Studies of the coat of the Romney lamb : practical and theoretical aspects of hair morphology, with special reference to the evolution of the fleece : thesis submitted by "Bourn" [for the degree of Master of Agricultural Science]
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(2) The is. ~ubmitted. by. conce rning ~ubsecti oo( ~) 608, of The Animal quEbBndry recti~n of the Uas ter of k 5 ricQltural Lcience Degree 1'340.. practical and theoret ic al aLpects of hair kith E~ecial refer e nce to the evoluti on of ~he tle ce.. mor p holo~y ,. ---000---. I. 1fifTfff111.
(3) CON1'ENI'S.. psge.. ... . .. .... INTRO DUCT I OlT. 1. SECTION I. .... I. MATERIAL ANt fSTHOD 1. DrytE Experim~ntal Romney Sheep 2. N-Ty pe 3. Non-N-type 4. Covering. ... .... .... 13 14. 5. Semoling Position G. 8amp li 1g ti ·r.e 7. Laboratory Equipment •• 8. .ruee.~ uro men t. 1. 9. Dyeing 10. Dyeing the ~heath of 11. jjenL.ol Te~, t. 12. Tatooing II.FI 1R~ ~YPE~. AND. FI~RE. 15 f i bres. o:T. . ... rIF"'E RE~TT. 16 16. 16 17. type arrays. III. FIBRE Tn>r::S ..t>•. ~hed. 14. 'f YPE AFRAY2. 1. IntroC.ucti .)U 2. Fibre ty es. 3. Fibr. ... . ... 9 9 11 12 13. 18 18 19 21. BOJJY REGI- 'S ;::;5. .Zre-Cll.rl.,y -Tip Grollp. .... 1. Su.b-?"3 lo-hair ~. ~al0 -h air Fhape (ai Falo-hair ~h&pe (b) 3r:~ - he1r lpng.h ( c) R& lo-! air Coaz· ~ene u._ •. 25 25 26 2b. 27 28. (d) HalJ -haj r abil.tldance on. the ~t. E~·. me boc..y regl ull. different lambs. {e) p·ab-hAir 3. r=utJ.<::r. nb·indan c~. 29 on. different bctu regions ~f the same lamb. 30. :'ic~~le. 34. A. (aJ Tip sha pe {'b) Super sic ~cle ( c ) S u pe r E i c k 1 e. ••. •••. shape .r. a u u.11 1ancP ?n the same body J::i. 35. 37. re gio n of d if ferea& l9mbe 37. >.
(4) -i i -. .... .... 4 . 2uper ickle A' (a) ibper s i ckle n' abundanc e • • 5 . Super Sickle B. . ... (aJ Super sickle B fibres chalky t hro ugho u. t \b) Super sickle B fibr e s fine thro.igho.it • • • {c } Super ~ickle B fibres chalky i a Lhe di etal portion and fine in the proximal porti.:>!1 ldJ Super Sickle B f i ore~ c halky in the proximal as uell as distal port i on , but fine in the midnle part >I the fibre ( e) Tip shape of s u.pe r .sic le 1 fibres •. • (f) super sickle ~ abundance 6. Sickle Fibre ••• ~a) Definitio'1 { b ) T i p ~ha pe. •••. • ••. (c} Length and Hairiness o Fie le ends .•• (d) Tuft snape influencing :::ha.oe of .Pre-nat a l part 0f :: iclde fibrec ••• {e) Port - ~a tal ~ o rtion of sickle 'ibre.... • ••. 40. 41 44 46 49 49. 53 54 55 62 63 63. 67 71 7'3. (i) Chalky throughout ( i 1 ) J' L , e th r o u.t.. ho u t (f)"To~ghnese " of havine array~. 75. (h) Remaining lO an~ c) cl ss~es of cickle fibres • • (j} Abundence of chalk;,- ~ickle. f:3. 74. 76 (g) Abundance af .11'ine sickle fibres 79 in valley arrayE. fibrec:. (· ). ~le. •••. tenu a rra f • (ii} SaddJ_e arravE •• (k ) Tnfluence of Body Re~ioc upon sic_le f ibre 8bQndaLce •.. f5. P5 86.
(5) ... ... -iii-. .... B. Curly-Tip Group ••• • •• 1. hairy-tip-curly-tip gibre ••• Tip shaY'e •• \ b ) I1e ng t h •••. •••. \S.I. • •• •••. 91 91 92 93. (c) Structure of the Eost - atal portion . • . •• \d} Ha iry tip c~rly tip fibre daace •••. • ••. 2. Curly-tip Fibre'"' ••••• (a) Tip form ••. {b). ~ost. atal. 95 a~uc-. 101. . ... Struct~re. l Oi. lu7. •••. 108. C. .P0s t-C.ur ly -'l' ip Group {Hi ste r.) tr i c hs) ••. 117 118 119. 1. ~ t ruct~re of flisterotric h fibres .• 2. Time of Rieterotrich Growth • 3. Uro.~th vf lliste .r:-o t.richs and the ven~ity of the fleece ••• 4 . ... ~d.e of lii~terotrich t.irowth. • •• 5. Ab~ndance of Eistera~rich f i br er ••. 122 12.3 H:5. fEC?Io:r I I. : :rrrR I. SHVDJHNG ·J~I BI RrfTC().AT FIBRES l\ 1,... 3 .... UCCES~ JRf, 1. &b e dding in the ?re -Curly-tip 0 l'Ollp; it~ var ia t io nB i l1 different Bo, ,y RegionE 134 o.nd arrays 146 2. ~hedding in Q11rly-Ti r - Grou.p 147 3. Shedding 8f Ei ::-tero trichs 14? lI. MODE OF S!~EDL'I::' G. . .. .... III. G2. .. ...... ... .... ... rray.... .... anc:. 1. .h'.ela t ion b;:: t·.veen G..,,. ..&.. f:. 2. Kels. t ion bet ''3en G~ an( Boey Re gi or:. J. He la ti on Bet Nee Kemp-bai ri r.ess (G2) anC. non-1{6 rn... ha irinet 1;:. \ pe1· .... istent cha 'c;1 ii ore:=-. 151. .. .... 151 151 ...,e:, P·". s::.:c11ION III. FIBRE TYPE ARRAY GR.b £ I ~NT l. Distribution of the Arrays over the l amb 'e Body. I.BRITCH- ¥ITHER~. 159 162.
(6) -iv-. II.. .... D I STRIBUTIO~ 1. tla ir ines~. JF ~A I RINESS in the same Arrays o n " iffe rent tsoC.y regL.> ' ~ • • ( a ) Plateau a r r eyr: (b) Sa~d le arrays •••. (c} Rav ine arraJs ••• \ Q} Valley ar r ays 2. Hai r i ne sE ~n Diff e re n t. 168 168 169 169 170. on. ~ r rays. 172. d ifferent Body r eg i )nE III. FORCE RE~Pon.::i r BLE FOR THE OF FAIR!NF~.._,S 1. The P reci pi ce • 2. .. Cr i s is th i nning " •••. 162. LITNUT ICJ N. .... 176 177 183. .... SECTic:: IV. I. N- TYl'E - FRO ·'1 THE f'TAN!HOINT OF 1. In tr oduction • ••. 2 • .1v -ty ;_le - ar. ~<.:xRmpl es. -i:'. T()J,U'llJ('-,;. of ..; tevu,·m. 191 191. 193. sr;CTIO'l'T v. 1ntrouQction. 1.. ••.. Ev o lution in the l:'r e 1..1urly f i ;J Group • -i:-v Jib.uti.J rl in ti1e vurly- t ip liro.i~' •.• ~ v olution in tl i terotrichs Helati1nehip betwee n Fibre Typ e Gr oups Evolutionary Tendency rnd the 1~neity. v.. 4.. 5. 6.. of the fleece •• •. L 2. ..,'~. .. "'.')R. FF. f'l'.'T filC .AL. 227. uo· ;:,l:En.ti rIO~;. u.:ic· Tl:!": 3RI'CrIIiG BJUT 1'H- ":VO il'F~ <'L-;J1C,.., De.uses of Bvolu tb· r. i nd cf 1 ~uta ti on tco1e of Mu tati on and. SelectL.rn in evolution (.Ti:·. WQRCt"!: .Al1 UJ::' 11111 OF. 219. 22!. II. FOT\CES A'I! 1-0RZ: r,?·Sf·JI~I2.I,f -C:VOLTJTJ-:;r O.tr THT: i: C!:.!NEY. III. ._QfF. 203 203 207 211 216. 1. 0~K. 231 231 233 241.
(7) -v-. 4. Selection an d its ~art in Evolution •.• 5. Role of "Small" ,\ltutations and Se l ection in the Evolution of the Fleece.. ~ECTI ON. 251. VI. I. THE CJ.:IliRACT ES! OF LA B'S r1 00L. 1. 2.. 249. ... .... lntroducti0n ••· Fineness ••. • •. (a i Rcl ~ tionship bet~een finenes8 of ool an d bo dy re3i~n ••. 254 254. 258 258. (b) Relation Fhi. p bet Ae en .il.. neness. of tJ.. 4.. Summary. ool an c Fibre Tyr e Array. .ia ve vrimp. .... . ... .... Biblvogre. phy .Appendix. 260 262 2i:.; 4. 272. 285. .... .... --- oJ o- --. 289.
(8) INTROD UCTION.. The original p ur pose of the precent thesis was to an s Ner t h is. "To wha t exte nt. q~e F ti o n:. i~. a. britch o f hi5h halo-hair abundance a n indicator of the hon-kem p hairiness of t. he fleec e ? " by ur. 1ry in the ligh t. of earlier on a britch. ~i th. very many. Ti p fibre s would be be. E O.. ve r ~. It wa'"' assumed observation ~ ,. halo-h~r E ,. that,. th e l arge Curly. hairy , a nd th i P ha e p roved to. Especi a lly d i e we want to lea rn ab out the. degree of hairiness on th e part of t he fleece nea r to Lhe britch.. The g ra die n t o ve r t he bo dy from the. britch ·as also much in mind , a l l the more because t he bounda ry bet we en the a r e a on th e britch with very. ma-i y. hal o ha ir e: nn<l. t he nei g hbour int;S region with f a r fe _, e r ~ttl.o hairf i f often abrup t. The a im o f the \ ork wa"' tha t ,i us t defi n ed , but t he. ana ly~i~. of t he samp l ec. for mat ion on a numbe r of other. e x s~ ine ~ mat ter ~ ,. provi ded insevera l of. which ma y be tho ugh t more in te re c ti ng th an the pro ble m propo s e d a t the o utset.. &re r e porte d in this thesis.. Thes e va r i ou E re f ultE.
(9) -2-. In carrying out this work it wae inevita b le that I e houl d r e vie w th e fac t s a nd i dea s co Gtributed by. earlier worke rs a t the J.rars ey Ag ricultural College.. Their. ~ ork. ha s all pa e sed throug h my mind .. little ban d of .. orke r~,. it. ha~. This. bee n po in ted ou t t o me,. ha s never oeen large ana i t ha e alway c been isola ted. I have e n tered t his school of thought fro m P rofe ~ sor Pra woche ncki' s De part me nt of the. U niver ~ ity. of. ~raco N. (Cra co w, Polan d) and ha ve s o,ne t i mes f ound my self offe ri ng i n ter pret at ion s "' lterre tive t o t hose put for ward 'l'hese i dea s ha ve. at t he Massey Ag ri c ultural Gollege.. been welc o e a , a lt holl6h t hoce h ol d i ng t he ol der vie ,s te l l me tha t they do no t expec L to prove .• ro ng every ti me . Thi s will. ex p la in. ~. hy I have de a lt 'Ni t h my. [ Ub j ect br oadly a nd at come lengt h.. Not only has t '-:e. method of presentati on helpe d m, work ing ac u . intance with t he cla cs of f a c t a nd mode of t houg ht of my ne w as s oci Ate s , bu t at the same time I have. ~ een. encoura ged. dlo revie \ critical ly the conce p ti on '"' a bJ llt the fleece hich I have encountered in Ne w. ~eal and ..
(10) -3,,.. );- ·". Guided thoughtfu.lly by Dr. F .• ~ •• Dry, and his closest a ssocia te, Mr .J.A . Sutherland , who. ~v ere. enoug h to teac h me th e principles of. Type Arrays ,. ~ ibre. kind. I was surp ri se d to fi nd the sheep's birth coat suc h a. wondefful obj e ct f or a fun dame n t s l s nudy of manifold one may think t hat the b irthcoat. aspects of wool.. --. is a "m i c roeosmos" of it"' own , and, i f Jne '1 as a Euit able method of app ro a ch the birthc oa t - I am sure Nill turn out to be an ind is pens able. ob ~e ct. l og ic c:.l studies.. fo r bio-. •. Dr . J.Jry 's p i:i neer i ng <r, ork has. lrea dy furnished. not only the co nce pt of a pproach, t hiE being the. ~ ibre. Type a r r ays , bu. t alsJ the m""terie l f or comp' ra t iv e study. This work and line of th ought ha ve op e ne d up quite a n e w br an c h of bio logic· l s cie nce, pa ra ti ve anatomy of co Lvi n ce d -. ~ ill. ~ ool " .. hich c £. OP. cr>l led ''Com-. Th i e new dic ci pli ne - I am. open a l arge f i el d. or i nves Ligati ons,. wh ich will he l p to get f ar dee per unce rr:tanding J f from the purely practical po int of. ./ .,,,-I. /. > , (. l> lfo, it dll,. vie ~ .. to my mi nd , throw light upon s uch vivi d. p ro ble m~. of. a~. v olds c h m i d ' ~. p~ysiolo g ical. que s ti on of evol u ti on . one exa mp le.. genetic s ,. rool,. ~e l l. as those as. the. Le t me pe permitted to g ive but. The pre-nat al -c heck, as . e understand i t. nJ w, i s a force, wh ich ma inly. c ause~. fib res at the head.
(11) -4 -. e c a n distinguish at least. of an arra y to be fi ne;. three importan t propertie s offohe check, namely : (i) the outset,. that i'"' the time when the pre-nata l check st rt s t') • ork;. (ii) The inte n sity, th u.t i f. th e whe t he r i t i. tre ngth of >- he c heck, c:. po e r f u l en? :ig h t o. cause tm. fibre to be Line t hroughout. or in the. n ee ~. regi on only,. an d finally, (iii) The exte ns ion of t he c heck, t ha t is, ho w f a r it exte nds in the Curly-Tip Group. The pre- nat a l c heck 1' h i. ~. i~. the r eru lt o f t he. ork we c an 11eaL1re ve r y a cc u. ra te l y .. ·~ ork. of g e ne s .. ..tror ins t a nee,. {. In Valle y arra.., s the out s et can be me as .ire d by the numb e r of halo-hairs a nd S llper s ickle number of fine. A. f i bres, the in ten s ity by. ickle fibr es end t he ex t ension by. .e. of c hecked curly-t ip fi b re s .. n~m b er. c Pn , fu r t her mo re, ex-. pre s s the mea sureme nt a c: a pe ccent age of cer tn i n fi b re. typer, t h i E' be i ng deci dedly more a vantag e ou i: a nd pro ba b l y G-. more accurate , t han ar b itrary. nv e stigi a l" c a"' e i n. ~r ') so p hi la .. t er m~. use d i o Goldschmidt' s. lit see ms to me t hat t he. work of ge n" s c ausing "ve s t i g i G. 1 11 i n L r :le o phi la, is very <:;. i mi l a r to the wo rk of gene s. in the oir th coa ti.. c au.~i. ng t he p re -na t a 1 c heck. 7 j.
(12) -5-. I have dealt t o some extent with the quest ion of the ev olllt ion of Lhe flee ce in the bre e d .. 'l'he. m~. t eri al for. Tew Zealand. Romney. pc. rt of my thesi i: : has. thi~. been gro - ing Qurin5 the c o ur Ee of my. inve~tigation,. 1hk h. i nc. ee d had n:)Lhing to do \ith the pr oble m of th e evoluti :m of the fleece and therefore I could hardly 1e a c cus e d of "having an ey e seeing . hat the eye it. wa~. 100 1.cs. for".. In deed,. too great a te mptation no t to use data o btained . re~en. in the course of the to outline the. ay. l. inve""tigatiJn in an. atte~pt. hich the evolution of the flee ce. .. has probably t aken pla ce . The. ~tudy. evidence from three 105ic al , and. my. the~is,. 6. of evolutionary ~oir ces:. eneti c al.. using. perh ap~. pro ces ~ e s. Pa le ontolJgi c aJ , embryo -. 'l'hese S..>.ir c es are c.is c rnsed in different word ing.. we may add, that by embryology of ool, L developmental changes a staple .. aE. In. p8~si~g ,. mea n t the. noted in the prenatal par t of. It i s fe lt , ho ever,. be g iven aE t o the firi:::. requires. source.. ha t c:.n exQlanati on muc-t Dr~'s. Exper i mental. sheep co uld hardly c laim to be paleontological examples. Nevertheless, I feel at ease t) s ay that tology serveE ch anges ,. ~ry's. ~c 1. a~. fc:::r as paleo.a-. a F0ur ce of evi dence of evolutionary xper imental flock may be regard e d as a. 2ource of phylogeneti c al eviden c e of. th~. ev)lution of. he. fl e ece, pr ovi ded, however, that bQ one . ill ag ree tJ regard.
(13) -6the Merino wool (being generally of most evolved. ~ 001. ~ild. c oat, ana on the other hand, the mixed. ~he a p 's. th&t i. lain array) as tt-e. -. , the most removed from the. J-ty pe - being. woolled !=:heep lBlackfa ce, ..t'olish Cakiel,. probably at least of Plateau array} ae the la t evolved wool, that is, not too far removed from the wild coat. This Jr ing so,. ~e. c an claim to be in a very. position to have the. 1. fo~t~na~e. pa leontologi c al" evi dence not in. the form of f.)"E ils, bllt in the form of liv e b~. ~h eep.. to me, I ha ve had the opportunity to 5e' acquainted. with. ~oli sh. mixed woolle d. ~ heep. ac well a!': Mer inos.. g a p between them was too broa d to bridge. ho t~ ever,. i '". The. The briage,. to be fol10d in Dry'::: Experirnent::.. l flock.. Inde e d , it 1iar worthwhile to sail save.Bl. at lea i? t,. ~eas,. to find thf> '' lost IT links in the c ha in ·of the fle e ce's evolut ion .. Ye t , although my ne ··. home is remote from. Central .i!:urope, i L is intere sting tD ha ve it pointe d ::>Ut to me that the work in Which l have drnme to take pa rt. has roots i n Vienna, in Told 1 s f ibre Lypes, and in sterdam,. in de. and developme n t.. ~eijere'. fund an-: ntal~. of ha ir ar r angement. I am told, besides, that. he voyage. from "'!i'urope should have been mad e by the 9ape, Dllerde n '~. check.. "b ir t h thinning 11. m-. ~oresha dowed. ·here. the pre-natal.
(14) -7-. My. philoEophy, it i s hinted t o me, occa si onally. showE signs of running ahead. of my factr. be true, if deeme d a. mer ~ t. 'i'hi.... ,. in the department where. if it l. work,. provi de d one oe a"ar e of the free rein g iv en to i ma g inati )Il. I t is a source of pri d. e rat her t han s hame in t hi s de pa rtme nt tha t a paper rn ce submitte a to ProfesC>or '.IT. 3 •• Ha ldane in his capacity of Ed itor of "The Journal. of Uenetics" was refuse d on the s core of be ing t oo sp ecul ati ve. ~p eculation. ferent. rea~on~.. i s value d he re for t wo. In the f ir et pla ce. quite dif-. pre s e nt knowle dge. ) fl. the surest scoring sho ts in lives to ck bree d i ng - one natura lly thinks of proge ny te sting - a re apt t o be clum y. 0. Deeper unlereta ndi.ng may make defter str oke s possllble, a nd this under stan d ing will be gained on ly. thro~gh. lively. i mag ina t i on. Speculation i s a l so welcomed because adventurous thinking i s e s senti a l if our ma te r i 2 l i s to yield some con tribution to fundament n l ge rn tic E.. 'hen i ndee d one reflec t s. how villing to speculate ab ut evolution comparative morphologiEts have been on provi des our j ustifi cation.. les~. '1' o. e xp lored be. ~. mat er i~ l,. thie. ur e , t he va 1 ue of.
(15) -8-. such phylogenetic '"'p eculation has ce e n called into aue f:"' tion Yet. by Bateson hi mfelf.. ~ha tever. a pa leontologis t c an. sho w c ause to believe ha s taken pla ce, it ir the businer s of ' he geneticist to d iscover the ge netic r ri ncipleQ therein in volve d , and it attem pt e d. i~. a very s imilar s ort of thing tha t is be i ng. i th the ma nif o ld va ri a tion c revealed in t he coat. of the "'hee p . The importanc e of t he pas toral At te nt i on to. ool. in a u ~ try. at t r a c t ed. Hence the i nt ensive st udy of. rese ~ rch.. the fibre typeQ of the Romney coa t and their developmen t . On ce th i s i n ve st i gat io n wa'"' well lau nche d , the conviction /. c a me to prevail Lhat th is cl ass of f a cts, ba. i s for t he. pl an nin~. ~erving. a. a. a nd in t er pt e tation of expe ri men t a l. bree d L ig , WJ uld oe of val ue to pure g e net ics.. That th is. is not a n unrea Qonable pr obab i l ity ,a s pointe d Ju t before, is apparent in t his the s is. At t he end of th i s i nt ro duction, I feel it is my duty t o apologise for having written this thesms in very poor Eng lish.. In s pite of having. Eng li Qh be fore be ing t a ught ou t that I I. 11. ~tarted. to learn. ha ir- Qp li tt i ngn i t haQ turned. m a f a r poorer linguist than a "hair-r p1it te r n.. hope i nd ulge nce. ~ ill. be g r ante d me all t he more , if one. is not apt to lay " •••••.•. blame on t. be pe t r-tree for not be a ri ng plums" . (Alain).. ---- oOo ----.
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