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The is ~ubmitted by
concerning ~ubsectioo(~) 608, of The Animal quEbBndry recti~n of
the Uaster of k 5 ricQltural Lcience Degree 1'340.
practical and theoretical aLpects of hair
morpholo~y , kith E~ecial reference to the evolution of ~he tle ce.
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I 1fif Tff f 111
CON1'ENI'S.
psge.
INTRO DUCT IOlT
. . . . . . ...
SECTION I
I. MATERIAL ANt fSTHOD
...
1. DrytE Experim~ntal Romney Sheep 2. N-Ty pe
3. Non-N-type 4. Covering
5. Semoling Position G. 8amp li 11g ti ·r.e
... ...
7. Laboratory Equipment ••
8. .ruee.~ uro men t.
9. Dyeing
. . .
...
10. Dyeing the ~heath of ~hed fi bres 11. jjenL.ol Te~, t
. . .
12. Tatooing
II.FI 1R~ ~YPE~ AND FI~RE 'fYPE AFRAY2
III.
1. IntroC.ucti.)U
. . .
2. Fibre ty es
3. Fibr type arrays
FIBRE Tn>r::S
o:T
rIF"'ERE~TT BOJJY REGI- 'S..t>• .Zre-Cll.rl.,y -Tip Grollp
1. Su.b-?"3 lo-hair
...
~. ~al0-hair Fhape (ai Falo-hair ~h&pe
(b) 3r:~-he1r lpng.h
( c) R& lo-! air Coaz· ~ene u._ •
(d) HalJ-haj r abil.tldance on the E~· me boc..y regl ull
~t different lambs {e) p·ab-hAir nb·indanc~ on
different bctu regions
~f the same lamb 3. r=utJ.<::r :'ic~~le A ••.
(aJ Tip shape •••
{ 'b) Super sic ~cle J::i shape ( c ) S u pe r E i c k 1 e .r. a u u.11 -
1ancP ?n the same body region of differea& l9mbe
1
9 9 11 12 13 13 14 14 15 16 16
16 17 18 18 19 21
;::;5
25 25 26 2b 27 28
29
30
34 35 37 37
>
-ii -
. . . ...
4. 2uper ickle
A'
(a) ibper sickle n' abundance •• 5. Super Sickle B
. . .
(aJ Super sickle B fibres chalky t hro ugho u. t
\b) Super sickle B fibres 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 fiore~ chalky
in the proximal as uell as distal porti 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-natal part 0f ::iclde fibrec •••
{e) Port-~atal ~ortion of sickle 'ibre... • ••
(i) Chalky throughout ( i 1 ) J' L , e th r o u.t.. ho u t
(f)"To~ghnese" of havine array~
(g) Abundance af .11'ine sickle fibres in valley arrayE
(h) Remaining lO an~ c) clss~es
of cickle fibres •• (j} Abundence of chalk;,- ~ickle
fibrec: •••
( · ) ~le tenu arra f •
(ii} SaddJ_e arravE ••
(k)Tnfluence of Body Re~ioc upon sic_le fibre 8bQndaLce •.
40 41 44 46 49
49
53 54 55 62 63 63 67
71
7'3 74
75 76 79 f:3
f 5 P5 86
. . . ...
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B. Curly-Tip Group ••• • ••
. . .
1. hairy-tip-curly-tip gibre •••
\S.I Tip shaY'e •• ••• • ••
\ b ) I1e ng t h • • • • • •
(c) Structure of the Eost- atal portion . • . ••
\d} Hairy tip c~rly tip fibre a~uc- daace ••• • ••
2. Curly-tip Fibre'"' •••••
. ..
(a) Tip form ••
{b) ~ost atal Struct~re •••
C. .P0s t-C.ur ly -'l' ip Group {Hi ste r.) tr i c hs) ••
1. ~truct~re of flisterotrich fibres .•
2. Time of Rieterotrich Growth •
3. Uro.~th vf lliste.r:-ot.richs and the
ven~ity of the fleece •••
4 . ... ~d.e of lii~terotrich t.irowth. • ••
5. Ab~ndance of Eistera~rich fi brer ••
fEC?Io:r I I
I. SHVDJHNG ·J~I BI RrfTC().AT FIBRES l\
: :rrrR
.... UCCES~ JRf,
. . .
1. &bedding in the ?re-Curly-tip 0 l'Ollp;
it~ var ia tionB i l1 different Bo, ,y RegionE
o.nd arrays
...
2. ~hedding in Q11rly-Ti r -Grou.p
.. . . .
3. Shedding 8f Ei ::-tero trichs
. ..
lI. MODE OF S!~EDL'I::' G
. . . . ..
III. G2
... . ..
1. .h'.ela t ion b;:: t·.veen G .... ..,, anc:. ..&. rray 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:=-
...
s::.:c11ION III
91 91 92 93 95 101 lOi lu7 108 117 118 119 122 12.3 H:5
1,...3
134 146 147 14?
151 151 151
P·" ...,e:,
I.BRITCH- ¥ITHER~ FIBRE
TYPE
ARRAY GR.b£I~NT 159 l. Distribution of the Arrays overthe lamb'e Body 162
II.
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DISTRIBUTIO~ JF ~AIRINESS
. . .
1. tlairines~ in the same Arrays on
"ifferent tsoC.y regL.>' ~ •• (a ) Plateau a r re yr:
(b) Sa~dle arrays •••
(c} Ravine arraJs •••
\Q} Valley arrays
2. Hair inesE ~n Different ~rrays on different Body regi )nE
III. FORCE RE~Pon.::irBLE FOR THE LITNUTICJN
OF FAIR!NF~.._,S
...
1. The Precipice •
. ..
2. .. Cr i s is thinning " •••
SECTic:: IV
162 168 168 169 169 170 172
176 177 183
I. N-TYl'E - FRO ·'1 THE f'TAN!HOINT OF -i:' T()J,U'llJ('-,; 191
1. Introduction • •• 191
2 • .1v-ty;_le - ar ~<.:xRmples of ..;tevu,·m 193
1.
v .
4. 5. 6.
sr;CTIO'l'T v
1ntrouQction ••.
Evolution in the l:'re 1..1urly f i;J Group • -i:-vJib.uti.Jrl in ti1e vurly-tip liro.i~' •.•
~v olution in tli terotrichs
Helati1nehip between Fibre Type Groups Evolutionary Tendency rnd the 1~neity
of the fleece •• •
II. FOT\CES A'I! 1-0RZ: r,?·Sf·JI~I2.I,f "'.')R FF -C:VOLTJTJ-:;r O.tr THT: i: C!:.!NEY
III. ._QfF (.Ti:· f'l'.'TfilC.AL
uo·
;:,l:En.ti1rIO~; u.:ic· Tl:!":WQRCt"!: .Al1 0~K 3RI'CrIIiG BJUT 1'H- ":VO- UJ::' 11111 OF il'F~ <'L-;J1C,..,
L De.uses of Bvolutb·
2.
.., '~
.
r.tco1e ind cf of Mu1~uta tatiti on on and. SelectL.rn in evolution203 203 207 211 216 219 22!
227
231 231 233 241
-v-
4. Selection and its ~art in Evolution •.• 249 5. Role of "Small" ,\ltutations and Selec-
tion in the Evolution of the Fleece. 251
~ECTION VI
I. THE CJ.:IliRACTES! OF LA B'S r100L
...
1. lntroducti0n ••·
...
2. Fineness ••. • •.
(ai Rcl~tionship bet~een finenes8 of ool and body re3i~n ••
(b) RelationFhi.p betAeen .il .. neness
of ool anc Fibre Tyre Array
tJ. .ia ve 4. vrimp
Summary
. . . ...
Biblvogre.phy
.Appendix
. . . ...
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...
254 254 258 258 260 262 2i:.;4
272 285 289
INTRODUCTION.
The original purpose of the precent thesis was to ansNer this q~eFtion: "To what extent i~ a britch of hi5h halo-hair abundance an indicator of the hon-kemp hairiness of t.he fleece?" It wa'"' assumed by ur. 1ry in the light. of earlier observation~ , that, on a britch ~ith very many halo-h~rE, the large Curly Tip fibres would be ver~ hairy, and thiP hae proved to be EO. Especially die we want to learn about the degree of hairiness on the part of the fleece near to
Lhe britch. The gradient over the body from the britch ·as also much in mind, a ll the more because t he boundary between the area on the britch with very ma-i y
halo haire: nn<l. the neighbourint;S region with far
fe _, e r ~ttl.o
hairf i f often abrupt.The aim of the \ ork wa"' that ,iust defined, but the analy~i~ of the samplec exs~ine~ provided in- formation on a number of other matter~ , several of which may be thought more interecting than the prob- lem proposed at the outset.. These var iouE ref ultE
&re reported in this thesis.
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In carrying out this work it wae inevitable that I ehould review the fac ts and ideas coGtributed
by earlier workers at the J.rarsey Agricultural College.
Their ~ork has all paesed through my mind. This
little band of . orker~, i t ha~ been pointed out to me, has never oeen large ana i t hae alwayc been isolated.
I have entered this school of thought from Profe~sor Prawochencki's Department of the Univer~ity of ~racoN
(Cracow, Poland) and have so,net imes found myself offer- ing interpretations "' lterre tive t o those put forward at t he Massey Agricultural Gollege. 'l'hese i deas have been welco ea, altholl6h thoce holding the older vie ,s
te ll me that they do not expecL to prove .• rong every time.
This will explain ~ hy I have dealt 'Nit h my
[ Ubject broadly and at come length. Not only has t'-:e method of presentation helped m, working ac u.intance with the clacs of fact and mode of thought of my new associAtes , but at the same time I have ~een encouraged dlo revie\ critically the conception'"' abJllt the fleece
hich I have encountered in New ~ealand.
./
.,,,-
I
/>
, (
-3-
,,..
);- ·"
Guided thoughtfu.lly by Dr. F .• ~ •• Dry, and his closest associate, Mr.J.A. Sutherland , who ~vere kind
enough to teach me the principles of ~ibre Type Arrays ,
I was surpri sed to find the sheep's birthcoat such a wondefful object f or a fundaments l snudy of manifold aspects of wool. one may think that the birthcoat
is a "microeosmos" of it"' own, and, if Jne '1as a Euit-
--
able method of approach the birthcoat - I am sure - Nill turn out to be an indispensable ob~ect for bio- logicc:.l studies.
•
Dr. J.Jry 's pi:ineering <r,ork has lready furnished not only the concept of approach, thiE being the ~ibre
Type arrays, bu.t alsJ the m""teriel for comp' rative study.
This work and line of thought have opened up quite a new branch of biologic· l science, hich c£ OP cr>lled ''Com-
parative anatomy of ~ool". Thie new diccipline - I am coLvinced - ~ill open a large field or i nvesLigations, which will help to get far deeper uncerr:tanding Jf rool, from the purely practical point of vie~. l>lfo, it d l l ,
to my mind , throw light upon such vivid problem~ as those of voldschmid'~ p~ysiological genetics, a~ ~el l as the question of evolution. Let me pe permitted to give but
one example. The pre-natal-check, as .e understand i t nJw, i s a force, which mainly cause~ fibres at the head
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of an array to be fine; e can distinguish at least three important properties offohe check, namely:
(i) the outset, that i'"' the time when the pre-natal check st rts t') • ork;
(ii) The intensity, thu.t i f. the trength of >-he check, w he t he r i t i c: po er f u l en? :ig h t o
cause
tm
fibre to be Line throughout or in the nee~ region only,and finally,
(iii) The extension of the check, that is, how far it extends in the Curly-Tip Group.
The pre-natal check i~ the rerult of the ·~ork of genes . 1'h i ~ ork we can 11eaL1re very accu.ra te ly. ..tror ins ta nee, {. In Valley arra.., s the outset can be meas .ired by the number of halo-hairs and Sllper sickle A f ibres, the intensity by number of fine ickle fibres end the extension by n~mber
of checked curly-tip fibres.
.e
cPn, fur thermore, ex-press the measurement ac: a pe ccentage of certnin fibre typer, thiE' being deci dedly more a vantageoui: and probably
G- more accurate, than arbitrary term~ used i o Goldschmidt's nvestigial" ca"'e in ~r')sophi la. lit seems to me that the work of gen" s causing "vest igiG. 111 in Lr:leo phi la, is very
7
<:; imi lar to the work of genes cau.~i ng the pre -na ta 1 check j
in the oirthcoa ti.
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I have dealt to some extent with the question of the evolllt ion of Lhe fleece in the Tew Zealand. Romney breed. 'l'he m~ terial for thi~ pc.rt of my thesii::: has been gro- ing Qurin5 the courEe of my inve~tigation, 1hk h inc.eed had n:)Lhing to do \ith the problem ofthe evoluti:m of the fleece and therefore I could hardly 1e accused of
"having an eye seeing . hat the eye 1001.cs for". Indeed, it wa~ too great a temptation not to use data obtained
in the course of the . re~en inve""tigatiJn in an atte~pt
to outline the ay l . hich the evolution of the fleece has probably taken place.
The ~tudy of evolutionary proces~es requires evidence from three ~oirces: PaleontolJgicaJ , embryo- 105ical, and 6enetical. 'l'hese S..>.irces are c.isc rnsed in my the~is, using perhap~ different wording. In p8~si~g ,
we may add, that by embryology of ool, L meant the developmental changes aE noted in the prenatal part of a staple. It i s felt , ho ever, hat c:.n exQlanation muc-t be given aE to the firi::: source. Dr~'s Experimental sheep could hardly claim to be paleontological examples.
Nevertheless, I feel at ease t) say that a~ fc:::r as paleo.a- tology serveE ~c a F0urce of evidence of evolutionary changes, ~ry's 1xperimental flock may be regarded as a 2ource of phylogenetical evidence of th~ ev)lution of he fleece, provided, however, that bQ one . i l l agree tJ regard
-6-
the Merino wool (being generally of lain array) as tt-e most evolved ~001 - th&t i , the most removed from the
~ild ~heap's coat, ana on the other hand, the mixed woolled !=:heep lBlackface, ..t'olish Cakiel, J-type- being probably at least of Plateau array} ae the la t evolved wool, that is, not too far removed from the wild coat.
This Jring so, ~e can claim to be in a very fo~t~na~e
position to have the 1paleontological" evidence not in the form of f.)"Eils, bllt in the form of live ~heep.
b~ to me, I have had the opportunity to 5e' acquainted with ~olish mixed woolled ~heep ac well a!': Merinos. The gap between them was too broad to bridge. The briage,
hot~ever, i'" to be fol10d in Dry'::: Experirnent::..l flock.
Indeed, it 1iar worthwhile to sail save.Bl ~eas, at leai?t, to find thf> ''lost IT links in the chain ·of the fleece's evolut ion. Yet , although my ne··. home is remote from Central .i!:urope, i L is interesting tD have it pointed ::>Ut
to me that the work in Which l have drnme to take part has roots in Vienna, in Told1s f ibre Lypes, and in m-
sterdam, in de ~eijere' fundan-:ntal~ of hair arrangement and development. I am told, besides, that he voyage from "'!i'urope should have been made by the 9ape, ·here
Dllerden'~ "birth thinning 11 ~oreshadowed the pre-natal check.
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My philoEophy, it is hinted t o me, occasionally showE signs of running ahead. of my factr. 'i'hi.. .. , if it be true, if deemed a mer~t in the department where l work, provided one oe a"are of the free rein given to imagina- ti )Il.
I t is a source of prid.e rather than shame in thi s department that a paper rnce submittea to ProfesC>or
'.IT. 3 • • Haldane in his capacity of Editor of "The Journal
of Uenetics" was refused on the score of being too specu- lative.
~peculation i s valued here for two quite dif- ferent rea~on~. In the f iret place ) fl present knowledge
the surest scoring shots in livestock breeding - one
naturally thinks of progeny testing - are apt t o be clum0y.
Deeper unleretandi.ng may make defter strokes possllble, and this understanding will be gained only thro~gh lively
imaginati on.
Speculation i s al so welcomed because adventurous thinking i s essential if our mater i2l is to yield some con- tribution to fundamentnl ge rnticE. 'hen indeed one reflects how villing to speculate ab ut evolution comparative
morphologiEts have been on les~ explored materi~ l, thie provides our justification. '1' o be ~ ur e , t he v a 1 ue of
/
-8-
such phylogenetic '"'peculation has ce en called into auef:"'tion by Bateson himfelf. Yet ~hatever a paleontologist can show cause to believe has taken place, it ir the businers of ' he geneticist to discover the genetic rrincipleQ therein involved, and it i~ a very similar sort of thing that is bei ng attempted i th the manifold variationc revealed in the coat of the "'heep.
The importance of the pastoral inau~try attracted At tention to ool rese~rch. Hence the intensive study of the fibre typeQ of the Romney coat and their development . Once thi s investigation wa'"' well launched, the conviction came to prevail Lhat this class of facts, ~erving a a ba is for the plannin~ and interptetation of experimental breedLig , WJuld oe of value to pure genetics. That this
is not an unreaQonable probabi l ity,as pointed Jut before, is apparent in this thesis.
At the end of this introduction, I feel it is my duty to apologise for having written this thesms in very poor Eng lish. In spite of having ~tarted to learn EngliQh before being taught 11hair-Qplitti ngn i t haQ turned out that I m a far poorer linguist than a "hair-rp1itter n.
I hope indulgence ~ ill be granted me al l the more, if one is not apt to lay " •••••.•. blame on t.be pet r-tree for not bearing plums". (Alain).
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