrceding of farm animals Rtcent developri1cnts in DNA technologies ha e mauc il r ossihle to uncover a large numher of gcnetic polymoq)hisms at the DNA sequellce level and to usc them as

marker for e aluation of Lhe gencti( basis ror the observed rhenotypic variability in quantita tive uaiL5 The markers revealing variaLiou~ at DNA level arc referred to as molecular markers( I )

Molecular markers ha e played a major role it the genetic lharactcritaLiofl and variation in genomic DNA A number of ledUliquc~ of cla~sical finger rrinling haw heen particulmly usefu l for genetic analysis Of tllest lCchniques Amplified fragrnen l ltuglh pol ymorphi~m (AFT P) which is I powerful DNA marker techn ique based on tile detecljoll or DNA restriction fragmellLlt by PCR amp I iticaLion(l9 nl)

This ludy was initiated i ll order to quantify thl level of polymllrphhm among the foUl local ecol ypes or cattle ge nel ic similarilY and g(n( tic relationships using AfLP lechuiqucs

Materials and Methods Blood samples from 60 allimaL~ represen ti ng Ilalive cattle tyre ill the SUUU1

were collected from their home-land Nyala K osLi Athara allli Ulllbcnaill for lhl Baggara Nilolic13u lana amJ Kenana callie rcspecl iv Iy

Genomic DNA was iso[ated and purified acconllng to lrnl1lcga C(lrpO ralion l~J by wizard Genomic DNA purification Kit AFL Igt ana l ys i~ was performed ugting Ihc Gibcobrl AFLP analysis syslcm and AI- Ji slarter primer kit (CaL 0 10544-01 3 and 10483-0 14 respectively wi lli minor IlI()uificali(lIl)

An marker analysi s was cmTied oul by J igC~ lillg lbe lolal genomic DNA (250 ng) of each of Ille ni ll e ralldom g ro ups of local callie wi lh 1~(oRI (Rare cullcr) alld Ms e1 (FreLJucll l cutler) an d then doubl - srrandd altIapcrs were ligal d to the fragment ends Elccn fc oRl and M~e clecti e prim r comhinatiolls wen ucd ro r fragmell t a I11rll ficalio n

Sud J Stlll- Metro Vol l (l ) 2007

Data analysis Scoring o[ the data was performecl on the bu is of the pres lice or a bsence of

cadl band for all genotypes by assign ing one ( I) to vis ible bands and zero (0) to absent bands Only clear and UI ltlmbiuous bandgt were visually scored the scoring did nol consider di ffc rences in intensity of the bauds among profile [rom di fferent samples

Each band was interpreted as one allele Bands with the same mo bili ty were assumed to be I omologous (17) The pol morphiltIl1 information content (PIC) was calculated as repOft d by Morgante (11) The percen tage of pol ymorphic loci illl d mean numb r of alleles were estimated by Nei and Li (1 3) The genetic similariti es (USs) and similarity matrices were calculated using the Jaccards coefficient (6)

Clus ler illlalys is based on similali ly matri x walt obtained with unwcighled pair gro up method of ari thmetic amiddotvcrage (lJPGM Rohlf 1990) and rela tionship

WiUlin and between types were vi sualized as d ndrogram displaying the hierarchical associaLiol1s among all genotypes Computations were performed using the SAS software for data analysis (SAS Institute Inc) by B io -RAD computer system

Results and Discussion 1 Level of polymorphism

FLP technique permitted the production of (726) selecti vely amp lified DN A fragments rallging fro m 30 bp to 1600 bp in s ize and the identification of 664 (9 15) polymOll)hic markers as shown in Table 1 illld Figurel Figure 2 shows example of two of the eleven combination used in the study

Table] Number of ampl1cons generalcd and percentage polymorphism among local types or callie revealed by AFLPs marker

Primer combination Primer

code

Total No of

amplicons

Pulymolllhic

IImplimns

Polymml)hism

1 E - AA G M middotCAG 2113 82 SO 97 6

2 E - M G M - CAe 2112 7S 62 795

J E- AGe x M -CAe 7112 lt)) 82 863

E - ACC 111 middotCAG 4113 9 1 76 81 5

5 E- AAG 1gt 1- CAT 2114 60 S9 10 3

h E - AG M -CIA 2115 75 69 lt)2 0

7 E - ACT M - CAT 6114 66 64 17 0

S o IC - AG emiddot gtl- CAG 8fJ 42 4 1 97 6

Y j - AGC VI- CAA 711

Sill

74 72 97J

[0 E - AGG M -CAi 49 4 1) ~ JY

11 E - Arc M middot- CAl 4112 14 13 929

TOlar 726 (64 9 l5

55

Slid I Simi M elro Vol 1(1) 2007

DNA finger-printing of Korean cattle using AFLP markers was st udied by Chung 1lt)

and they reported that in the IIanw()o breed the numbel of N~LP bmlds per primer combination varied from 37 to 102 bands with an averag of 66 1 hallds using a 13 primer combination A tota l of 859 markers were generated ami 568 bands were polymorphic (66)- In Italian Holstei n dairy cows AFLP finger-p rin ting was used by Ajmone (3) They employed 16 primer com bi nations and generate d 1098 hands out of which 248 were polyrilOrphic averaging 155 bands pcr primcr combinat ion and ranging between 6 and 27 bmds Ajmone (2) ill vestigated variation aDOIlg Italian Friesian Italian Brown and Maremmana cattle and reported that 3J6 fragments were produced when they used AFLP to sludy lile polymorp l1isD md I J 1 of these fragment were polymorphic shy

l()()shy

90

80

7)

Amplicons 60

50

40

30

20

10

-r-

-- - -~I

r- I

F i r ~

Ill ~ i-

I ~

I

I

~

2111 2112 712 4 J 21M

-

----- 2115 6114 13m 7m

Primer combination

D Total No of llmplicolls

Polymorphic amplico[J

Fig 1 AFLP histogram showing the level of pnlymorphlsm as revealed

by each AFLP primer w mbination

56

Slid I Stnds Metro Vol 1(1) 2007

M 1 -----~ 9 9 1----shy

A B

Fig 2 Polyachrylamide gel showing AfLP profiles of four ocallypes Lane (1 2) Hagganl- Nia lawy (34) Nilulic - Majock (5 6) Bulana and (789) Kenana callie M

molecular weight slandard (lOU bp ladder) A primer- combination 23 B primer combination 24

2 Genetic similarity The breeder can use genetic similarity (Genetic distance = 100 - genetic

similarity) informaLion to make informed dec isions regarding the choice of genotypes to cross for the development of populations in order to maximize the expression of heterosis The genetic similariLies between pairs of type lines were calculiltcd using Jaccard s coefficient Pair wise similarities ranging from 46 9 to 803 with an average of 636 (Table 2) Maximum similarity of 8()Yc) was observed within Nilotic type Jines both share the same genetic background Nilotic (Majock) while the lowest genetic similarity of 46 9 was observed between Raggara and Kenana cattle which are different types from different locations

The mean genetic distance within Italian Friesian Italian Brown and Maremmana cattle studied by Aj mone (l J was 8YXi Jiang (7 ) showed that the genetic similarity within six autochthonus goat population s US ilig AFLP ranged from 0745 to 0 758 and 0951 to 0 970 between populations In Ital y Ajmone (2) investigated

57

Slid I Sinds Metro bull Iol 1([) 2007

the genelic variability in 4 goat breeds and r porld that with in hreed genetic diversity was 87 for the majori ty or breeds alld genetic similaritis hdweclI pltlirs of individuals of the same breeds averaged 070

Table 2 Genetic similarity matrix computed according to Jaccards cllcfficienl bmed on AFLP data between 9 lines of 4 local types lfiaggara - Nialawy Lines 1 and Nilutlcshy

Majock lines 3 and 4 Butana lines 5 and 6 Kenana callie breed lines 7 Sand 9)

Types Nialawy M aj ock I3 utana Kena na

1 2 3 4 5 6 7 8 9

Nialawy 1 I00 0 2 668 1000

Majock 3 651 598 1000 4 62 8 567 80 3 1000

Butana 5 603 56lt) 746 760 1000 6 54 1 520 62 1 65 1 76~ 1000

Kenana 7 527 544 559 5~ 5 5~ 2 52 6 1000 H 517 -469 _569 597 68 4 765 51 1 J 000 9 495 537 550 554 540 4lt) 2 63 470 100 0

3 GtIletic relationship

The dendrogram constructed from UPGMA cluster analysis betwe n [he local breeds based on AFLP dala resolved the 9 breed Iille~ into Ulree mail dusters a shown in Figure 3 The first cl uster with 053 genetic relationships is bctwecli Western I3aggara-N ialawy and NiloLic-Majock breeu Then the genctic relatiollship heLween Western Baggara and Nj jotic li nes are 067 amI 06 1 respectively The second cluster with 057 genetic re JaLi onship is between Western Haggara UUIula and Kenana breeus The genetic rela tionship between BUlltUla li lles is nRO The third cluster group wi th 066 genetic relaLionships is between DulanH emu KCIlltlnu catlle breed The genetic relationship betweclI line 9 8 and 7 is 072 wh ile between inc 8 and 7 of Kenana caltle breed the rd aLionship is 077 as lt110 vn in Figure 3

Fig3 Dendrogram of genelic relationship gcoclntcd om AFLP data constructed h) IJ1lGllA among 9 lines of 4 local types (Klnana nulana Nilotic - MaJock (N I) and Bag~ara - NialawyWN)

------------------~

1 ~ - t __~ I ~

M 1 Mshy

BN nN i3UI ltlIlltl

13 l1Lana

K nanltl

Kcnltlnltl

Kell am

Slid J ~11l1A Metrol Vlll I( 1) 2007

The genet ic relationltlbip~ he tween and witbin these loeal lypcs )f callIe wert IJIvesligated using duster analysis based )f) silllilarily JIlalricc~ of UPGMA and tllt dendrogram constructed resolved the 9 groups lines llllO three main cJuslcn as shown in Figure 3 The firs t with a )~ genetic relation hip hetween Baggara (Nialawy) am Nil otic (Mujock) Iype while Ihe genetic relationship iL)l1 Baggafa and Nilol ic types were 067 ltLillI 061 respecllvely The second cluster vith 0_7 genetic relationship was between Baggara (Nialawy) and Butana type whik tlle genetic relationship wi thin the Bulalla type was ORO The third luster group willI 066 gcn~tic rclaLioml1ip be tween I3utana and KClIana cat lie while [JIC range of genetilt relationship witllin KCllana caule was 072 10 077 hom t11i~ resull it is oh vious that the genetic relationship betweell KCllana and I3utana is the closest then foll oweu hy the genetic relationship hetween Baggara Kenalla I3utana Niloljc (Majo k) anu Buggma (Nialawy) me lllQre ui~tan t This result j ltonsislcllt Willl their geographic locallon and disuibutioll aou coinciues to some degree willl Illcir historical dassification mentiOlleu by Ma~oJl and Maule (iIll Rouse ( I ~ Payne (I)

anu Joshi ~) Joshi (8) esLimated phylogenetic relatjon~hip among [lanwou hincsc Yanbian unu foreign breeds They reponed thal the Hanwoo breed was Illlt~ most closely related to tbe Chinese Yanhian atlle followed by the lIereford and lIolstein hreeds

Ajmone (2) presented a d endrogram lmd concluded t11at t1le Italian Friesian and Italian Brown catlle were slightl y more closely related to each olher than tJley were to MaremmU1a caule Cluster analysis in six autochthonous goal populations ludieu hy Jiang et II (2003) using AfLP mdicaled t11al Chengdu Grey goal (eGG) is the most uislant popu lation while Chuunuong White goaL (CWO) and Yangtze Riwr Oel t~l White goat (ROWG) were IllC dosest popu lations followed hy Boujio gnat ( B(J ) Hui goat (HG)ltUld Malou goat (MCi)

References I AjmonemiddotMarsan II Negrinl R Milancsi E Bozzi R Nijman I 1 bullbull Ilunljcr J fl

Valentin A and LenstJ3 J A (211110) G ne ti c di stances wi tiJin and )Cross cat ile hrecus as illuitlt1 led b biallcl ic rLP lllatkcr Ani ma] Gelle(ics 33 2~O-2R6

2 Ajll1one-l1lttJsan P bull Otsen M bullbull NegJini R Vecchiulli-Anlald Cbull KUiper Mbull Len~lla JA and Piva C 0(99) Genetic diycrsily iLlIin an d betwlcn cIU le bneds a ~ measured wi th AFLP nlm)crs Recent 1 rore in limal production Science 1 Procttdi l1 ~ oj lbl AS]A XlU Congress PinCCII 7lt1 tt aly 21 - 2-l Jllllt

t 000 1 55 - 1 57

3 Ajmonc-lfanIII P Valentini A Ca~sandro M VccchioltjmiddotAntll ldl C Bcrtoni G and Kuiper M (1 997 )IFLP mlJ-kcrs Ill DNA fingerpri nl ing ill Cltlu it Anima l Genetics 2~ 4 18 - 420

4 Bennett S C J John J R and nwi~Hn J W (1954) nillwl Iusbdildry III At-ric nll lm in the S lId ll1 [o ll1ill l D (td ) pr 6n - 667 ()x ro rd l Jni erStt) Prcs~ UK

i)

SlIn ] SCnn Metrol Vol 1() 2J07

5 Chung ER Kim WT Kim YS and nan SK (2000) DN middot fingerprintin g of Korenn cattle llsiug ArLP markers Korean JOllrnal of Animal Science 42 19 1 shy406

6 Jaccard P (l9()8) ~ NOllve lles recherches sur la dislribution flora Ie Buil -SOc Vauu Sci Nat 44 223 - 270

7 Jiang XP Liu GO Ding JT Yang LG Cau Sx and Cheng S() (2003) Diversity in six gont populatiolls in the middle and lower Yangtze River Vallcy Asian Australian Journal of Animal Sciences 16 277 - 2~ I

8 Joshi C G Rank D N Brahmakshtrl It P Palel A V Vataliya P II Muraleed haran P Khoda V K and Soiankii 1 V (1998) RJ PO analsys by PCR llsing arbitrary primers in Jifferent animal species Indian Vet 1 75 1029-1031

9 Joshi N R McLaughlin E A and Phillips R W (l1S7) Types Hnd breeds of African Cattle Agric Stud No 37 FAO Rome

10 Mason 1 L and Maule J P (]960) T he indi ge nous li vestock of Eastern and Southern Africa Conunon Bnr An ima l Breed Genet Tech Com-mun No 14 Commonwealth Agricultural Bureaux Farnham Royal UK

] 1 Mitra A Schlee P Krause I Blusch bull J Werner T Balakrishnan C R and Pirchner r (1998) Molec ular marker and their applications in livestock improvement Anim B iotech 9 81- 87

12 Morganle M Raralski J A lliddlc P Tingey S and Olivieri A M (1994) Genetic mapping and variability of seven soybean simple sequenee repea r loci Genome 37 763 - 769

]3 Net M and Lt W H (1979) Mathematical model for studying genetic variation in tenus of restriction endonnceases Proceedings of the al io nal Academy of Sciences US A 76 5269 - 5273

14 Payne W J A (1970) Caltle Prodllction in the Tropics Vo l I Breeds and Breeding Longman Gronp Ltd Lo ndon P 65

15 Promega Corporation (1999) WiLltlld GeIli)mic DNA purificltioll Kit Technica l Manual 2800 Woods Ho llow Road Matli so n US A

16 Ruhlf F J (1990) NTS S-P Numerical Taxonomy and Multivariate Analysis System Version 17 Owners Manual

17 Rouppe Van der VoorlJNAM Van Zandvoort P Van Eck if 01 Fulkerlsma R T Hutten R n c Draainrabull1 Gummers (1 bull 1 Jaco h~en E Helder bullJ and Baker J (1997) Use of allele specificity of comi grating AFLP markers to align genetic maps from different potato ge notypes Mol (l cn l icnet 255 438 shy477

Ill Rouse J (1972) World Cattle Il Cattle of Afriea and AsiR Okhlhoma Uni versity Press OkLlhoma USA

19 Vos Igt Rogers R Bleeker M Rcijms M Van de Lee T Uorncs M Fijlers A Pot 1 Peleman J Kuiper M and Z3ueau Me ( 1I95) AfLP a Hew

technique for DNA fingerprinting Nuc Acids Res 23 4407 - 4414

20 Zabeml M and Vos P (1993) Selective restrictioll fragment ampliiicalion a eeneral method for DNA fingcrprinllng EuropcMl Patent Application of 92402G 2SJ Pub

o 053485~A 1 bull

60

Page 3: » 1, · cluster anRlysis resolved tIm:" main clnsters wilh gcm:tic relationships of 53 %, 57'i{ and ()(i';( belweell l1 aggara (Niala wy) and Nilotic (Maioek): between Haggara (l'I,1Iawy)

Sud J Stlll- Metro Vol l (l ) 2007

Data analysis Scoring o[ the data was performecl on the bu is of the pres lice or a bsence of

cadl band for all genotypes by assign ing one ( I) to vis ible bands and zero (0) to absent bands Only clear and UI ltlmbiuous bandgt were visually scored the scoring did nol consider di ffc rences in intensity of the bauds among profile [rom di fferent samples

Each band was interpreted as one allele Bands with the same mo bili ty were assumed to be I omologous (17) The pol morphiltIl1 information content (PIC) was calculated as repOft d by Morgante (11) The percen tage of pol ymorphic loci illl d mean numb r of alleles were estimated by Nei and Li (1 3) The genetic similariti es (USs) and similarity matrices were calculated using the Jaccards coefficient (6)

Clus ler illlalys is based on similali ly matri x walt obtained with unwcighled pair gro up method of ari thmetic amiddotvcrage (lJPGM Rohlf 1990) and rela tionship

WiUlin and between types were vi sualized as d ndrogram displaying the hierarchical associaLiol1s among all genotypes Computations were performed using the SAS software for data analysis (SAS Institute Inc) by B io -RAD computer system

Results and Discussion 1 Level of polymorphism

FLP technique permitted the production of (726) selecti vely amp lified DN A fragments rallging fro m 30 bp to 1600 bp in s ize and the identification of 664 (9 15) polymOll)hic markers as shown in Table 1 illld Figurel Figure 2 shows example of two of the eleven combination used in the study

Table] Number of ampl1cons generalcd and percentage polymorphism among local types or callie revealed by AFLPs marker

Primer combination Primer

code

Total No of

amplicons

Pulymolllhic

IImplimns

Polymml)hism

1 E - AA G M middotCAG 2113 82 SO 97 6

2 E - M G M - CAe 2112 7S 62 795

J E- AGe x M -CAe 7112 lt)) 82 863

E - ACC 111 middotCAG 4113 9 1 76 81 5

5 E- AAG 1gt 1- CAT 2114 60 S9 10 3

h E - AG M -CIA 2115 75 69 lt)2 0

7 E - ACT M - CAT 6114 66 64 17 0

S o IC - AG emiddot gtl- CAG 8fJ 42 4 1 97 6

Y j - AGC VI- CAA 711

Sill

74 72 97J

[0 E - AGG M -CAi 49 4 1) ~ JY

11 E - Arc M middot- CAl 4112 14 13 929

TOlar 726 (64 9 l5

55

Slid I Simi M elro Vol 1(1) 2007

DNA finger-printing of Korean cattle using AFLP markers was st udied by Chung 1lt)

and they reported that in the IIanw()o breed the numbel of N~LP bmlds per primer combination varied from 37 to 102 bands with an averag of 66 1 hallds using a 13 primer combination A tota l of 859 markers were generated ami 568 bands were polymorphic (66)- In Italian Holstei n dairy cows AFLP finger-p rin ting was used by Ajmone (3) They employed 16 primer com bi nations and generate d 1098 hands out of which 248 were polyrilOrphic averaging 155 bands pcr primcr combinat ion and ranging between 6 and 27 bmds Ajmone (2) ill vestigated variation aDOIlg Italian Friesian Italian Brown and Maremmana cattle and reported that 3J6 fragments were produced when they used AFLP to sludy lile polymorp l1isD md I J 1 of these fragment were polymorphic shy

l()()shy

90

80

7)

Amplicons 60

50

40

30

20

10

-r-

-- - -~I

r- I

F i r ~

Ill ~ i-

I ~

I

I

~

2111 2112 712 4 J 21M

-

----- 2115 6114 13m 7m

Primer combination

D Total No of llmplicolls

Polymorphic amplico[J

Fig 1 AFLP histogram showing the level of pnlymorphlsm as revealed

by each AFLP primer w mbination

56

Slid I Stnds Metro Vol 1(1) 2007

M 1 -----~ 9 9 1----shy

A B

Fig 2 Polyachrylamide gel showing AfLP profiles of four ocallypes Lane (1 2) Hagganl- Nia lawy (34) Nilulic - Majock (5 6) Bulana and (789) Kenana callie M

molecular weight slandard (lOU bp ladder) A primer- combination 23 B primer combination 24

2 Genetic similarity The breeder can use genetic similarity (Genetic distance = 100 - genetic

similarity) informaLion to make informed dec isions regarding the choice of genotypes to cross for the development of populations in order to maximize the expression of heterosis The genetic similariLies between pairs of type lines were calculiltcd using Jaccard s coefficient Pair wise similarities ranging from 46 9 to 803 with an average of 636 (Table 2) Maximum similarity of 8()Yc) was observed within Nilotic type Jines both share the same genetic background Nilotic (Majock) while the lowest genetic similarity of 46 9 was observed between Raggara and Kenana cattle which are different types from different locations

The mean genetic distance within Italian Friesian Italian Brown and Maremmana cattle studied by Aj mone (l J was 8YXi Jiang (7 ) showed that the genetic similarity within six autochthonus goat population s US ilig AFLP ranged from 0745 to 0 758 and 0951 to 0 970 between populations In Ital y Ajmone (2) investigated

57

Slid I Sinds Metro bull Iol 1([) 2007

the genelic variability in 4 goat breeds and r porld that with in hreed genetic diversity was 87 for the majori ty or breeds alld genetic similaritis hdweclI pltlirs of individuals of the same breeds averaged 070

Table 2 Genetic similarity matrix computed according to Jaccards cllcfficienl bmed on AFLP data between 9 lines of 4 local types lfiaggara - Nialawy Lines 1 and Nilutlcshy

Majock lines 3 and 4 Butana lines 5 and 6 Kenana callie breed lines 7 Sand 9)

Types Nialawy M aj ock I3 utana Kena na

1 2 3 4 5 6 7 8 9

Nialawy 1 I00 0 2 668 1000

Majock 3 651 598 1000 4 62 8 567 80 3 1000

Butana 5 603 56lt) 746 760 1000 6 54 1 520 62 1 65 1 76~ 1000

Kenana 7 527 544 559 5~ 5 5~ 2 52 6 1000 H 517 -469 _569 597 68 4 765 51 1 J 000 9 495 537 550 554 540 4lt) 2 63 470 100 0

3 GtIletic relationship

The dendrogram constructed from UPGMA cluster analysis betwe n [he local breeds based on AFLP dala resolved the 9 breed Iille~ into Ulree mail dusters a shown in Figure 3 The first cl uster with 053 genetic relationships is bctwecli Western I3aggara-N ialawy and NiloLic-Majock breeu Then the genctic relatiollship heLween Western Baggara and Nj jotic li nes are 067 amI 06 1 respectively The second cluster with 057 genetic re JaLi onship is between Western Haggara UUIula and Kenana breeus The genetic rela tionship between BUlltUla li lles is nRO The third cluster group wi th 066 genetic relaLionships is between DulanH emu KCIlltlnu catlle breed The genetic relationship betweclI line 9 8 and 7 is 072 wh ile between inc 8 and 7 of Kenana caltle breed the rd aLionship is 077 as lt110 vn in Figure 3

Fig3 Dendrogram of genelic relationship gcoclntcd om AFLP data constructed h) IJ1lGllA among 9 lines of 4 local types (Klnana nulana Nilotic - MaJock (N I) and Bag~ara - NialawyWN)

------------------~

1 ~ - t __~ I ~

M 1 Mshy

BN nN i3UI ltlIlltl

13 l1Lana

K nanltl

Kcnltlnltl

Kell am

Slid J ~11l1A Metrol Vlll I( 1) 2007

The genet ic relationltlbip~ he tween and witbin these loeal lypcs )f callIe wert IJIvesligated using duster analysis based )f) silllilarily JIlalricc~ of UPGMA and tllt dendrogram constructed resolved the 9 groups lines llllO three main cJuslcn as shown in Figure 3 The firs t with a )~ genetic relation hip hetween Baggara (Nialawy) am Nil otic (Mujock) Iype while Ihe genetic relationship iL)l1 Baggafa and Nilol ic types were 067 ltLillI 061 respecllvely The second cluster vith 0_7 genetic relationship was between Baggara (Nialawy) and Butana type whik tlle genetic relationship wi thin the Bulalla type was ORO The third luster group willI 066 gcn~tic rclaLioml1ip be tween I3utana and KClIana cat lie while [JIC range of genetilt relationship witllin KCllana caule was 072 10 077 hom t11i~ resull it is oh vious that the genetic relationship betweell KCllana and I3utana is the closest then foll oweu hy the genetic relationship hetween Baggara Kenalla I3utana Niloljc (Majo k) anu Buggma (Nialawy) me lllQre ui~tan t This result j ltonsislcllt Willl their geographic locallon and disuibutioll aou coinciues to some degree willl Illcir historical dassification mentiOlleu by Ma~oJl and Maule (iIll Rouse ( I ~ Payne (I)

anu Joshi ~) Joshi (8) esLimated phylogenetic relatjon~hip among [lanwou hincsc Yanbian unu foreign breeds They reponed thal the Hanwoo breed was Illlt~ most closely related to tbe Chinese Yanhian atlle followed by the lIereford and lIolstein hreeds

Ajmone (2) presented a d endrogram lmd concluded t11at t1le Italian Friesian and Italian Brown catlle were slightl y more closely related to each olher than tJley were to MaremmU1a caule Cluster analysis in six autochthonous goal populations ludieu hy Jiang et II (2003) using AfLP mdicaled t11al Chengdu Grey goal (eGG) is the most uislant popu lation while Chuunuong White goaL (CWO) and Yangtze Riwr Oel t~l White goat (ROWG) were IllC dosest popu lations followed hy Boujio gnat ( B(J ) Hui goat (HG)ltUld Malou goat (MCi)

References I AjmonemiddotMarsan II Negrinl R Milancsi E Bozzi R Nijman I 1 bullbull Ilunljcr J fl

Valentin A and LenstJ3 J A (211110) G ne ti c di stances wi tiJin and )Cross cat ile hrecus as illuitlt1 led b biallcl ic rLP lllatkcr Ani ma] Gelle(ics 33 2~O-2R6

2 Ajll1one-l1lttJsan P bull Otsen M bullbull NegJini R Vecchiulli-Anlald Cbull KUiper Mbull Len~lla JA and Piva C 0(99) Genetic diycrsily iLlIin an d betwlcn cIU le bneds a ~ measured wi th AFLP nlm)crs Recent 1 rore in limal production Science 1 Procttdi l1 ~ oj lbl AS]A XlU Congress PinCCII 7lt1 tt aly 21 - 2-l Jllllt

t 000 1 55 - 1 57

3 Ajmonc-lfanIII P Valentini A Ca~sandro M VccchioltjmiddotAntll ldl C Bcrtoni G and Kuiper M (1 997 )IFLP mlJ-kcrs Ill DNA fingerpri nl ing ill Cltlu it Anima l Genetics 2~ 4 18 - 420

4 Bennett S C J John J R and nwi~Hn J W (1954) nillwl Iusbdildry III At-ric nll lm in the S lId ll1 [o ll1ill l D (td ) pr 6n - 667 ()x ro rd l Jni erStt) Prcs~ UK

i)

SlIn ] SCnn Metrol Vol 1() 2J07

5 Chung ER Kim WT Kim YS and nan SK (2000) DN middot fingerprintin g of Korenn cattle llsiug ArLP markers Korean JOllrnal of Animal Science 42 19 1 shy406

6 Jaccard P (l9()8) ~ NOllve lles recherches sur la dislribution flora Ie Buil -SOc Vauu Sci Nat 44 223 - 270

7 Jiang XP Liu GO Ding JT Yang LG Cau Sx and Cheng S() (2003) Diversity in six gont populatiolls in the middle and lower Yangtze River Vallcy Asian Australian Journal of Animal Sciences 16 277 - 2~ I

8 Joshi C G Rank D N Brahmakshtrl It P Palel A V Vataliya P II Muraleed haran P Khoda V K and Soiankii 1 V (1998) RJ PO analsys by PCR llsing arbitrary primers in Jifferent animal species Indian Vet 1 75 1029-1031

9 Joshi N R McLaughlin E A and Phillips R W (l1S7) Types Hnd breeds of African Cattle Agric Stud No 37 FAO Rome

10 Mason 1 L and Maule J P (]960) T he indi ge nous li vestock of Eastern and Southern Africa Conunon Bnr An ima l Breed Genet Tech Com-mun No 14 Commonwealth Agricultural Bureaux Farnham Royal UK

] 1 Mitra A Schlee P Krause I Blusch bull J Werner T Balakrishnan C R and Pirchner r (1998) Molec ular marker and their applications in livestock improvement Anim B iotech 9 81- 87

12 Morganle M Raralski J A lliddlc P Tingey S and Olivieri A M (1994) Genetic mapping and variability of seven soybean simple sequenee repea r loci Genome 37 763 - 769

]3 Net M and Lt W H (1979) Mathematical model for studying genetic variation in tenus of restriction endonnceases Proceedings of the al io nal Academy of Sciences US A 76 5269 - 5273

14 Payne W J A (1970) Caltle Prodllction in the Tropics Vo l I Breeds and Breeding Longman Gronp Ltd Lo ndon P 65

15 Promega Corporation (1999) WiLltlld GeIli)mic DNA purificltioll Kit Technica l Manual 2800 Woods Ho llow Road Matli so n US A

16 Ruhlf F J (1990) NTS S-P Numerical Taxonomy and Multivariate Analysis System Version 17 Owners Manual

17 Rouppe Van der VoorlJNAM Van Zandvoort P Van Eck if 01 Fulkerlsma R T Hutten R n c Draainrabull1 Gummers (1 bull 1 Jaco h~en E Helder bullJ and Baker J (1997) Use of allele specificity of comi grating AFLP markers to align genetic maps from different potato ge notypes Mol (l cn l icnet 255 438 shy477

Ill Rouse J (1972) World Cattle Il Cattle of Afriea and AsiR Okhlhoma Uni versity Press OkLlhoma USA

19 Vos Igt Rogers R Bleeker M Rcijms M Van de Lee T Uorncs M Fijlers A Pot 1 Peleman J Kuiper M and Z3ueau Me ( 1I95) AfLP a Hew

technique for DNA fingerprinting Nuc Acids Res 23 4407 - 4414

20 Zabeml M and Vos P (1993) Selective restrictioll fragment ampliiicalion a eeneral method for DNA fingcrprinllng EuropcMl Patent Application of 92402G 2SJ Pub

o 053485~A 1 bull

60

Page 4: » 1, · cluster anRlysis resolved tIm:" main clnsters wilh gcm:tic relationships of 53 %, 57'i{ and ()(i';( belweell l1 aggara (Niala wy) and Nilotic (Maioek): between Haggara (l'I,1Iawy)

Slid I Simi M elro Vol 1(1) 2007

DNA finger-printing of Korean cattle using AFLP markers was st udied by Chung 1lt)

and they reported that in the IIanw()o breed the numbel of N~LP bmlds per primer combination varied from 37 to 102 bands with an averag of 66 1 hallds using a 13 primer combination A tota l of 859 markers were generated ami 568 bands were polymorphic (66)- In Italian Holstei n dairy cows AFLP finger-p rin ting was used by Ajmone (3) They employed 16 primer com bi nations and generate d 1098 hands out of which 248 were polyrilOrphic averaging 155 bands pcr primcr combinat ion and ranging between 6 and 27 bmds Ajmone (2) ill vestigated variation aDOIlg Italian Friesian Italian Brown and Maremmana cattle and reported that 3J6 fragments were produced when they used AFLP to sludy lile polymorp l1isD md I J 1 of these fragment were polymorphic shy

l()()shy

90

80

7)

Amplicons 60

50

40

30

20

10

-r-

-- - -~I

r- I

F i r ~

Ill ~ i-

I ~

I

I

~

2111 2112 712 4 J 21M

-

----- 2115 6114 13m 7m

Primer combination

D Total No of llmplicolls

Polymorphic amplico[J

Fig 1 AFLP histogram showing the level of pnlymorphlsm as revealed

by each AFLP primer w mbination

56

Slid I Stnds Metro Vol 1(1) 2007

M 1 -----~ 9 9 1----shy

A B

Fig 2 Polyachrylamide gel showing AfLP profiles of four ocallypes Lane (1 2) Hagganl- Nia lawy (34) Nilulic - Majock (5 6) Bulana and (789) Kenana callie M

molecular weight slandard (lOU bp ladder) A primer- combination 23 B primer combination 24

2 Genetic similarity The breeder can use genetic similarity (Genetic distance = 100 - genetic

similarity) informaLion to make informed dec isions regarding the choice of genotypes to cross for the development of populations in order to maximize the expression of heterosis The genetic similariLies between pairs of type lines were calculiltcd using Jaccard s coefficient Pair wise similarities ranging from 46 9 to 803 with an average of 636 (Table 2) Maximum similarity of 8()Yc) was observed within Nilotic type Jines both share the same genetic background Nilotic (Majock) while the lowest genetic similarity of 46 9 was observed between Raggara and Kenana cattle which are different types from different locations

The mean genetic distance within Italian Friesian Italian Brown and Maremmana cattle studied by Aj mone (l J was 8YXi Jiang (7 ) showed that the genetic similarity within six autochthonus goat population s US ilig AFLP ranged from 0745 to 0 758 and 0951 to 0 970 between populations In Ital y Ajmone (2) investigated

57

Slid I Sinds Metro bull Iol 1([) 2007

the genelic variability in 4 goat breeds and r porld that with in hreed genetic diversity was 87 for the majori ty or breeds alld genetic similaritis hdweclI pltlirs of individuals of the same breeds averaged 070

Table 2 Genetic similarity matrix computed according to Jaccards cllcfficienl bmed on AFLP data between 9 lines of 4 local types lfiaggara - Nialawy Lines 1 and Nilutlcshy

Majock lines 3 and 4 Butana lines 5 and 6 Kenana callie breed lines 7 Sand 9)

Types Nialawy M aj ock I3 utana Kena na

1 2 3 4 5 6 7 8 9

Nialawy 1 I00 0 2 668 1000

Majock 3 651 598 1000 4 62 8 567 80 3 1000

Butana 5 603 56lt) 746 760 1000 6 54 1 520 62 1 65 1 76~ 1000

Kenana 7 527 544 559 5~ 5 5~ 2 52 6 1000 H 517 -469 _569 597 68 4 765 51 1 J 000 9 495 537 550 554 540 4lt) 2 63 470 100 0

3 GtIletic relationship

The dendrogram constructed from UPGMA cluster analysis betwe n [he local breeds based on AFLP dala resolved the 9 breed Iille~ into Ulree mail dusters a shown in Figure 3 The first cl uster with 053 genetic relationships is bctwecli Western I3aggara-N ialawy and NiloLic-Majock breeu Then the genctic relatiollship heLween Western Baggara and Nj jotic li nes are 067 amI 06 1 respectively The second cluster with 057 genetic re JaLi onship is between Western Haggara UUIula and Kenana breeus The genetic rela tionship between BUlltUla li lles is nRO The third cluster group wi th 066 genetic relaLionships is between DulanH emu KCIlltlnu catlle breed The genetic relationship betweclI line 9 8 and 7 is 072 wh ile between inc 8 and 7 of Kenana caltle breed the rd aLionship is 077 as lt110 vn in Figure 3

Fig3 Dendrogram of genelic relationship gcoclntcd om AFLP data constructed h) IJ1lGllA among 9 lines of 4 local types (Klnana nulana Nilotic - MaJock (N I) and Bag~ara - NialawyWN)

------------------~

1 ~ - t __~ I ~

M 1 Mshy

BN nN i3UI ltlIlltl

13 l1Lana

K nanltl

Kcnltlnltl

Kell am

Slid J ~11l1A Metrol Vlll I( 1) 2007

The genet ic relationltlbip~ he tween and witbin these loeal lypcs )f callIe wert IJIvesligated using duster analysis based )f) silllilarily JIlalricc~ of UPGMA and tllt dendrogram constructed resolved the 9 groups lines llllO three main cJuslcn as shown in Figure 3 The firs t with a )~ genetic relation hip hetween Baggara (Nialawy) am Nil otic (Mujock) Iype while Ihe genetic relationship iL)l1 Baggafa and Nilol ic types were 067 ltLillI 061 respecllvely The second cluster vith 0_7 genetic relationship was between Baggara (Nialawy) and Butana type whik tlle genetic relationship wi thin the Bulalla type was ORO The third luster group willI 066 gcn~tic rclaLioml1ip be tween I3utana and KClIana cat lie while [JIC range of genetilt relationship witllin KCllana caule was 072 10 077 hom t11i~ resull it is oh vious that the genetic relationship betweell KCllana and I3utana is the closest then foll oweu hy the genetic relationship hetween Baggara Kenalla I3utana Niloljc (Majo k) anu Buggma (Nialawy) me lllQre ui~tan t This result j ltonsislcllt Willl their geographic locallon and disuibutioll aou coinciues to some degree willl Illcir historical dassification mentiOlleu by Ma~oJl and Maule (iIll Rouse ( I ~ Payne (I)

anu Joshi ~) Joshi (8) esLimated phylogenetic relatjon~hip among [lanwou hincsc Yanbian unu foreign breeds They reponed thal the Hanwoo breed was Illlt~ most closely related to tbe Chinese Yanhian atlle followed by the lIereford and lIolstein hreeds

Ajmone (2) presented a d endrogram lmd concluded t11at t1le Italian Friesian and Italian Brown catlle were slightl y more closely related to each olher than tJley were to MaremmU1a caule Cluster analysis in six autochthonous goal populations ludieu hy Jiang et II (2003) using AfLP mdicaled t11al Chengdu Grey goal (eGG) is the most uislant popu lation while Chuunuong White goaL (CWO) and Yangtze Riwr Oel t~l White goat (ROWG) were IllC dosest popu lations followed hy Boujio gnat ( B(J ) Hui goat (HG)ltUld Malou goat (MCi)

References I AjmonemiddotMarsan II Negrinl R Milancsi E Bozzi R Nijman I 1 bullbull Ilunljcr J fl

Valentin A and LenstJ3 J A (211110) G ne ti c di stances wi tiJin and )Cross cat ile hrecus as illuitlt1 led b biallcl ic rLP lllatkcr Ani ma] Gelle(ics 33 2~O-2R6

2 Ajll1one-l1lttJsan P bull Otsen M bullbull NegJini R Vecchiulli-Anlald Cbull KUiper Mbull Len~lla JA and Piva C 0(99) Genetic diycrsily iLlIin an d betwlcn cIU le bneds a ~ measured wi th AFLP nlm)crs Recent 1 rore in limal production Science 1 Procttdi l1 ~ oj lbl AS]A XlU Congress PinCCII 7lt1 tt aly 21 - 2-l Jllllt

t 000 1 55 - 1 57

3 Ajmonc-lfanIII P Valentini A Ca~sandro M VccchioltjmiddotAntll ldl C Bcrtoni G and Kuiper M (1 997 )IFLP mlJ-kcrs Ill DNA fingerpri nl ing ill Cltlu it Anima l Genetics 2~ 4 18 - 420

4 Bennett S C J John J R and nwi~Hn J W (1954) nillwl Iusbdildry III At-ric nll lm in the S lId ll1 [o ll1ill l D (td ) pr 6n - 667 ()x ro rd l Jni erStt) Prcs~ UK

i)

SlIn ] SCnn Metrol Vol 1() 2J07

5 Chung ER Kim WT Kim YS and nan SK (2000) DN middot fingerprintin g of Korenn cattle llsiug ArLP markers Korean JOllrnal of Animal Science 42 19 1 shy406

6 Jaccard P (l9()8) ~ NOllve lles recherches sur la dislribution flora Ie Buil -SOc Vauu Sci Nat 44 223 - 270

7 Jiang XP Liu GO Ding JT Yang LG Cau Sx and Cheng S() (2003) Diversity in six gont populatiolls in the middle and lower Yangtze River Vallcy Asian Australian Journal of Animal Sciences 16 277 - 2~ I

8 Joshi C G Rank D N Brahmakshtrl It P Palel A V Vataliya P II Muraleed haran P Khoda V K and Soiankii 1 V (1998) RJ PO analsys by PCR llsing arbitrary primers in Jifferent animal species Indian Vet 1 75 1029-1031

9 Joshi N R McLaughlin E A and Phillips R W (l1S7) Types Hnd breeds of African Cattle Agric Stud No 37 FAO Rome

10 Mason 1 L and Maule J P (]960) T he indi ge nous li vestock of Eastern and Southern Africa Conunon Bnr An ima l Breed Genet Tech Com-mun No 14 Commonwealth Agricultural Bureaux Farnham Royal UK

] 1 Mitra A Schlee P Krause I Blusch bull J Werner T Balakrishnan C R and Pirchner r (1998) Molec ular marker and their applications in livestock improvement Anim B iotech 9 81- 87

12 Morganle M Raralski J A lliddlc P Tingey S and Olivieri A M (1994) Genetic mapping and variability of seven soybean simple sequenee repea r loci Genome 37 763 - 769

]3 Net M and Lt W H (1979) Mathematical model for studying genetic variation in tenus of restriction endonnceases Proceedings of the al io nal Academy of Sciences US A 76 5269 - 5273

14 Payne W J A (1970) Caltle Prodllction in the Tropics Vo l I Breeds and Breeding Longman Gronp Ltd Lo ndon P 65

15 Promega Corporation (1999) WiLltlld GeIli)mic DNA purificltioll Kit Technica l Manual 2800 Woods Ho llow Road Matli so n US A

16 Ruhlf F J (1990) NTS S-P Numerical Taxonomy and Multivariate Analysis System Version 17 Owners Manual

17 Rouppe Van der VoorlJNAM Van Zandvoort P Van Eck if 01 Fulkerlsma R T Hutten R n c Draainrabull1 Gummers (1 bull 1 Jaco h~en E Helder bullJ and Baker J (1997) Use of allele specificity of comi grating AFLP markers to align genetic maps from different potato ge notypes Mol (l cn l icnet 255 438 shy477

Ill Rouse J (1972) World Cattle Il Cattle of Afriea and AsiR Okhlhoma Uni versity Press OkLlhoma USA

19 Vos Igt Rogers R Bleeker M Rcijms M Van de Lee T Uorncs M Fijlers A Pot 1 Peleman J Kuiper M and Z3ueau Me ( 1I95) AfLP a Hew

technique for DNA fingerprinting Nuc Acids Res 23 4407 - 4414

20 Zabeml M and Vos P (1993) Selective restrictioll fragment ampliiicalion a eeneral method for DNA fingcrprinllng EuropcMl Patent Application of 92402G 2SJ Pub

o 053485~A 1 bull

60

Page 5: » 1, · cluster anRlysis resolved tIm:" main clnsters wilh gcm:tic relationships of 53 %, 57'i{ and ()(i';( belweell l1 aggara (Niala wy) and Nilotic (Maioek): between Haggara (l'I,1Iawy)

Slid I Stnds Metro Vol 1(1) 2007

M 1 -----~ 9 9 1----shy

A B

Fig 2 Polyachrylamide gel showing AfLP profiles of four ocallypes Lane (1 2) Hagganl- Nia lawy (34) Nilulic - Majock (5 6) Bulana and (789) Kenana callie M

molecular weight slandard (lOU bp ladder) A primer- combination 23 B primer combination 24

2 Genetic similarity The breeder can use genetic similarity (Genetic distance = 100 - genetic

similarity) informaLion to make informed dec isions regarding the choice of genotypes to cross for the development of populations in order to maximize the expression of heterosis The genetic similariLies between pairs of type lines were calculiltcd using Jaccard s coefficient Pair wise similarities ranging from 46 9 to 803 with an average of 636 (Table 2) Maximum similarity of 8()Yc) was observed within Nilotic type Jines both share the same genetic background Nilotic (Majock) while the lowest genetic similarity of 46 9 was observed between Raggara and Kenana cattle which are different types from different locations

The mean genetic distance within Italian Friesian Italian Brown and Maremmana cattle studied by Aj mone (l J was 8YXi Jiang (7 ) showed that the genetic similarity within six autochthonus goat population s US ilig AFLP ranged from 0745 to 0 758 and 0951 to 0 970 between populations In Ital y Ajmone (2) investigated

57

Slid I Sinds Metro bull Iol 1([) 2007

the genelic variability in 4 goat breeds and r porld that with in hreed genetic diversity was 87 for the majori ty or breeds alld genetic similaritis hdweclI pltlirs of individuals of the same breeds averaged 070

Table 2 Genetic similarity matrix computed according to Jaccards cllcfficienl bmed on AFLP data between 9 lines of 4 local types lfiaggara - Nialawy Lines 1 and Nilutlcshy

Majock lines 3 and 4 Butana lines 5 and 6 Kenana callie breed lines 7 Sand 9)

Types Nialawy M aj ock I3 utana Kena na

1 2 3 4 5 6 7 8 9

Nialawy 1 I00 0 2 668 1000

Majock 3 651 598 1000 4 62 8 567 80 3 1000

Butana 5 603 56lt) 746 760 1000 6 54 1 520 62 1 65 1 76~ 1000

Kenana 7 527 544 559 5~ 5 5~ 2 52 6 1000 H 517 -469 _569 597 68 4 765 51 1 J 000 9 495 537 550 554 540 4lt) 2 63 470 100 0

3 GtIletic relationship

The dendrogram constructed from UPGMA cluster analysis betwe n [he local breeds based on AFLP dala resolved the 9 breed Iille~ into Ulree mail dusters a shown in Figure 3 The first cl uster with 053 genetic relationships is bctwecli Western I3aggara-N ialawy and NiloLic-Majock breeu Then the genctic relatiollship heLween Western Baggara and Nj jotic li nes are 067 amI 06 1 respectively The second cluster with 057 genetic re JaLi onship is between Western Haggara UUIula and Kenana breeus The genetic rela tionship between BUlltUla li lles is nRO The third cluster group wi th 066 genetic relaLionships is between DulanH emu KCIlltlnu catlle breed The genetic relationship betweclI line 9 8 and 7 is 072 wh ile between inc 8 and 7 of Kenana caltle breed the rd aLionship is 077 as lt110 vn in Figure 3

Fig3 Dendrogram of genelic relationship gcoclntcd om AFLP data constructed h) IJ1lGllA among 9 lines of 4 local types (Klnana nulana Nilotic - MaJock (N I) and Bag~ara - NialawyWN)

------------------~

1 ~ - t __~ I ~

M 1 Mshy

BN nN i3UI ltlIlltl

13 l1Lana

K nanltl

Kcnltlnltl

Kell am

Slid J ~11l1A Metrol Vlll I( 1) 2007

The genet ic relationltlbip~ he tween and witbin these loeal lypcs )f callIe wert IJIvesligated using duster analysis based )f) silllilarily JIlalricc~ of UPGMA and tllt dendrogram constructed resolved the 9 groups lines llllO three main cJuslcn as shown in Figure 3 The firs t with a )~ genetic relation hip hetween Baggara (Nialawy) am Nil otic (Mujock) Iype while Ihe genetic relationship iL)l1 Baggafa and Nilol ic types were 067 ltLillI 061 respecllvely The second cluster vith 0_7 genetic relationship was between Baggara (Nialawy) and Butana type whik tlle genetic relationship wi thin the Bulalla type was ORO The third luster group willI 066 gcn~tic rclaLioml1ip be tween I3utana and KClIana cat lie while [JIC range of genetilt relationship witllin KCllana caule was 072 10 077 hom t11i~ resull it is oh vious that the genetic relationship betweell KCllana and I3utana is the closest then foll oweu hy the genetic relationship hetween Baggara Kenalla I3utana Niloljc (Majo k) anu Buggma (Nialawy) me lllQre ui~tan t This result j ltonsislcllt Willl their geographic locallon and disuibutioll aou coinciues to some degree willl Illcir historical dassification mentiOlleu by Ma~oJl and Maule (iIll Rouse ( I ~ Payne (I)

anu Joshi ~) Joshi (8) esLimated phylogenetic relatjon~hip among [lanwou hincsc Yanbian unu foreign breeds They reponed thal the Hanwoo breed was Illlt~ most closely related to tbe Chinese Yanhian atlle followed by the lIereford and lIolstein hreeds

Ajmone (2) presented a d endrogram lmd concluded t11at t1le Italian Friesian and Italian Brown catlle were slightl y more closely related to each olher than tJley were to MaremmU1a caule Cluster analysis in six autochthonous goal populations ludieu hy Jiang et II (2003) using AfLP mdicaled t11al Chengdu Grey goal (eGG) is the most uislant popu lation while Chuunuong White goaL (CWO) and Yangtze Riwr Oel t~l White goat (ROWG) were IllC dosest popu lations followed hy Boujio gnat ( B(J ) Hui goat (HG)ltUld Malou goat (MCi)

References I AjmonemiddotMarsan II Negrinl R Milancsi E Bozzi R Nijman I 1 bullbull Ilunljcr J fl

Valentin A and LenstJ3 J A (211110) G ne ti c di stances wi tiJin and )Cross cat ile hrecus as illuitlt1 led b biallcl ic rLP lllatkcr Ani ma] Gelle(ics 33 2~O-2R6

2 Ajll1one-l1lttJsan P bull Otsen M bullbull NegJini R Vecchiulli-Anlald Cbull KUiper Mbull Len~lla JA and Piva C 0(99) Genetic diycrsily iLlIin an d betwlcn cIU le bneds a ~ measured wi th AFLP nlm)crs Recent 1 rore in limal production Science 1 Procttdi l1 ~ oj lbl AS]A XlU Congress PinCCII 7lt1 tt aly 21 - 2-l Jllllt

t 000 1 55 - 1 57

3 Ajmonc-lfanIII P Valentini A Ca~sandro M VccchioltjmiddotAntll ldl C Bcrtoni G and Kuiper M (1 997 )IFLP mlJ-kcrs Ill DNA fingerpri nl ing ill Cltlu it Anima l Genetics 2~ 4 18 - 420

4 Bennett S C J John J R and nwi~Hn J W (1954) nillwl Iusbdildry III At-ric nll lm in the S lId ll1 [o ll1ill l D (td ) pr 6n - 667 ()x ro rd l Jni erStt) Prcs~ UK

i)

SlIn ] SCnn Metrol Vol 1() 2J07

5 Chung ER Kim WT Kim YS and nan SK (2000) DN middot fingerprintin g of Korenn cattle llsiug ArLP markers Korean JOllrnal of Animal Science 42 19 1 shy406

6 Jaccard P (l9()8) ~ NOllve lles recherches sur la dislribution flora Ie Buil -SOc Vauu Sci Nat 44 223 - 270

7 Jiang XP Liu GO Ding JT Yang LG Cau Sx and Cheng S() (2003) Diversity in six gont populatiolls in the middle and lower Yangtze River Vallcy Asian Australian Journal of Animal Sciences 16 277 - 2~ I

8 Joshi C G Rank D N Brahmakshtrl It P Palel A V Vataliya P II Muraleed haran P Khoda V K and Soiankii 1 V (1998) RJ PO analsys by PCR llsing arbitrary primers in Jifferent animal species Indian Vet 1 75 1029-1031

9 Joshi N R McLaughlin E A and Phillips R W (l1S7) Types Hnd breeds of African Cattle Agric Stud No 37 FAO Rome

10 Mason 1 L and Maule J P (]960) T he indi ge nous li vestock of Eastern and Southern Africa Conunon Bnr An ima l Breed Genet Tech Com-mun No 14 Commonwealth Agricultural Bureaux Farnham Royal UK

] 1 Mitra A Schlee P Krause I Blusch bull J Werner T Balakrishnan C R and Pirchner r (1998) Molec ular marker and their applications in livestock improvement Anim B iotech 9 81- 87

12 Morganle M Raralski J A lliddlc P Tingey S and Olivieri A M (1994) Genetic mapping and variability of seven soybean simple sequenee repea r loci Genome 37 763 - 769

]3 Net M and Lt W H (1979) Mathematical model for studying genetic variation in tenus of restriction endonnceases Proceedings of the al io nal Academy of Sciences US A 76 5269 - 5273

14 Payne W J A (1970) Caltle Prodllction in the Tropics Vo l I Breeds and Breeding Longman Gronp Ltd Lo ndon P 65

15 Promega Corporation (1999) WiLltlld GeIli)mic DNA purificltioll Kit Technica l Manual 2800 Woods Ho llow Road Matli so n US A

16 Ruhlf F J (1990) NTS S-P Numerical Taxonomy and Multivariate Analysis System Version 17 Owners Manual

17 Rouppe Van der VoorlJNAM Van Zandvoort P Van Eck if 01 Fulkerlsma R T Hutten R n c Draainrabull1 Gummers (1 bull 1 Jaco h~en E Helder bullJ and Baker J (1997) Use of allele specificity of comi grating AFLP markers to align genetic maps from different potato ge notypes Mol (l cn l icnet 255 438 shy477

Ill Rouse J (1972) World Cattle Il Cattle of Afriea and AsiR Okhlhoma Uni versity Press OkLlhoma USA

19 Vos Igt Rogers R Bleeker M Rcijms M Van de Lee T Uorncs M Fijlers A Pot 1 Peleman J Kuiper M and Z3ueau Me ( 1I95) AfLP a Hew

technique for DNA fingerprinting Nuc Acids Res 23 4407 - 4414

20 Zabeml M and Vos P (1993) Selective restrictioll fragment ampliiicalion a eeneral method for DNA fingcrprinllng EuropcMl Patent Application of 92402G 2SJ Pub

o 053485~A 1 bull

60

Page 6: » 1, · cluster anRlysis resolved tIm:" main clnsters wilh gcm:tic relationships of 53 %, 57'i{ and ()(i';( belweell l1 aggara (Niala wy) and Nilotic (Maioek): between Haggara (l'I,1Iawy)

Slid I Sinds Metro bull Iol 1([) 2007

the genelic variability in 4 goat breeds and r porld that with in hreed genetic diversity was 87 for the majori ty or breeds alld genetic similaritis hdweclI pltlirs of individuals of the same breeds averaged 070

Table 2 Genetic similarity matrix computed according to Jaccards cllcfficienl bmed on AFLP data between 9 lines of 4 local types lfiaggara - Nialawy Lines 1 and Nilutlcshy

Majock lines 3 and 4 Butana lines 5 and 6 Kenana callie breed lines 7 Sand 9)

Types Nialawy M aj ock I3 utana Kena na

1 2 3 4 5 6 7 8 9

Nialawy 1 I00 0 2 668 1000

Majock 3 651 598 1000 4 62 8 567 80 3 1000

Butana 5 603 56lt) 746 760 1000 6 54 1 520 62 1 65 1 76~ 1000

Kenana 7 527 544 559 5~ 5 5~ 2 52 6 1000 H 517 -469 _569 597 68 4 765 51 1 J 000 9 495 537 550 554 540 4lt) 2 63 470 100 0

3 GtIletic relationship

The dendrogram constructed from UPGMA cluster analysis betwe n [he local breeds based on AFLP dala resolved the 9 breed Iille~ into Ulree mail dusters a shown in Figure 3 The first cl uster with 053 genetic relationships is bctwecli Western I3aggara-N ialawy and NiloLic-Majock breeu Then the genctic relatiollship heLween Western Baggara and Nj jotic li nes are 067 amI 06 1 respectively The second cluster with 057 genetic re JaLi onship is between Western Haggara UUIula and Kenana breeus The genetic rela tionship between BUlltUla li lles is nRO The third cluster group wi th 066 genetic relaLionships is between DulanH emu KCIlltlnu catlle breed The genetic relationship betweclI line 9 8 and 7 is 072 wh ile between inc 8 and 7 of Kenana caltle breed the rd aLionship is 077 as lt110 vn in Figure 3

Fig3 Dendrogram of genelic relationship gcoclntcd om AFLP data constructed h) IJ1lGllA among 9 lines of 4 local types (Klnana nulana Nilotic - MaJock (N I) and Bag~ara - NialawyWN)

------------------~

1 ~ - t __~ I ~

M 1 Mshy

BN nN i3UI ltlIlltl

13 l1Lana

K nanltl

Kcnltlnltl

Kell am

Slid J ~11l1A Metrol Vlll I( 1) 2007

The genet ic relationltlbip~ he tween and witbin these loeal lypcs )f callIe wert IJIvesligated using duster analysis based )f) silllilarily JIlalricc~ of UPGMA and tllt dendrogram constructed resolved the 9 groups lines llllO three main cJuslcn as shown in Figure 3 The firs t with a )~ genetic relation hip hetween Baggara (Nialawy) am Nil otic (Mujock) Iype while Ihe genetic relationship iL)l1 Baggafa and Nilol ic types were 067 ltLillI 061 respecllvely The second cluster vith 0_7 genetic relationship was between Baggara (Nialawy) and Butana type whik tlle genetic relationship wi thin the Bulalla type was ORO The third luster group willI 066 gcn~tic rclaLioml1ip be tween I3utana and KClIana cat lie while [JIC range of genetilt relationship witllin KCllana caule was 072 10 077 hom t11i~ resull it is oh vious that the genetic relationship betweell KCllana and I3utana is the closest then foll oweu hy the genetic relationship hetween Baggara Kenalla I3utana Niloljc (Majo k) anu Buggma (Nialawy) me lllQre ui~tan t This result j ltonsislcllt Willl their geographic locallon and disuibutioll aou coinciues to some degree willl Illcir historical dassification mentiOlleu by Ma~oJl and Maule (iIll Rouse ( I ~ Payne (I)

anu Joshi ~) Joshi (8) esLimated phylogenetic relatjon~hip among [lanwou hincsc Yanbian unu foreign breeds They reponed thal the Hanwoo breed was Illlt~ most closely related to tbe Chinese Yanhian atlle followed by the lIereford and lIolstein hreeds

Ajmone (2) presented a d endrogram lmd concluded t11at t1le Italian Friesian and Italian Brown catlle were slightl y more closely related to each olher than tJley were to MaremmU1a caule Cluster analysis in six autochthonous goal populations ludieu hy Jiang et II (2003) using AfLP mdicaled t11al Chengdu Grey goal (eGG) is the most uislant popu lation while Chuunuong White goaL (CWO) and Yangtze Riwr Oel t~l White goat (ROWG) were IllC dosest popu lations followed hy Boujio gnat ( B(J ) Hui goat (HG)ltUld Malou goat (MCi)

References I AjmonemiddotMarsan II Negrinl R Milancsi E Bozzi R Nijman I 1 bullbull Ilunljcr J fl

Valentin A and LenstJ3 J A (211110) G ne ti c di stances wi tiJin and )Cross cat ile hrecus as illuitlt1 led b biallcl ic rLP lllatkcr Ani ma] Gelle(ics 33 2~O-2R6

2 Ajll1one-l1lttJsan P bull Otsen M bullbull NegJini R Vecchiulli-Anlald Cbull KUiper Mbull Len~lla JA and Piva C 0(99) Genetic diycrsily iLlIin an d betwlcn cIU le bneds a ~ measured wi th AFLP nlm)crs Recent 1 rore in limal production Science 1 Procttdi l1 ~ oj lbl AS]A XlU Congress PinCCII 7lt1 tt aly 21 - 2-l Jllllt

t 000 1 55 - 1 57

3 Ajmonc-lfanIII P Valentini A Ca~sandro M VccchioltjmiddotAntll ldl C Bcrtoni G and Kuiper M (1 997 )IFLP mlJ-kcrs Ill DNA fingerpri nl ing ill Cltlu it Anima l Genetics 2~ 4 18 - 420

4 Bennett S C J John J R and nwi~Hn J W (1954) nillwl Iusbdildry III At-ric nll lm in the S lId ll1 [o ll1ill l D (td ) pr 6n - 667 ()x ro rd l Jni erStt) Prcs~ UK

i)

SlIn ] SCnn Metrol Vol 1() 2J07

5 Chung ER Kim WT Kim YS and nan SK (2000) DN middot fingerprintin g of Korenn cattle llsiug ArLP markers Korean JOllrnal of Animal Science 42 19 1 shy406

6 Jaccard P (l9()8) ~ NOllve lles recherches sur la dislribution flora Ie Buil -SOc Vauu Sci Nat 44 223 - 270

7 Jiang XP Liu GO Ding JT Yang LG Cau Sx and Cheng S() (2003) Diversity in six gont populatiolls in the middle and lower Yangtze River Vallcy Asian Australian Journal of Animal Sciences 16 277 - 2~ I

8 Joshi C G Rank D N Brahmakshtrl It P Palel A V Vataliya P II Muraleed haran P Khoda V K and Soiankii 1 V (1998) RJ PO analsys by PCR llsing arbitrary primers in Jifferent animal species Indian Vet 1 75 1029-1031

9 Joshi N R McLaughlin E A and Phillips R W (l1S7) Types Hnd breeds of African Cattle Agric Stud No 37 FAO Rome

10 Mason 1 L and Maule J P (]960) T he indi ge nous li vestock of Eastern and Southern Africa Conunon Bnr An ima l Breed Genet Tech Com-mun No 14 Commonwealth Agricultural Bureaux Farnham Royal UK

] 1 Mitra A Schlee P Krause I Blusch bull J Werner T Balakrishnan C R and Pirchner r (1998) Molec ular marker and their applications in livestock improvement Anim B iotech 9 81- 87

12 Morganle M Raralski J A lliddlc P Tingey S and Olivieri A M (1994) Genetic mapping and variability of seven soybean simple sequenee repea r loci Genome 37 763 - 769

]3 Net M and Lt W H (1979) Mathematical model for studying genetic variation in tenus of restriction endonnceases Proceedings of the al io nal Academy of Sciences US A 76 5269 - 5273

14 Payne W J A (1970) Caltle Prodllction in the Tropics Vo l I Breeds and Breeding Longman Gronp Ltd Lo ndon P 65

15 Promega Corporation (1999) WiLltlld GeIli)mic DNA purificltioll Kit Technica l Manual 2800 Woods Ho llow Road Matli so n US A

16 Ruhlf F J (1990) NTS S-P Numerical Taxonomy and Multivariate Analysis System Version 17 Owners Manual

17 Rouppe Van der VoorlJNAM Van Zandvoort P Van Eck if 01 Fulkerlsma R T Hutten R n c Draainrabull1 Gummers (1 bull 1 Jaco h~en E Helder bullJ and Baker J (1997) Use of allele specificity of comi grating AFLP markers to align genetic maps from different potato ge notypes Mol (l cn l icnet 255 438 shy477

Ill Rouse J (1972) World Cattle Il Cattle of Afriea and AsiR Okhlhoma Uni versity Press OkLlhoma USA

19 Vos Igt Rogers R Bleeker M Rcijms M Van de Lee T Uorncs M Fijlers A Pot 1 Peleman J Kuiper M and Z3ueau Me ( 1I95) AfLP a Hew

technique for DNA fingerprinting Nuc Acids Res 23 4407 - 4414

20 Zabeml M and Vos P (1993) Selective restrictioll fragment ampliiicalion a eeneral method for DNA fingcrprinllng EuropcMl Patent Application of 92402G 2SJ Pub

o 053485~A 1 bull

60

Page 7: » 1, · cluster anRlysis resolved tIm:" main clnsters wilh gcm:tic relationships of 53 %, 57'i{ and ()(i';( belweell l1 aggara (Niala wy) and Nilotic (Maioek): between Haggara (l'I,1Iawy)

Slid J ~11l1A Metrol Vlll I( 1) 2007

The genet ic relationltlbip~ he tween and witbin these loeal lypcs )f callIe wert IJIvesligated using duster analysis based )f) silllilarily JIlalricc~ of UPGMA and tllt dendrogram constructed resolved the 9 groups lines llllO three main cJuslcn as shown in Figure 3 The firs t with a )~ genetic relation hip hetween Baggara (Nialawy) am Nil otic (Mujock) Iype while Ihe genetic relationship iL)l1 Baggafa and Nilol ic types were 067 ltLillI 061 respecllvely The second cluster vith 0_7 genetic relationship was between Baggara (Nialawy) and Butana type whik tlle genetic relationship wi thin the Bulalla type was ORO The third luster group willI 066 gcn~tic rclaLioml1ip be tween I3utana and KClIana cat lie while [JIC range of genetilt relationship witllin KCllana caule was 072 10 077 hom t11i~ resull it is oh vious that the genetic relationship betweell KCllana and I3utana is the closest then foll oweu hy the genetic relationship hetween Baggara Kenalla I3utana Niloljc (Majo k) anu Buggma (Nialawy) me lllQre ui~tan t This result j ltonsislcllt Willl their geographic locallon and disuibutioll aou coinciues to some degree willl Illcir historical dassification mentiOlleu by Ma~oJl and Maule (iIll Rouse ( I ~ Payne (I)

anu Joshi ~) Joshi (8) esLimated phylogenetic relatjon~hip among [lanwou hincsc Yanbian unu foreign breeds They reponed thal the Hanwoo breed was Illlt~ most closely related to tbe Chinese Yanhian atlle followed by the lIereford and lIolstein hreeds

Ajmone (2) presented a d endrogram lmd concluded t11at t1le Italian Friesian and Italian Brown catlle were slightl y more closely related to each olher than tJley were to MaremmU1a caule Cluster analysis in six autochthonous goal populations ludieu hy Jiang et II (2003) using AfLP mdicaled t11al Chengdu Grey goal (eGG) is the most uislant popu lation while Chuunuong White goaL (CWO) and Yangtze Riwr Oel t~l White goat (ROWG) were IllC dosest popu lations followed hy Boujio gnat ( B(J ) Hui goat (HG)ltUld Malou goat (MCi)

References I AjmonemiddotMarsan II Negrinl R Milancsi E Bozzi R Nijman I 1 bullbull Ilunljcr J fl

Valentin A and LenstJ3 J A (211110) G ne ti c di stances wi tiJin and )Cross cat ile hrecus as illuitlt1 led b biallcl ic rLP lllatkcr Ani ma] Gelle(ics 33 2~O-2R6

2 Ajll1one-l1lttJsan P bull Otsen M bullbull NegJini R Vecchiulli-Anlald Cbull KUiper Mbull Len~lla JA and Piva C 0(99) Genetic diycrsily iLlIin an d betwlcn cIU le bneds a ~ measured wi th AFLP nlm)crs Recent 1 rore in limal production Science 1 Procttdi l1 ~ oj lbl AS]A XlU Congress PinCCII 7lt1 tt aly 21 - 2-l Jllllt

t 000 1 55 - 1 57

3 Ajmonc-lfanIII P Valentini A Ca~sandro M VccchioltjmiddotAntll ldl C Bcrtoni G and Kuiper M (1 997 )IFLP mlJ-kcrs Ill DNA fingerpri nl ing ill Cltlu it Anima l Genetics 2~ 4 18 - 420

4 Bennett S C J John J R and nwi~Hn J W (1954) nillwl Iusbdildry III At-ric nll lm in the S lId ll1 [o ll1ill l D (td ) pr 6n - 667 ()x ro rd l Jni erStt) Prcs~ UK

i)

SlIn ] SCnn Metrol Vol 1() 2J07

5 Chung ER Kim WT Kim YS and nan SK (2000) DN middot fingerprintin g of Korenn cattle llsiug ArLP markers Korean JOllrnal of Animal Science 42 19 1 shy406

6 Jaccard P (l9()8) ~ NOllve lles recherches sur la dislribution flora Ie Buil -SOc Vauu Sci Nat 44 223 - 270

7 Jiang XP Liu GO Ding JT Yang LG Cau Sx and Cheng S() (2003) Diversity in six gont populatiolls in the middle and lower Yangtze River Vallcy Asian Australian Journal of Animal Sciences 16 277 - 2~ I

8 Joshi C G Rank D N Brahmakshtrl It P Palel A V Vataliya P II Muraleed haran P Khoda V K and Soiankii 1 V (1998) RJ PO analsys by PCR llsing arbitrary primers in Jifferent animal species Indian Vet 1 75 1029-1031

9 Joshi N R McLaughlin E A and Phillips R W (l1S7) Types Hnd breeds of African Cattle Agric Stud No 37 FAO Rome

10 Mason 1 L and Maule J P (]960) T he indi ge nous li vestock of Eastern and Southern Africa Conunon Bnr An ima l Breed Genet Tech Com-mun No 14 Commonwealth Agricultural Bureaux Farnham Royal UK

] 1 Mitra A Schlee P Krause I Blusch bull J Werner T Balakrishnan C R and Pirchner r (1998) Molec ular marker and their applications in livestock improvement Anim B iotech 9 81- 87

12 Morganle M Raralski J A lliddlc P Tingey S and Olivieri A M (1994) Genetic mapping and variability of seven soybean simple sequenee repea r loci Genome 37 763 - 769

]3 Net M and Lt W H (1979) Mathematical model for studying genetic variation in tenus of restriction endonnceases Proceedings of the al io nal Academy of Sciences US A 76 5269 - 5273

14 Payne W J A (1970) Caltle Prodllction in the Tropics Vo l I Breeds and Breeding Longman Gronp Ltd Lo ndon P 65

15 Promega Corporation (1999) WiLltlld GeIli)mic DNA purificltioll Kit Technica l Manual 2800 Woods Ho llow Road Matli so n US A

16 Ruhlf F J (1990) NTS S-P Numerical Taxonomy and Multivariate Analysis System Version 17 Owners Manual

17 Rouppe Van der VoorlJNAM Van Zandvoort P Van Eck if 01 Fulkerlsma R T Hutten R n c Draainrabull1 Gummers (1 bull 1 Jaco h~en E Helder bullJ and Baker J (1997) Use of allele specificity of comi grating AFLP markers to align genetic maps from different potato ge notypes Mol (l cn l icnet 255 438 shy477

Ill Rouse J (1972) World Cattle Il Cattle of Afriea and AsiR Okhlhoma Uni versity Press OkLlhoma USA

19 Vos Igt Rogers R Bleeker M Rcijms M Van de Lee T Uorncs M Fijlers A Pot 1 Peleman J Kuiper M and Z3ueau Me ( 1I95) AfLP a Hew

technique for DNA fingerprinting Nuc Acids Res 23 4407 - 4414

20 Zabeml M and Vos P (1993) Selective restrictioll fragment ampliiicalion a eeneral method for DNA fingcrprinllng EuropcMl Patent Application of 92402G 2SJ Pub

o 053485~A 1 bull

60

Page 8: » 1, · cluster anRlysis resolved tIm:" main clnsters wilh gcm:tic relationships of 53 %, 57'i{ and ()(i';( belweell l1 aggara (Niala wy) and Nilotic (Maioek): between Haggara (l'I,1Iawy)

SlIn ] SCnn Metrol Vol 1() 2J07

5 Chung ER Kim WT Kim YS and nan SK (2000) DN middot fingerprintin g of Korenn cattle llsiug ArLP markers Korean JOllrnal of Animal Science 42 19 1 shy406

6 Jaccard P (l9()8) ~ NOllve lles recherches sur la dislribution flora Ie Buil -SOc Vauu Sci Nat 44 223 - 270

7 Jiang XP Liu GO Ding JT Yang LG Cau Sx and Cheng S() (2003) Diversity in six gont populatiolls in the middle and lower Yangtze River Vallcy Asian Australian Journal of Animal Sciences 16 277 - 2~ I

8 Joshi C G Rank D N Brahmakshtrl It P Palel A V Vataliya P II Muraleed haran P Khoda V K and Soiankii 1 V (1998) RJ PO analsys by PCR llsing arbitrary primers in Jifferent animal species Indian Vet 1 75 1029-1031

9 Joshi N R McLaughlin E A and Phillips R W (l1S7) Types Hnd breeds of African Cattle Agric Stud No 37 FAO Rome

10 Mason 1 L and Maule J P (]960) T he indi ge nous li vestock of Eastern and Southern Africa Conunon Bnr An ima l Breed Genet Tech Com-mun No 14 Commonwealth Agricultural Bureaux Farnham Royal UK

] 1 Mitra A Schlee P Krause I Blusch bull J Werner T Balakrishnan C R and Pirchner r (1998) Molec ular marker and their applications in livestock improvement Anim B iotech 9 81- 87

12 Morganle M Raralski J A lliddlc P Tingey S and Olivieri A M (1994) Genetic mapping and variability of seven soybean simple sequenee repea r loci Genome 37 763 - 769

]3 Net M and Lt W H (1979) Mathematical model for studying genetic variation in tenus of restriction endonnceases Proceedings of the al io nal Academy of Sciences US A 76 5269 - 5273

14 Payne W J A (1970) Caltle Prodllction in the Tropics Vo l I Breeds and Breeding Longman Gronp Ltd Lo ndon P 65

15 Promega Corporation (1999) WiLltlld GeIli)mic DNA purificltioll Kit Technica l Manual 2800 Woods Ho llow Road Matli so n US A

16 Ruhlf F J (1990) NTS S-P Numerical Taxonomy and Multivariate Analysis System Version 17 Owners Manual

17 Rouppe Van der VoorlJNAM Van Zandvoort P Van Eck if 01 Fulkerlsma R T Hutten R n c Draainrabull1 Gummers (1 bull 1 Jaco h~en E Helder bullJ and Baker J (1997) Use of allele specificity of comi grating AFLP markers to align genetic maps from different potato ge notypes Mol (l cn l icnet 255 438 shy477

Ill Rouse J (1972) World Cattle Il Cattle of Afriea and AsiR Okhlhoma Uni versity Press OkLlhoma USA

19 Vos Igt Rogers R Bleeker M Rcijms M Van de Lee T Uorncs M Fijlers A Pot 1 Peleman J Kuiper M and Z3ueau Me ( 1I95) AfLP a Hew

technique for DNA fingerprinting Nuc Acids Res 23 4407 - 4414

20 Zabeml M and Vos P (1993) Selective restrictioll fragment ampliiicalion a eeneral method for DNA fingcrprinllng EuropcMl Patent Application of 92402G 2SJ Pub

o 053485~A 1 bull

60