THE INHERITANCE OF HORNS IN . SOME FUt~THE:~ DATA.

B~z A. D. BUCHANAN SMITH. A~d,mal B~'eedin9 Resec~'eh Depa~'t'men4 Unive,rsi~y of Edinbu~9 h.

(With One Plate and Four Text-figures.)

Wm~N tile writer :first :heard of certain exceptions to the well-estab- lished rule of simple dominance of the polled condition in earle, he paid littIe attention to them. Further records of such exceptions, however, were encountered, and it was then felt that the subject merited fuller consideration. The following report concerning such exceptions came from two main smtrees: from Northern Rhodesia, where the crosses concerned were Aberdeen-Angus bulls and native cows, and from England, where Crosses involving the Park Cattle, between their two types and various other breeds, appeared to give anomalous resuRs.

A. RttO.DES:[ANNATIVE CATTLE. For precise information regarding the case in Northern Rhodesia, the writer is indebted to N_r R. A. S. ~Iacdonald, M.R.C.V.S., of the Veterinary Experimental Station, Chilanga. His facts relate mainly go two cases, and are as follows: 1..In a herd of Angoni and hfashukulumbwe cows, the majority of which were pure native bred (though a few showed slight traces of Hereford or blood, bug none had any Aberdeen-Angus blood), a p~e bred registered Aberdeen-Angus polled bull was placed (see photograph). The cows were alI horned, ani[ in colour were red, red- and-white or dun. There were practically no blacks. FiRy-one of g.heir progeny so produced were traced as under: 27 heifers all compleSely polled. 9 bulls all completely polled. 15 bulls mostly with distinct horns or buds. All were completely black with the exception of a few white ~aces. This was the case in 120 such calves. 2. In another herd, a pure bred registered Aberdeen-Angus btdl (polled) was mated to native cows, all of which possessed the typical 3(16 [-[or~8 in Cattle heavy horn. In this ease, all f;he progeny were completely black, practi- cally all the heifers polled, and practically all the males with buds, short drooping horns, or ordinary short horns. It was found diNcuR to obtain more detailed information as regards exact munbers, and rather than obtain figtu'es which could not be guaranteed to be correct, it was considered advisable merely to give the bare outline of facts. The following conclusion concerning these two herds, which are typical of several others, may, i~herefore, be con- sidered indubitable and has been confirmed by correspondence: That a registered Aberdeen-Angus bull (polled) when mated to the native cows of Northern iRhodesia (horned) produced calves, all the heifers being polled and withou~ scars, and the majority of the bull calves possessing horns of various shapes, .a few being only scurred, and a very few being polled. As the cattle of Northern l~hodesia are being rapidly "graded" up, it is tmlikely that the native breed will retain their purity much longer. Andsince it seemed improbable that more exact facts were likely to be obtained, or that the reciprocal cross was likely to be made, it is con- sidered advisable to record these facts as they stand. The conclusion, therefore, is that in rantings with the indigenous cattle of Africa, horns are inherited as a sex-Iimited character; that ~s to say that the factors for their expression, are either dominant in the male and recessive in the femaIe, or otherwise governed by sex. Sinces in the ordinary domesticated breeds of cattle, the polled condition behaves as a simple dominant to the horned, and since there was ~ome blood of domesticated breeds in certain of the native cows used in this cross, it would appear logical to presmne that either the sex-limited transmission of horns, or the simple dominance of the polled condition, is the rule in cattle, and the one condition may be altered to ~he other by the action of one or more pairs of modifying factors. A point worth noting in this connection is that although ~his cross reveals a sex difference in )espec~ of horns, castration of the males does not appear to modify the horns to any great extent, as is the case in other animals whose horns are modified or conditioned by sex, e. 9. (Narshall and Hammond, 1914), though Arkell (1912) shows this to be not so marked in the case of Merinos.

B. PAP~I(CATTLE. The report concerning the other exception to the rnle of simple dominance of the polled condition in cattle, came from the Wild White A. D. BUOt{ANAN Sm'rn 367 Park Ca~le breeders in ~heir opera,ions towards bee formation of ~he domesCicaied breed of Park @stale. Since 1923, the writer has been in Couch wigh several breeders in an effort go obtain daga regarding ghe inheritance of coat colour in crosses of Park Cattle and[ other breeds. AC first, no record was taken regarding the inheritance of horns, as the figures appeared merely go be abnormal ratios. But, since practically nil the breeders with whom contact was made had obtMned qniCe nnexpec~ed ratios of horned and[ polled calves, resulting from certain crosses, iC was considered a,dvisable Co obtain some fro:Chefreliable data. The dabs which follow were obtained from Sir Claud Alexander, Ba,rt., of FaygaCe, Sussex, and Nit A. H. L. Bohrmann, F.Z.S., of Ide Hill, Keng, both of whom are leading breeders of Park @attic, and reeognised authorities on the breed. Before tabulating their experiences, ig musg be staged Chat there are, at the very leas$, ewe disCincg types of Wild White Park Ca~Cle which form ~he foundation stock for these experiments. These are called the horned and the polled. The horned were formerly ~he more common in ~he notch and wesg of , and are now best represented by ghe Chiilingham Herd in Northumberland belonging to ~he Earl of Tankerville, and ~he @adzow1 herd in , the property of the Duke of I-Iamilton. The former herd has had very ligCle, if any, addition of foreign blood within ~he pas~ ewe huncked years. The latter is no5 qnige so pure, yet has a definite type, though some West I-Iighland blood has been in~rodueed in~,o ig as well as some Chillingham. The 1 Of ghis herd, t~{r Brown, Ohamberlain go ghe Duke of Hamilton, reported about ~he year 1886, f~haL "The cows seldom have horns" (8~orer, 1879, p. 339). This was corroborated by Sir John Orde of Iiihnory, u ~fgegilllvray (in his prize essay), and R[r Ohandos- Pole-@ell. The latter in Oegober 1874 fmmd ghe herd engirely horned. Sir John Orde sugges~ sghag bhe herd acquired horns after ghe ingrodnetion of a Highland bali (SCorer, pp. 341-845}. (See also Auld, 1888, pp. 507-508.) The earliest Mmwn reference to ghis herd being potled is in 1809 when a local poeg, l~oberg Burns of tic,mitten, in the note go his poem, "Cadzow Casgle," dedieaf~ed go Sir Wal~er Seo~, wrote ~llag'~4e bulls ~hen in use were mostly "humble" (i.e. polled) and ~ha~, some of ~he cows had "/~ne moony horns." Sir Wglger Beobg himself in his early poem, "Cadzow C~slAe," wriggen a year or ~wo previously, bug dealing wi~h evengs of ghe year 1569, describes the death of a wild bull "wigh black hoof and horn," despi~e f.he fac~ ~ha~ he was well acqnMnged wigh polled eagtle, and a friend of ~[r Hugh Wagson, one of ghe founders of Aberdeen-Angus eaggle. As the original wild Caledonian e~ttle were undoubtedly horned, ig would appear as ghongh ghe Cadzow herd became polled eigher by a mutagion or ghe infusion of polled blood, imssibly after ~he Oromwellian period, when, a~ ~r Brown says, ig was "nearly extirpgged." Ig is suggesgive tha~, according ~o Burns, in 1809 ghe bulls were polled and cows horned, where~s on ghe evidence of Iager aughorigies, e.g, h{r Brown, ~he females were more fre- quengly polled. TMs mighg indicate ghat polled blood was ingroduced through ghe bulls ~bout bhe beginning of bhe sengury, jusg as bhe honied blood was several decades Inter. 368 Horns in Uatt[e

:Fig. I. :Horned :Park bull from the Char~ley I-Ierd, "Fayg~te :Brace." Though ma~ed to many polled cows, this bull left only one polled calf, and ~h~ a twin to a horned cMf. (:By courtesy of Sir Claud Alexander, Bart.) (Figs. 1-4 by courtesy of her ~. A. S. ~Iaedonald, ~f.t~.O.V.S.)

Fig. 2. "FaygM;s C4ar~er," a ~ypieM horned P~rk cow by a horned Park bull, and out of a polled cow from ~ pure polled herd. (Photograph by the owner, Mr A. H. L. ]3oln,m~nn,) A. D. BlJm-iAm~.~ S~H,r~t o69r b

Fig. 3. Horned heifer, out, of a horned Shorthorn-Ayrshire eow by ~ polled Park bull. This bull, when mated to a pedigree Shorthorn cow, produced a polled calf. (Photo- graph by Mr A. }:L L. Bohrmanm)

iFig. 4. I{orned blIll (14~ monghs old) by a horned Park bull (whi~o wigh black points) and out of an Aberdeen-Angus cow (polled, bh~ek). (Photograph by Mr J. Cob.) 370 fIo,r~s i'~ 6~c~ttle Ghartley herd, now belonging to the Duke of Bedford, is also horned. It contains some Longhorn bl.ood (introduced after removal from Chartley go Wobm'n). The polled Wild Park @attle were ioeat~ed chiefly in the middle and eastern counties of England. The Somerford herd, one of the oldest, belonging to Sir Walter Shakerley, Bart., dispersed in 1925, probably occasionally tllrew horned calves. The Woodbastwick herd, belonging to John Cater, Esq., has some Stl.orthorn blood in it. I~or many years the latter herd provided the former wittl bulls. There are now seven horned and ten polled herds, and two con- taining both kinds. The two sections of the herd book are kept qtfite distinct, and breeders who keep both horned and polled Park @attfle, have to register ahem as two separate herds. Entry is, however, allowed from inter-herd breeding, as also is admixture of blood from other breeds, provided the progeny are of the required type. It must therefore be remembered that many of the polled ammals are heterozygous for that condition. Sir Glaud Alexander states that horned individuals crop up occasion- ally in polled herds. Ke fm'ther states that his herd was founded (1908) on cows culIed from the original polled herds, mated to pare Cadzow and Ghartley bulls (horned). The herd was.occasionally replenished with cows from the polled herds, but of the calves produced by this cross over a considerable period of years, only one was polled, and that a twin, the sire being the horned pttre old Chartley bull, "traygate ]Brace." Nr Dawkins at Wileote bred Park Cattle for many years, mostly founded on white Pembrokes with crosses of other horned and polled Park Cattle. I-Ie crossed horned Park Cattle with polled Park Cattle, and also, experimentally, horned Park bulls with polled cows of various other breeding (ineNding Aberdeen-Angus). ]from such horned-polled crosses he obtained 21 calves 1 of which 19 were horned and two polled (one bulI calf and one heifer calf). In the Lyme herd (horned) which belonged to Sir Piers Leigh, when a polled @isbttrne bull f~om the Gisb~trne herd (now extinct) was mated to a horned cow, the progeny was fully horned, bug the horns grew downwards instead of outwards. Whether there were any polled calves from this cross does not appear to be known. Of precise rantings it is difficult to obtain, data, as reliable figttres as regards sex, etc., do not appear to have been kept. 1 The experimeng~lerossbreds were a.I1, later, eulled from the herd. A. D. BO0~{ANaN SMI'l't:~ 371 Mr A. H. L. Bohrmann 1 has supplied the following: An Irish black ~oolie cow (polled) cMved to a Friesian-Park • Shorthorn bull (horned) and produced a, horned heifer calf. A horned bull of various of the best Park strains and with one British Friesian grandparent, mated to an Aberdeen-Angus (polled) cow, has produced a horned bull oaK. These together with other results are given in Table I.

TABLE I. Su'mm, a,,ry of O'rosses of Pa,r/(: Cattle, etc. }[erd Sire and D~m Offspring ]~enlarlr ii'aygaee Park H* (Cadzow) x Park P-l" All t-I Frmn I908 to about t918, 30 calves Park l:t (Chartley) x :Park P All H s~ve t (a twin) Wileote Park H • P~rkP } 19H(8d, 99) Park H .'< various P including Aberdeen-Angus ~ 2 P (1 d, 1-9) Lyme Park P (Oisburne) x Park H AllH Perhaps one or two P Ide Friesiau/Park-ShorHmrn H x Irish Moolie P H 9 Park (some Friesian) H • Aberdeen-Angus P I-I Park H (Ohartley) x Park P I-1-9 (a) P~ed Poll-Park P x (a) K P 9 Park P • (o.) K P c~ With sours ? g x(a) H ~c~ Park P x Shorthorn-Ayrshire H I-1.9 Same bull mated to produced only polled * H=Horned. t P=Polled. The proper interpretation of these cases really rests on the fact that an abnormally small number of polled calves have been produced by a mating of horned • polled, even when the polled ~oarent was registered in s polled herd, or of s pure polled breed. Nevertheless, it must be remembered that normally these uneven ratios do not ocettr often, even in s large popular, ion, and that therefore, since the population from which they are culled is extremdy small, their significance is the greater. No reliable dates have been obtained of any cases where horned x horned have given polled offspring. Only the very general conclusion can be dra~vn that the Wild appear to carry certain factors which modify the normal mode of inheritance of horns.

DISCUSSION. An experience similar to these was recorded by Wentworth (1911) in the case of the famous White Shorthorn x breeding 1 ~Ir Bohrmann has ])red a considerable number of crossbred cattle for experimental purposes only (not to augment his herd). gourn, of Gen. xwr* 25 972 fflorgz, in Ua, tt[e experiment a~ Iowa State College. WRh two exceptions, the 70 F l animals were all blue grey and polled. A polled 1#1 bu.ll was mated back to three galloway cows (polled). By one, he sired only polled offspring, and all the calves this cow produced by horned bulls were also polled. By another, he produced a horned bull calf, though this cow had six poIIed offspring afterwards. By the third, he produced a horned bull and a horned heifer calf at diifferen~ births, as well as four polled calves. It is probable that these last two cows were heterozygous for the polled condition, and that it was a oh.shoe that two such cows should ]].ave been used. But the fact that the horns of one of these calves resembled not a bit the horns of any ptu'e breed of oattie, may lend some doubt to this statement. ~ossam Boyd (1906) mated a potled ~nz~tcsna Herefm'd bull to horned Hereford cows, and obtained the curious proportion of 22 polled to 6 horned ealves. He also records (1908) data drawn from crosses of Bison bulls on 14 polled cows which seem to show that in this cross ~he inheritance of horns is the same as in domestic breeds of cattle, with the polled condition a simple dominant. Oowen (1918, 1927) states that when the factors for the polled con- dition are in the simplex sta~e, "they tend to produce the horned con- difiion more frequently in the males than they do in ~he females." He gives an example of a black bull with shor~ horns produced by a mating of an Aberdeen-Angus bull (potIed) on an Ayrshire (horned) cow. Spillman (1905) had already suggested this but without adducing any evidence. Watson (1921)also found the polled condition to be occasionally in- completely dominant in the male. There is a further point to which reference may be made, namely, the condition as regards horns of early forms of cattle. The majority of authorRies recognise tha~, it was a polled animal that was the fore- runner of the presen~-day cattle. It has, however, been pointed ou~ by many writers that these polled types were succeeded by species in which. the names were horned and the females polled. Prof. J. Cossar Ewart (1909) states: "Several species of cattle with horned males, but hornless females, occur in the lower Pliocene deposits. In Leptobos Fa~cone~'i of India, L. Rtrz~sczts and L. Lat~ts of Italy, France and Spain, we have early Pliocene species in which horns were probably always absent in the females. In the deer group, the females as a rule have continued hornless, but dt~ring the later Pliocene period, female as well as male oxen acquired horns, and the horns seem to have been retained by both the males and the females tmtil cattle were domesticated during the A. D. BU0~:IAN~N SMIT~I 373 neolithic age." This view is substantiated by Boyd, Dawkins, Auld, Lydekker, l%utimeyer, Arenander and others. This therefore inclines one to the conclusion that the simple domin- ance of the polled condition is derived from an earlier sex-limited con- dition which is still to be found in certain types of cattle, especially those in which there has been little or no admixtm'e of the recognised modem domesticated breeds. In the genetic a,cquisition of horns, cattle seem to illustrate a generM evolutionary principle. The mutation ~rom the polled to the horned condition was first exhibited by the male, and only much later in ~he history of the species by the female. As no accurate data were obtained concerning scars or slugs, no study of these has been made, though the results of Watson (1921), Cole (192~) and Landauer (1925) would seem to show that their ex- pression is largely sex-controlled.

OONCLUSION. There are n~maerous records which confirm as a general rule the simple dominance of the polled condition in cattle, and it is unnecessary to quote them in detM1 as the fact has been well established by the common practice of cattle breeders. The purpose of this paper is to show that there appear to be factors which modify the normal mode of inheritance. The writer in no way claims that the data presented in this paper deny the established rule, but rather suggests that in certain crosses there are additional complications, and that these complications appear to arise when blood strains other than those of the commoner domesticated breeds are present. This appears to be a perfectly logical evolution of the study of the inheritance of horns, and may be compared to the progress of the studies of many other heritable characters in both plants and animals. ACK~COWLEDe~IENT. The writer wishes to acknowledge the assistance he has obtMned in the compilation of this paper from Nr ~. A. S. NacDonald, M.I~.C.V.S., Sir Claud Alexander, Bart., Mr A. H. L. Bohrmann, F.Z.S., :Prof. J. Cossar Ewart, F.P~.S. and Dr F. A. E. Crew.

25-2 37-i Items '[~ Cattle

LITEI~ATUI~E CITED.

A aK:~LL, T. I{. (1912). "Some Da~a on the Inheritance of ]~Iorns in Sheep." New Hampshire Agri~. Ex23. ~ta. B'ull. 160. AOLD, I~. C. (1887). "Hornless ~uminanl~s." Amer. Nat. xxr. 730-748, 885-902, 1076-1098.

-- (1888). "Wild Cattle of C4rea.t Britain." Ibid. xx~. 498-509, (1888). "Derivation of the Domestic Polled Breeds." Ibid. XX~. 784-802. BOYD, Moss~ (1906). A'merican Breeders' Association, Annual 2~e2ort, IT.. 198-201.

-- (1908). "A Shor~ Aecmm~ of ~m Experiment in 6~ossh~g the American Bison with Domestic Cattle." Ibid. zv. 324-331, COLE, L. J. (1924). "The Wisconsin Experiment in Crossbreeding Cattle." Paper from the D~paTtment of Crenetlcs, Wiseon,~in Agricultural Ex2eriment Station, No. 40. Ca~w, ~. A. E. (1924). Animal Genetics. Edinburgh, O~ver and Boyd. EwA~% J. Coss~ (1909). Hornless Cattle. Edfl~burgh. Quo~ed inHistory of Aberdeen- . By ~Iaedon~ld and Sinclair, London, 1910, 9. 14. See also Cattle .Breeding, :Edinbtn'gh~ 1925, pp. 2-16. Gown~, J. W. (1918). "Inheritance Studies of Colour and I-Iorn CharacterisMes." Maine Agrie. Ex2. Sta. Bull. 272. "

-- (1927). "A B,6sum6 of Cattle Inheritance." Bibliog. Genet. ~. 87-138. :L~nAUEl~, W. (].925). "Ergebnisse in der E rbanalyso der Behornung yon l:Lind, Seha~ and Ziege." Zeit. indulct. Abst. xxxlx. 294-322. i~I.~I~SKAL~, F. H. A. and I-Lu~mlqn, J. (1914). "On ~he Effects of Complete and ]hcomplete Castration upon Horn Growth in ~erdwick Sheep2' Journ. of Physiol. x~wz. I7I-I76. S~m~, W. J. (1905). "Mendel's Law in relation to Animal Breeding," American .Breede~'g Assoc. I. 171-177. S~o~, The ~ev. Jom~ (1879). The Wild White Cattle of C~'eat J~ritai~. London, Cassell, Petter and Galpin. }V~soN, S. A. S. (1921). "A Mendelian Experiment with Aberdeen-A_agus and West Highland Ca~tle." Joum. of GeneEcs, :~. 59-67. W~TWO~:~, E. N. (1911). "Se~ega~ion in Cattle." American .Breeders' Assoc., Annual Re2ort, viii. 572-580. See also LLOYD-JoI~ES, OI%I~E~ and EVVAI~D, J. l~i. (1916). "Inheritance of Colour and I~orns iu ]3lue-g~rey Cat~le." A~Tricultural Nx2~eriment ~tation, Iowa 5'tare 6'ol~ege, Rese~'eh ~ulIetin, No. 30.

EXPLANATION OF PLATE XXII.

:Fig. 1. "~hodesdale Magistrate," s pure bred Aberdeen-Angus bull used in case l. :Fig. 2. A ~ypieal Mashukalumbwe cow showing ~he horns, wi~h a 6-months old grade calf at foot, Fig. 3. The male progeny of a cross of such a bull as shown in Fig. 1, and bhe cow as shown in. Fig. 2, and others like her. The bullocks are ~wo years old. Note ~he various ~ypes of hor~ls. Fig. 4. The female progeny of the same cross, all completely polled. They are ~wo years old. ,JOURNAL OF GENETI08, VOL. XVlll. NO. 3.

Fig t

Fig. 2. PLATE XXII

Fig, 3.

Fig. 4,