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MASSEY AGR!CU!... TI.JRJI.L COLLEGE 
LIBRARY PAUKi�STON NORTH, N.Z. 
ENERGY l\1ETABOLISM, RANGING BEF..AVIOUR 
AND 
H��TOLOGICAL STlmiES 
WITH 
ROMNEY MARSH AND CHEVIOT SHEEP 
being a thesis 
presented for the Degree 
of Doctor of Philosophy in the 
University of New Zealand 
Eric Cresswell 
Massey Agricultural College 
University of New Zealand 
19.58 
ACKNOWIEOOEMENTS 
This work was carried out during the tenure of a Senior 
Research Fellowship of the Research Committee of the University 
of New Zealand. Financial assistance was also supplied byMassey 
Agricultural College. 
GratefUl recognition is made of the generosity of the Nuffield 
Foundation and Cooper McDougal and Robertson Ltd., England. 
It is desired to extend particular thanks for all advice, 
support, assistance and facilities to the following:-
Professor G.S. Peren, C.B.E., Professor A.L. Rae, Mr1 M.W. 
Webster, Dr. C.R. Barnicoat, Miss M.G. Campbell and the Library 
staff, Mr. C.H. Fairless and Mr. T. Crotty, all of Massey Agricultural 
College. 
Grateful acknowledgement is made of the generosity of the 
Medical Superintendent, Dr. J.H. North; the Pathologist, Dr. T.H. 
Pullar; the Principal Bacteriologist, Mr. H.E. Hutchings and the 
Instrument Maker, Mr. G.D. Kemp of the Palmerston North Public 
Hospital. 
The counsel of Professor R.W. Dougherty of the University of 
Cornell, U.S.A. and Dr. A.T. Phillipson of the Rowett Research 
Institute, Scotland was a source of great enco�agement. 
The typing labours of Mrs. A.'w. Warren, Massey Agricultural 
College, and Miss s • .Ai+en and Mrs. G. Montrose of Utah State University 
are no less appreciated, 
CONTENTS 
List of lllustrations 
List o£ Table 
Division "A" Background to project 
"B" Energy utaboli studies 
Addend\111 
"C" RaJti1nc behaviour studies 
•n• B:aematological studies 
',:• 
•E" Diacussion and conclusions 
ot the overall project 
{DiV'isions "A", "B", "C", "D") 
"P'" Bibliography 
Appendix A 
Appendix B 
(In sections corresponding to project 
divisions "A", "B", "C", "D", "E") 
Pages 
1 • iv 
V- vi1. 
1 - 26 
27 - 101 
102 - 130 
131- 168 
169 - 191 
192- 204 
20.5 - 214 
21.5 - 226 
227 - 2J6 
LIST OF ILLUSTRATIONS 
Page 
Figure I. The shoulder pro£iles of 10 South Country 7 
{Bord r) Cheviot and 10 North Country Cheviot 
ewes at Sourhope hill research station, 
· 
Scotland (Wan.'lop 195/b). 
Figure II. Total 24-hour basal heat production referred 31 
to body weight - sheep. The solid dots 
represent a u. s. A. cross between t.he 
Sout.hdown and Rambouillet and the hollow 
circl.es Au tralian Merinos (Benedict 1938b). 
Figure III. Showing (1) weather data, (2) average 161 
distances traveled in miles per week for 
Cheviot and Romney Marsh ewes • (3) periods 
and locations, (4) dates or recording 
periods. 
Plate 1. Typical Border Cheviot ewe. 
2• Typical Romney Marsh ewe. 
3• Cheviot Hills 20/6/);.6. 
4• Romney Marsh 31/7/54. 
.5• Cheviot Rllls 16/10/4.5. 
6. Romney Marsh 19/4/55· 
1. • Field • pattern tracheotomy tube (a), w1 th 
introducer (b) and rubber balloon (c). 
8. x ... ray plat showing inflated balloon occluding 
the upper tr che of a sheep. 
9• Sheep befor th insertion of a tracheal balloon, 
4 
4 
15 
16 
17 
18 
39 
41 
(a). 42 
10. Sheep after the insertion !llld i.ntlation of a trachea1 
balloon. ot .. pUot balloon (a) connection tube (b). 42 
1 
Page 
Plate 11. Traebeal oannula with corrugated rubber cuff (a). 44 
12. A ap1ra:uter recording in progren usina the 46 
mocSU'ttcl 'Field' pattern traoheal cermUla. (a) 
Exte:rnal •eal.ing pa4a, (b) SpirCMter, (e) Two 
Wa7 'I' lve COI"l"elponding te the ant ul t s noatrtla. 
13. Modified 'Jleld' pattern traoheot� tube with 
tracheal support• (a) and (b). 
14. Modified •Field• pattern tracheotOJO" tube with 49 
rubber sheath (a} nd introducer cones (b) and (c). 
lS. Connection tube for linking a beep to CDf'gen 
ttPlrGIII.eter. 
16. •r• cannula open-showing (a) Grocwe4 edge, 
(b) C.pUlary tube, (o) Roll-on rubbet' cuff, 
(d) ubber extension tube, (c) Pilot balloon, 
(t) Cutting edge to fit groove (a), and Cbamte!"ed 
tube (c). 
S2 
17. •t• cannula closed. 54 
lB. A aptr-eet.er rec4rd1ng 1n progr ss using the •r• 56 
cannula. Not the lack ot external sealing pads 
on the sheep' n ck. 
19. lnterior of •be p shed. 62 
20. Typical �gen spirometer graph. Note the 66 
downw&J'd bUps •x• produced during eructation. 
Tbeae blips are d e to eea•t1on of breathing 
abd not to tbe entry of eructated. gas into the 
apb'C��eter. 
21. Met•boU . charts 1llu8trating:- 1. The lack of 67 
etteet of pr-olonged tabollsm recording on tbe 
reapiratar,r �tt m and rate ot �gen oonaumption-
2. The repee.tlb1Uty or recordingat ohart B •• 
ilad• 8 Jdnutee after cbart A; ). Enetation 
blip •x• . 
22. The effect of stan tlon on the reap1rat017 pattern 75 
�f a Cbertot ...  
u. 
Plate 23. The energy metaboli experiment sheep 1/5/57 
(prior to slaughter on 8/5/5?). 
Page 
77 
24. The respiratory pattern of a sheep before and 106 
a.t"ter shearing 
25. The respiratory pattern of a shorn sheep before 108 
and arter wetting• 
26. The respiratory pattern of an unahorn sheep before 109 
and after wetting. 
21. Oqgen spirometer graphs made with an eight minute 110 
g p between recordings using the same sh ep. 
28. A tracheotomised calf .  127 
29. The respirator;y patt rn of a ruminating sheep. 128 
The points of regurgitation are marked by the 
larg upward blips "X". 
30. Showing details of the design and construction 14 3 
of the Rang eter. 
31. Showing details of the design and construction 144 
of the Rangemeter. 
32• Showing detail of th design and con truction 145 
of the Rang eter. 
J3. Showing details of the design and construction 1..46 
of th Rang eter• 
)4. A vi w of tb holding paddock, Tua Paka, tak n 1.50 
from the central ridge. 
35• Range ter flock sh ep ustered for mileage 152 
recording to b not d• 
)6• h-ont and rear views ot sh ep in 
harness• 
emet r 
37. Ranney Marsh nUl" sing lamb while 1n hamess. 
111 
15) 
151� 
Page 
Plate )8. Rang eter flock ewes on their way out to the 155 
hill after mustering. 
39. Rangemeter flock ewes on the bill. 156 
40. Reading the Rangemeters. 157 
41. Measuring the height at withers. 
42. Blood dye injection. Note the sh ven neck. 
iT 
157 
182 
LIST OF TABLES 
Table l. G. B. Meteorological Office weather data 
(average figures) for the recording stations 
nearest to Romn y Marsh and the Cheviot Hills • 
Page 
19 
2. Chronological events--energy metabolism sheep-- ?0 
19/9/56 to 8/8/5�. 
). Showing for the energy metabolism xperiment ewes: 85 
a. M an llveweights in kilograms (exp riment Blocks 
I • VI). b. Square metres computed surface areas 
(experiment Block I .. IV). c. Final liveweights 
in kilograms. d. Square metres measured surface 
area. 
4. Showing (1) Th breed means and St. D1s per 86 
experiment for o�gen consumption in 11 tree per 
five minutes. (2) The F. values derived f'rom 
between breed analyses of variance (Appendix A, 
tables 8, 9, 10, ll) carried out on the oJcygen 
conswnption data related to the following bases; 
{a) Animal, (b) Kg. body weight, (c) Sq. metres 
surface ar as(computed), (d) Kg. body weight to the 
power o.n. (3) Significance of dif'f renoe between 
breeds. ( 4) Mean body weights for periods 1n 
ld:klgrams • • 
5. Showing (1) The lamb production data of ewes 1-10 88 
in the energy taboll experiments. (2) The 
�sis of variance between breeds for 1 b rate 
of gain. 
6. The standard d viations of the br ed mean for o2 90 
consumption Within blocks xpr�ssed as percentages 
of the m ans (to n arest �). 
?. Showin ·•figur s in the lit rature and those 98 
calcul ted tros the data in 'his project tor the 
cal.Qries/24 hour heat production of sheep• 
8. Carcass and ol"' an data. nergy etabollsm exper1- 99 
nt • ea. 
9. Ha tological data of nergy etabolism xperiment lOO 
ewes ?/S/S7. 
T 
Pag 
Table 10. Showing the o:x;ygen consumption records (litres 10? 
o2/ S minutes) of' eleven she p; (l) Pr and post snear1n , (2) Pre and post wetting af'ter shearing, 
(3) Routine recordings with an eight minute time 
interval, ( 4) One f'inal routine recording. 
ll. The mc;ygen con umption records in litres consumed/ 114 
five lllinutes of three sheep each fed l lb. oats after 
twelve l urs tarvation and a ain eight hours later. 
12. The f'f'eet of' thyroXin tr tment on the oxygen ll? 
consumption, respiration rate d liveweight 
of one Rclnney Marsh ewe. 
13. The results of analyses of' gas samples drawn 120 
after the sh ep had exhausted half the resevoir 
capacity of a closed circuit oxygen spirometer. 
14. The daily distances walked by different heep 134 
breeds singly and in variou combinations (Louw 
n. AY:.· 194a>. 
1.5. The total d1 tance {in yarcl ) tr veled by ach 135 
sheep during tw nty four hour period {England 
19.54). 
16. Peroenta e t e p t r sting 1n the hours 7:00 136 
a .m. - 7:00 P• .• by lowland and hill beep, 
April - Sept ber (Hunwr 19.54). 
1?. The av rag claiq distance walked by tour sh ep 136 
br (Van R nsburg 1956). 
18. et r beep weights 24/8/57/ 158 
19. M11 a r cord , Rangem t r sh p, 28/7/56 - 160 
2A-/8/57. 
20. (1) Avera � gr t r w ekl.y mileage travel 16) 
by Cheviot over ey she p, within periods. 
( 2) Signiticanc of ditf'er ne in e kly 1 g 
bet n bre ds within periods u.si k 
(J) SignU'icanc of ditt renc b tw en we 
within periods within breeds. 
vi 
Page 
Table 21. Le�els of significance of difference in weekly 165 
mil ages between consecutive periods within 
breeds using sh ep means. 
22. Comparison of foreleg measurements, H�ematology 168 
and Rangemeter sheep. 
2). Comparison of height at withers measurements 168 
( unshom sheep) von Borstel (1951) and HaematolC?gy 
flock this project. 
24. The distribution of hi h potassium �alues in the 17J 
red blood oells of British breeds of sheep (Evans 
and Mounib 1957). 
25. Summarising data in the literature on haematological 174 
values for heep. 
26. S arisin data 1n the lit rature on blood volumes 176 
or she p (Hansard � �. - 195J). 
27. The general production data of the Ha tology 185 
flock. 
28. Haematologieal data of the Haematology flock 8nd 186 
the results of analyses of �ariance carried out 
cm the sam data. 
29. Blood volum and r levant data-Haematology flock, 189 
February/ 57. 
JO. The m an concentrations of.Potassium in tb whole 191 
blood and red blood cells, Haematology flock J/10/ 
56, also data ex Evans (19.54). 
DIVISION "A" 
BACKGROUND TO PROJECT 
CONTENTS 
A-I Introduction 
A-n Review of 11 terature and some discussion of it 
A-III Breed historie 
(a) The Romney Marsh or Kent Sheep 
(b) The Cheviot Breed of sheep 
2 
Page 
8 
11 
A-IV Topography, general aspect and climate of Romney Marsh 14 
and the English/Scottish Border 
A-V Discussion 20 
A-I INTRODUCTION 
Youatt (1776-1847 a) in commenting on British breeds of sheep, 
writes:-
Hin all the different districts of the Kingdom we find various 
breeds of sheep beautifully adapted to the locality which they 
occupy. No one knows their origin; they are indigenous to the 
soil, climate and pasturage, the locality on which they graze; 
they seem to have been formed for it and by it." 
Some present day students of animal husbandry are now examining 
the characteristics of these various breeds in the light of the par-
ticular environment in which each originated. Their object is to 
3 
discover what characteristics can fairly be ascribed to particular sets 
of conditions, or in other words, what possible functional adaptations 
to environment can be revealed by study of this unique collection of 
soil stable breeds. 
Interest in the background of the development of these breeds and 
its possible influence on them is not. howev r limited to the United 
Kingdom as British breeds of livestock have been taken to all the cor-
-
ners of the world in the wake of the migratory ovements of the British 
people. For example, in New Zealand, at the time of writing, attention 
is being focused on the Romney-Cheviot crossbred ewe which is competing 
against the Romney Marsh for certain hill country and this has prompted 
consideration of the parent stocks (Plates l and 2) from the standpoint 
of their original habitats. This project was undertaken to augment 
what has already been done i.n New Zealand on this subject. 
MASSEY AGRIC:Ul TURAL COLLEGE 
LIBRARY PALtJIERSTOi\J NORTH, N.Z. 
Plate 1. Typical Border Cheviot ewe. 
Plate 2. Typical Romney Marsh ewe. 
4 
A-II REVIEW OF LITERATURE AND SOME DISCUSSION OF IT 
5 
von Borstel (1951) studied the forequarter anatomy of samples of 
ten animals from each of the Romney Marsh and Cheviot breeds. He found 
that the Cheviot has comparatively sharp, pointed withers and rather 
sloping shoulders and that, while the shoulder muscles believed to be 
concerned with the flight phase of locomotion were very much the same 
in both breed types, those believed to be concerned with the work 
phase ( producing locomotive power) were significantly heavier in the 
Cheviot group • . Yon Borstel views these breed differences as having 
possibly been brought about through the Cheviot, a hill breed. deve­
loping shoulders for active hill work that the Romney, a lowland breed, 
would not require. He contends that locomotion studies, linked with 
behaviour investigations, would enlighten the subject of anatomical 
form as related to adaptation. 
Unfortunately however, for the student of functional anat�, the 
agricultural literature abounds with circumstantial--but nonetheless 
concrete-�evidence to the effect that sheep of many British breeds have 
had their b� conformation changed to the breeders' concept of what it 
should be in place of what it was. Bakewell (1725-1795) pioneered 
odern alterations in sheep breeds and his disciples have been numerous, 
to s� th least of it, the yardsticks of desirability being supplied 
by the wool merchants and the butcher and his customers. Thus it would 
appear that lowland breeds as well as hill types may have originally 
possessed high, and/or light shoulders. 
6 
That high shoulders in sheep should at the present time be indi­
cative of an active disposition is also questioned by Wannop (1957 a) 
who writes:-
"The early breed society (Cheviot) statements on type said that 
the withers should be high enough to give style but not so high 
that there was any indication of slackness of back. Nowad�s, 
little if any reference is made to this featm·e and there is 
certainly no special emphasis placed on the high withers or any 
desire to accentuate them. They occur in the breed, but the 
desire to avoid slackness in the back would, if�ything, tend 
to reduce the' wither height. Re. our newly shorn North Country 
and South Country (Border) Cheviots run together on the same 
hirsel• on Sourhope. Both are equally prominent in the withers, 
but the South Country tends to have a more rounded shoulder and 
the North Countey a more pointed shoulder. From this one would 
expect the North Country to be more active whereas in practice 
it is the South Country which ranges more freely to the top of 
the hill and, unless herded, the North Country would keep to 
the lower slopes. The Scotch Blackfaced sheep tends to be more 
level from neck to tail. In practice it has to be more agile 
than the Cheviot, grazing steeper, higher and more rugged hills." 
-· \. 
Wannop(l957 a) confirmed the more pointed nature of the North Country 
Cheviot's shoulder by making a number of shoulder moulds according to 
the method of von Borstel (1951). Scaled down by �iannop, these shoulder 
moulds are presented in Figure I. They show that the South Country 
Cheviot does tend to have the ore rounded shoulder. Att mpts to re-
late present day anatomy to the particular environments of British 
sheep breeds are therefore beggared fro the very start by the unrival-
led enthusiasm with which the United Kingdom animal breeder has unques-
' 
tionabl.y remoulded the native stocks. For this reason the writer 
• Pasture divisions. 
Figure I. The shoulder profiles of 10 South Country (Border) Cheviot and 10 North Country 
Cheviot. ewes at Sourhope hill research station, Scotland (Wannop 195'b>. 
-.J 
decided to look for some other approach to the question of how ances­
tral background m� still affect New Zealand Romney Marsh and Cheviot 
sheep. In seeking for clues to what m� be basic differences, other 
than conformation, between the two breeds recourse was had to their 
breed histories. 
A-III §REED HISTORIES 
(a) The Romney Marsh or Kent Sheep 
8 
Romney Marsh occupies the South East corner of Kent. The 
district includes not only ROliUley Marsh proper, 24,000 acres; but 
also the adjoining and similar areas of Walland Marsh, 15,000 
acres, the New Romney Level, 1,500 acres, and Denge Marsh, 9,000 
acres, making altogether roughly 50,000 acres in the County of 
Kent. There are also approximately 9,000 acres in the county of 
Sussex, a total of 59,000 acres. 'The Marsh' is a marsh in name 
only. Drainage has made the land fit to graze and plough, and it 
has become a farming area of first class value although the bulk 
of it lies below the high water level of spring tides. It is pre­
eminently a sheep grazing area and carries so e of the best sheep 
pastures in the world; but there has always been some arable land, 
the ount varying with the prosperity or ·otherwise of ar bl 
farming. The Marsh is isolated but has a present dB¥ population 
of 50 to 500 people per square mile (Bartholomew 1953 a). Generally 
9 
speaking, the land is flat, large areas being almost treeless and 
without hedges to break the force of the wind. 
Fields in the Marsh are divided by water dykes which carry 
off the drainage w ter into sewers, the sewers conveying the water 
either into the Royal Militar.y Canal or into the sea at low tide. 
The tendency is for the fields to be rather larger than in most 
parts of Kent. A fair average figure may be fifteen acres with 
very many in the 15 - 20 • 30 acre range whereas on the South Downs 
the divisions would be in the five to fifteen acre range and on the 
North Downs seven to t n acres (Wyatt 1957). 
Taken as a whole the Marsh is inherently fertile and capable 
of producing not only excellent grazing but also arable crops com­
parable with those obtained in any other part of Great Britain. 
In summer the Marsh is very heavily stocked with sheeP--eight to 
ten ewes per acre. At the end of August or the beginning of Se 
tember it is customar,y to relieve the pastures of so e of the sheep 
by sending most of the lambs aw33', either to be sold in the autumn 
lamb sales or to be wintered until the following April on th uplands 
of Kent, Surrey, and Sussex or farther afield. Nevertheless the 
breeding ewes and wether tegs and ome of the lambs remain in the 
Marsh. Even in winter, therefore, the Marsh pastures are not 
really bein rested fro sheep--they are stocked up to capacity, 
usually at the rate of two and a half sheep per acr • The Marsh 
grazier likes to have a fair cov ring of grass on hi · pastures in 
10 
the autumn in order that he may have sufficient winter food for 
his ewe and wether tegs which find their own sustenance on the 
grass throughout the year, generally without the help of h�. 
Romney Marsh is the home of the Romney Marsh or Kent breed of 
sheep, several other breeds having been tried in the area but with­
out success. The breed has possibly one of the longest traceable 
histories of sheep in the United Kingdom for the isolated position 
of Romney Marsh {it was in earlier d;qs separated from the other 
�rts of the Kingdom by the vast Kent Wealden forest) must have 
assisted in maintaining the local breed pure and untainted from 
outside strains until comparatively modern times. 
In making particular reference to the Romney Marsh or Kentish 
sheep, Youatt (1776-1847 b) says that a long wooled and highly 
valuable breed of sheep has been kept on Romney Marsh from time 
immemorial. This breed demanded a pasture of unusual richness and 
found it on these Marshes. It was not uncommon for six or seven 
or eight f ttening wethers to b placed'�n one aer • 
Evidenc of interference with the original conformation of the 
breed through the introduction of the Leicester is also given by 
the s e authority (Youatt 1776-1847 c). He writes that th pure 
Leicest r would not be sufficiently hardy for the Marsh lands but 
they would aff et some valuable service in producing greater depth 
and roundness, symmetr,y of form, earlier maturity, and greater pro­
pensity to fatten. 
ll 
The 'Romney' is not only an inhabitant of the Marsh but, as 
its other name implies, was and is one of the main breeds of the 
whole county of Kent. The Kent Sheep as the Ronmey is known in 
that county generally, was bred essentially for grazing pastures 
and the li'ne of improvement chosen by the old Kentish flock masters 
was to hold grazing contests rather th� to ?arade the sheep for 
judging at show. During the earlier part of the nineteenth 
century many flock masters turned their attention to careful selec­
tion, rams from the best flocks being let out to hire. (Garrard 1954). 
A-III (b) The Cheviot Breed of Sheep 
The Cheviot Hills are part of an elevated and extensive range 
which extends from Galway through Northumberland and occupies a 
space of approxUnately 112,000 acres. 
If Romney Marsh can be referred to as "isolated" the Cheviots 
can be described as extremely remote. Roman rule was never per­
manently extended beyond Hadrian's Wall which lies some )0 to 40 
miles south of Cheviot and, fro the recall of the Legions to the 
seventeenth century, the area knew rel ti ve peace only under the 
Northumbrian branch of the Saxon heptarchy. Wllllam' s Doomsday 
recorders (1086) did not enter Northumb rland as, subsequent to 
his harrying of the North (1068), the area lay a desert* tor upwards 
of two centuries. After this both England and Scotland tacitly 
* An apt descript�on used by early historians. 
12 
encouraged the Border warfare which raged unceasingly between clan 
and clan, family and family, until the Act of Union (1707). This 
family warfare resulted in little intercourse between the outside 
world and the Border peoples. It can thus be reasonably assUllled 
that the sheep stooks of th area were free from extensive external 
influences until quite modern times. Some idea of the remoteness 
of the district is gained from the fact that Soot� (1771-1832) was 
the first an to tak a wheeled carriage into J..iddlesda.le, a large 
centre of Cheviot stock. At the present time the human population 
is in many places less than one per square mile with a maximum of 
fifty in the vicinity of small villages and townships {Bartholomew 
1953 b). 
Sir John Sinclair (Youatt 1776-1847 d ) described as follows 
the native sheep of the Cheviot Hills as they were in 1792 before 
the admixture of Leicester blood:-
"Perhaps there is no part of the Island where, at first 
sight, fine wooled breed is less to be expected than 
among the Cheviot Hills. Many parts of the sheep walks 
consist of nothing but peat bogs and deep morasses. Dur. 
ing winter the hills are covered with snow for two, three, 
and sometimes four months and they have an ample proportion 
of bad weather during the other seasons of the year, and 
yet a sheep is to be found th t will thrive in the wildest 
part of it. Their. shape is excellent and their forequarter 
in p rtioular is distinguished by such justness of propor­
tion as to be equal in weight to the hind one. Their limbs 
are of a length to fit th for tr velling and enable them 
to pass over bogs and snows, through which a shorter legged 
an1m eould not netrate. They are excellent snow travel-
lers and are accustomed to procure their food by scraping 
snow off the ground with their f et, ev n when the top is 
hardened. by frost." 
lJ 
Mr. Culley (Youatt 1776-1847 e ) reports the breed as, 
up·orequa.rter wanting depth in the chest and breadth both there 
and on the chine." Youa.tt comments, nThis might be correct as to 
the Cheviot of that d� but the system of crossing with the Lei­
cesters has remedied this def et." Here again is evidence of 
alterations in the external conformation of a native sheep. 
Conditions in the Cheviot Hills have changed little since 
Sir John Sinelair's time. Winter is still har3h with many long 
weeks of snow and ice and eaaterlies.which blow across the North 
Sea from Siberia often continuously from December to May. Hirsels 
are still large, ranging from lOO to 600 acres (Richards 1957), and 
the tru hill pasturage can in no wtq compare in quality, quantity 
and length of growing season with Romney Marsh. The average stock­
ing per acre in the Cheviot area is one ewe ' : two acres (H.F.R. 
1951 a). Ewe lambs are wintered on lowland farms the mature sheep 
being left to face semi-starvation for a period of some months. 
The doubtful quality of mueh hill herbage , the lack of arable crops, 
adequate shelters, buildings and enclosur s, l arsh climate and the 
remoteness of many farms ther for all continue to militate again t 
th less suited types ot sheep. 
The odern Cheviot 1 described by Thomas (194' a) as be1ng 
on of t.h ost important br s in Britain. Not only ha it a 
gr at regional importanc in the Engli h/::>oottish Border country, 
14 
but in Northern Scotland it is promin nt, and in many English coun­
ties it is found to such an extent that it can claim to be more 
cosmopolitan than any other breed thin the United Kingdom. Vast 
numbers of Cheviots invaded the chalk lands of Southern England when 
the reduction in ploughed land between the wars caused the giving up 
of Down breeds kept in arable flocks; they have even inYaded Wales 
and a colony has been established on Exmoor . Within the breed a 
strain possessing a bigger body, a larger head, a more open coat, 
and a requirement for more and better food has been evolved in 
Caithness (Thomas 1945 b; H.F.R. 1951 b). 
A-IV TOPOG HY GENERAL ASPECT AND CLIMATE OF ROMNEY MARSH 
AND THE ENGLISH SCOTTISH BORDER 
Plates J, 4, 5, and 6• illustrate the great difference in topog-
raphy and gen ral aspect between typical sections of Romn y }.1arsh . 
and the Cheviot Hills in winter and summer. 
Unfortunat � there are no meteorological stations located 
exactly in these areas but Table 1 summarizes th information on 
temperature, frost, snow, rain, sunshine, and wind for the station 
n arest to th required districts (G.B. Meteorological Offio 1957). 
In ach case th station nearest to Romn y Marsh is giv n first and 
the station nearest to th Cheviots i s oond. 
This tabl clearly d on tr t s th severitY. of climatic condi­
tions on the English/Scotti h Bo� r in comparison with those of 
•Sy courtesy of the Royal Air Force. 
15 
Plate 3 - Cheviot Hills 20/6/46 
Plate 4 - Romney Marsh 31/8/54 
16 
Plate 5 - Cheviot Hills 16/10/45 17 
18 
Plate 6 - Romney Marsh 19/4/55 
., 
Table 1. G. B . Meteorological Office weather data (average figures) for the recording stations 
nearest+ to Romney Harsh and the Cheviot Hills. 
Fo1kstone 
Kelso 
Folkstone 
Kelso 
Folks tone 
Kelso 
Folks tone 
Kelso 
Folks tone 
Kelso 
Folks tone 
Eskdal.emuir 
Lympne 
\.Jest Linton 
Lympne 
West Linton 
Dover 
Eskdalemuir 
DATES OF FIRST AND LA.T SCREEN FROSTS 
First Frost Last Frost 
Folks tone* 19th Nov. 21st Harch / 
Kelso** 6th Oct. 7th May 
Mean Temperature °F 
Jan. Feb. Mch. Apl. :4ay Jun. Jul. Aug. Sept. Oct. Nov. Dec. Y, ar 
40.0 40.) 43.1 46.8 52.7 57.3 61.4 61.6 58.9 52.8 45.5 41.7 50.2 
37.8 38.) 40.5' 44.2 50.1 54.9 58.3 57.5 53.2 47.4 40.) 38.4 46.7 
56 
59 
17 
1 
58 
59 
15 
-3 
67 
71 
12 
6 
76 
74 
27 
15 
Extremes of Temperature OF 
83 
81 
32 
24 
Maximum 
85 
86 
38 
31 
88 
88 
42 
37 
Rainfall in inches 
90 
86 
44 
35 
86 
88 
28 
75 
79 
29 
18 
63 
62 
27 
10 
59 
57 
12 
12 
90 
88 
l2 
-3 
2.25 2.03 2.17 1.66 1.68 1.99 2.10 2.39 2.37 4.03 3.25 3.21 29.13 
1.75 1.70 1.95 1.57 1.93 2.11 2.63 2.95 1.90 2.91 2.)1 2.32 26.03 
16 14 15 13 11 ll ll l2 ll 16 15 18 �.6 
17 15 18 15 14 13 15 18 13 18 16 17 15.7 
56 
37 
3.8 
7.7 
1.9 
82 
60 
4.7 
6.9 
2.4 
Average Sunshine (Total Hours per 1-!onth) 
138 
97 
2.8 
8.3 
1.4 
167 
126 
224 
167 
236 
171 
231 
137 
216 
125 
Number of Days Snow Falling and Lying 
Snm-1 Falling 
1.8 
o.1 
1.8 
0.1 
1.7 
Snow Lying 
o.o 
0.1 
Average Wind Speed 
l!.P.H. 
164 
99 
119 
80 
o.1 
1.2 
o.o 
0.8 
67 
54 
1.0 
3.8 
0.3 
3.6 
53 
36 
3.2 
6.4 
146.1 
99.1 
17.5 
41.7 
1.6 7.7 
7.2 39.3 
16.6 15.6 12.5 13.5 12.1 n.9 11.6 10.6 11.8 13.7 14.1 14.7 13.2 
12.6 13.6 13.2 ll.3 10.2 10.8 9-7 8.5 9.7 11.0 12.4 12•0 11.2 
+ 
* 
n-.e Station nearest to Romney Marsh is given first under each sub-heading 
.Height 128' 
** Height 1931 
Southern K nt. Derived from Table 1, the following facts are of 
particular interest: 
20 
(1) Both districts have approrlmately the same rainfall and num­
ber of rainy days but snow lies in the North on an average 
for J9.J days and only 7.7 days in Kent. 
(2) The m an y arly temperature for the Borders is only 4�. below 
that of Kent but the mean minimum temperatures show a wide 
difference, temperatures at Kelso dropping to below zero and 
showing only July and August free of frost. Kelso is at 
approximately the same h ight s Folkstone, Lympne, and 
Dover. The Cheviots range 2000 feet higher than Kelso and 
therefore must have an even more severe climate. 
(3) The hours of sunshine total for the year at Folkstone is 17.53 
whereas in Eskdalemuir there are only 1189 hours. 
A-V DISCUSSION 
An e�amination of the backgrounds of the two breeds reveals a 
number of basic differences. Ort · the one hand is the Romney, a long­
wooled, south-country, lowland sheep, accusto ed to rich pasturage on 
land which will carry up to ten sheep per acre in SWIIlller and two and 
a half sheep per aor in wint r. On the other hand is the Cheviot, a 
medi length-wooled, north-uountey hill and moorland sheep accustomed 
to tough living conditions on land which t it best will carry no 
mor than one sheep per acr in summer, While for a sheep to survive 
Border winter conditions it ust be able to withstand b� wastage 
21 
which m� run from ten to forty-six percent of its Autumn live weight 
(Cresswell 1951; annop 1957 b). 
"At the present juncture it seems that it is the extent to which 
boc1y wastage can be tolerated by the sheep that decides most of 
the problems concerned with the successful running of a hill 
flock." (Jones 1945 a). 
The rate of stocking of the Cheviot Hills and Romney Marsh indi-
oates that the CheViot may requir to coYer a great deal more country 
in search of food than the Romney. It must be admitted though that 
the system of husbandry followed in the Border hill country may force 
the Cheviot to range more freely than is necessary. The Cheviot• s 
pastures are large, running up to 600 acres, and the instinct of the 
flocks makes them move to the lowest point in the early part of th 
� and from there work their w� back to the highest point by evening. 
Thus the sheep may cover quite high mil.eages. per day just moving to a 
preordained pattern. Hunter (19.54) makes some interesting observations 
on this type of phenomenon in hill sheep. The average size of th 
fields of Romney )�sh (15 - 20 - JO acres) is certainly large enough 
to give opportunity for raking* but the conditions are not co parabl 
to the Border sheep walks. 
These differences in the background of the breeds suggest that 
they m:q be endowed with diff ring physiological mechanisms related to 
th ir ability to survive in their r speetiv environments and which 
the breeder ay not have radically altered in their selective breeding 
• WB.l.king and grazing. 
22 
for carcass improvement.asJones (194.5 b ) in writing on sheep and their 
enVironments comments·, "It is well known that it spells ruin evecy 
so often if the demands of the breed or type are bigger than the capa­
city of the land at its lowest;" and Nichols (1928) wrote:-
"The two chief classes of British sheep which can b distinguished 
are the mountain and lowland types, the latter cannot success­
fully contest for the natural habitat of the former, while owing 
to the development of the i'l.ying flock, the former, frequently 
and in la.rg.e numbers encroach upon the habitat of the latter. 
The distinction in type is complicated and embodies physiological 
and genetical differences which at the present tim cannot be 
expressed in strict t rms and therefore cannot be subjected to 
definit analysis�" 
The analysis of breed characteristics has still not yet proceeded 
far, and evidence is still lacking as to the precise nature of the 
breed adaptations to environment; but in the present project the ani-
mals and their backgrounds indicated thr e lines of approach which 
would be suitable for intensive study :-
L Energr Met.pholi sm • 
That characteristic which enables one animal to live wher 
another may die due to its inability to gather and to make good 
use of limited food of possibly poor quality• is a factor which 
is  obviously oonnected with th ov r-all metabolic processes of 
the animal as for xample :- Herbivorou animals differ in th ir 
ability to survive on the toughened d a·ioc ted vegetation of the 
desert• "The burro of the desert and th 11 of the arid puna 
thrive on provender which the cow and the horse are unwilling to 
t and possibly unable to digest" (Dill 19)8). 
23 
The Cheviot has been evolved in an area where the winter and 
even summer climate and pasturage can be classed as desert when 
compared with Southeast England and the breed is therefore recog­
nized as being able to make adequate use of a comparatively poor 
food supply. 
This may be accomplished in one of two ways: 
(i ) By an adaptation in th metabolic rate; 
( ii ) By an increased efficiency in the manner in which the 
digestive system deals with the food available. 
The last mentioned possibility was beyond the training of 
the author to deal with but the first was deemed to hold some 
promise and was examined further. 
Keyes !,1 !!• (1950) define "adaptation" as a useful adjust ... 
ment to altered circumstances. When the total metabolic rat 
decreases in starvation , as it indubitably does,  it is certainly 
a favourable change in that it reduces the caloric deficit as 
compared with what it would be in the absence of such changes in 
the basal metabolism. To the starving individual the reduced 
metabolic r te means that, at a given caloric intake, his rate of 
loss of strength and endurance is diminished and thus he will 
survive longer. 
The present day Cheviots 1 ancestors were subjected to compara­
tively severe seasonal starv tion each and very year and so far 
as it and the Romney are concerned it was th refore felt r asonable 
24 
to look for a possible breed difference in the effect of starva-
tion on their energy metabolism . However� the Cheviots • ancestral 
standard of living being at the best of times comparatively poor, 
it was felt to be reasonable to also look for a differential be-
tween the metabolic rates of the breeds on adequate rations. 
2. Ranging behayiour 
Ranging behaviour studies were initiated because the follow-
ing points indicate that the Cheviot is a much more active animal 
than the Romney. 
(i) The difference in field size between Romney Marsh and 
the Cheviot Hills. 
(ii) The possible need of the Cheviot to cov r more ground 
in search of sustenance. 
(iii) The comparatively more dynamic , nervous and intelli gent 
disposition of the Cheviot as opposed to the Romney 
Marsh sheep. 
), Haematology 
As it was hoped to show th t one of th breeds (Cheviot) was 
more active than the other it w s thought it :q be useful to 
include a study based on some accepted measures of uscular work 
capacity e.g. blood volume and haemoglobin. 
Brody (1945) states that:-
"Muscular-work cap city is d pendent on many factors such as 
soundness of limbs d muscles , on body' build and trength, 
on skill and intelligence, on temperament and ambition and 
2.5 
so on. These f ctors are more or less judgeable ext rnally 
by inspection. There are however other factors and aptitudes 
which are not externally evident, measur ble only functionally. 
These are concerned mostly with the oxygen supply to the 
tissues by the cardio-respiratory system. "  
Stamina, reserve power, endurance, appear to be directly propor-
tional to factors such as pulmonary ventilation, blood circulation, 
quantity of haemoglobin (quantity of blood) , oxygen capacity of 
the blood, and so on. 
In discussing factors causing physiological variations in 
plasma and r d cell volume , Reeve (1948 ) mentions uscular activity.  
He states that he knows of little work on the effects of prolonged 
muscular training on the blood volume but his impression is that 
p�sically fit men such as front line soldiers have a greater 
blood volume, particularly red cell volume, than civilians. 
The inclusion of haematological studies was also of value 
from another viewpoint. Ha atological data are used in every-
day clinical medicine whether with the hUIIlan being or farm animal 
a an indication of the physical well-being or otherwise of sub-
jects believed to be under physiological stress. 
Between the Cheviot and the Romney sheep on certain are s of 
hill country in New Zealand it is the Romney which is expOsed to 
th less favourable environment when the breeds are considered 
from the viewpoint of their ancestral habit ts. Under these con-
ditions the Romney is performing in a disappointing anner parti­
cularly in lazb prOO.uctio:t1 {rere� !lt al .. 1951 ) ,  and it was hoped 
that this would be reflected in the h ematological picture of the 
breed. 
26 
These three natural divisions of the project , energy meta­
bolism, ranging behaviou.r, and haem tologioal studies are now 
dealt wi th  in tu m. 
DIVISION "B" 
ENERGY METABOLISM STUDIES 
CONTENTS 
B-I Introduction and review of llt.erature 
B-II Techniques tor easur1ng energy M tabolis 
(a)  Principles 
(b) Technique development 
B-UI Pre-requisites tor nergy metabolism measurements 
B-IV Experimental arrangements 
B-V Experilllental d sign 
B-VI Chronological events : 29/9/56 to 8/8/57 
28 
Page 
)0 
32 
32 
37 
59 
60 
68 
69 
B-VII Bases tor presentation ot en rgy m taboll exp riment 79 
r sult.s 
B-VIII Re ults 
(a) Stat.istical proc dures 
(b ) Introduction ot new animals 
(c )  Body ights and surface area 
8) 
8) 
84 
87 
B-IX 
(d) Position in recording sequence 
(e)  Lability 
(f)  Br ed comparisons 
(g)  Carcass data 
Addendum 
( ) The effect of shearing and drenching w1 th simulated 
rain on the oxygen consumption and r spiratory pattern 
of sheep 
(b) The !feet on their metabolic rates of feeding one 
pound of oats to each of three sheep after twelve 
hour starvation 
( c )  Th eff et of 1 thyroxine on the oxygen consumption, 
respiratory rat and b� weight of one Romney 
Marsh ewe . 
(d.) Methane exhalation in sheep 
(e) Regurgitation in sheep 
29 
89 
89 
90 
97 
102 
104 
112 
ll5 
ll9 
121 
30 
B·I INTRODUCfiON AND REV IEW  OF UTEMTURE 
The main aim of this division of the project was to compare th 
energy metabolism of Cheviot with .Romney lo!arsh sheep under standardized 
oondi.tions of environment and feed over a p riod of approximately one 
year. Incidental studies which were added as the study proceeded and 
,, , 
opportunity presented are reported 1n the Addendum to this Division B.  
Benedict ( 1 938 a)  writes c- "Within a few animal species striking 
racial d11'fer nces in aeta'bollsm have be n noted. " Th.is author pre-
sents a oompr hensive summary of available knowledge on the subject. 
Brody ( 1945 a)  tound statistically s1gn111cant difterenc s between the 
resting metabolism ot some Holstein and Jersey heUers . Unfortunat ly 
Brody did little work with sheep and then only on a few Merinos prior 
to 1940 (Brody 1956 ) . 
Prosser � �. ( 1952 ) present a review of intra-specific differ-
ences in oxygen consumption which includes Drosophila and humans .  
However, as the present project is  primarily concerned with sheep , 
only the position a regards the beep is to be laborated on. 
Cited from Bened.ict ( 1938 b) the d ta pres nted in Fig. II repre-
aent two races of sheep, tho 1n Australia being the Merino and those 
at Durham, U.S,A . , a erose between the Southdown and the Rambouillet 
breed • 
Benedict ar es that the pronounc d labill ty 1n the bas .. t -
boli of thes anj.mals miaht well expl 1n the scatter of th d ta above 
Cal. 
1100 0 0 • • 
• 
0 
0 
1400 
0 0 
• 0 
1200 • • 
0 
• l> OA 
1 000 & 
00 0 
800 0 
800 
25kg. 1 0  5 5  e o  8 5 
Figure II.  Total 24-hour basal heat production referred 
to body weight � sheep . The solid dots 
represent a U. s .  A .  cross between the 
Southdown and Rambouillet and the hollow 
circles Australian Merinos (Benedict l9J8b) . 
31 
32 
and below the average curve , but lt cannot explain the difference in 
the two groups of sheep as a whole. Consequently he writes , "It 
appears that there is  a distinct racial difference in that the energy 
metabolism of the Australian race m� b thirteen percent lower than 
the Durham. " However, Marston ( 1948 a) feels that there was no re son 
for Benedict to assume a distinct racial difference. 
The probl in the comparison of the data from Australia with 
that in America is that the etabolism of the Australian sheep was 
measured after a 48 hour Jrast and the American sheep after a 24 hour 
fast . Also Marston argu�s that both bases for reporting the metabolism 
of these sheep are unacc,pt ble and insecure in the absence of a eo 
plete 1efinition of the �revious level of feeding and the nature of 
the fodder. 
B-II TECHNIQUES FOR MEASURING ENERGY METABOLISM 
(a) Principle 
The two jor biocalorimetric t chniques are designated as 
direct and indirect. Both direct and indirect methods were ori-
gin ted by Lavoisier in rrn • Lavoisier demonstrated that living 
involves oxidation ; he defined lite as a chemical process and 
introduced th essentially odern nomenclature especially as it 
rel t s to OxYgen in life proce ses. 
Direct calortmetr.y involves the me suring of the heat o -
put of an organi confined in a sui table chamber ( caloriJR ter) • 
33 
This chamber commonly takes the form of a well insulated box, 
the interior of which is kept at a constant temperature by water 
circulating through pipes attached to the lining. The heat ab­
sorbed by the water is computed from the amount of water flowing 
per unit time and from the tern rature differences between in­
coming and outgoing water . 
A current of air is drawn through the chamber to provide the 
subject with fresh air to breathe . This air is warmer when it 
comes out than when it enters 1 and consequently carries heat away 
with it . The amount of heat thus removed is calculated by- mul­
tiplying the volume of air drawn through the chamber by its den­
sity , by the differ ne in temperature between the entering and 
issuing air and the specific heat of air. 
A certain amount of heat is also carried out of the chamber 
in the issuing air in the form of latent heat of the water vapour 
exhaled by the subject. The weight of water vapour thus removed 
multiplied by the latent heat of vaporisation of water gives the 
amount of heat removed from the chamber in this form" These three 
quantities of heat add d togeth r give the total amount of he t 
lost by the subject. 
Other methods of direct calorimetry are reported by Benedict 
and Le (193? )  and Kurlin and Burton (1935) . The former authors 
pl ced the subject in one chamber while another similar chamber had 
electric-r sistant wir s ade to produce exactly th s e ount of 
heat as that emitted by the subject in the other chamber. The 
latter authors used a Pyrex glass cylinder for the respiration 
chamber. 
Indirect calorimetr.y is based on the fact that the heat out­
put of some organisms can be calculated from their 02 consumption, 
or 002 production ,and the caloric values of those gas s .  
The commonly adopted procedures in indirect calorimetry are 
therefore :-
(1 )  To connect the subject to an oxygen spirometer and 
measure its oxygen consumption . 
( 2 )  To collect and analyse the air expired by the subject . 
However , the caloric equivalent of 02 consumed .and C02 pro­
duced varies with the nature of the substance being oxidised. It 
is  therefore theoretically necessary to know the composition of 
the fuel mix (carbohydrate , fat , protein ) being oxidised . The 
relative amounts of fat and carbohydrate oxidised are determined 
fro the non-protein r spiratory quotient {R . Q. ). 
The respirator.y quotient is a te� pplied by Pfluger 
(Cantarrow and Trumper 1949 a) to the ratio of the volume of car­
bon dioxide expelled to the volume of oxygen inspired during t.le 
same interval of time . The value of the respiratory quotient varies 
with the nature of the foodstuffs metabolised , the determining 
factor being their rel tive content of hydrogen and oxygen. 
35 
In considering the relative values of direct and indirect 
ealorimetry, Cantarrow and Trumper (1949 b) write to the effect 
·that direct calorimetry. though the most accurate , is too costlY 
and impracticable for routine clinical use but indirect ealori­
metry is aff:i cientl:r acc!1r�.te to meet the ·acting requirements 
of clinical tests in man. 
Brody (1945 b)  considers that direct and indirect calori­
metry are equally simple in principle but since the direct method 
in practice is much more expensive and complicated it is compara­
tively rarely used for farm and other animals. 
In addition to high cost and complicated procedure , direct 
calorimetry has the following disadvantages : -
( ! )  The long period required to put each animal through the 
respiration chamber procedure . 
(2 )  With ruminants , the heat of fermentation in the rumen 
cannot be distinguished from true body heat. 
(J) The lack of control of the animal. 
(4) Possible p�siological upsets stemming from the psycho­
logical shook of confinement in a totally enclosed chamber. 
Indirect calorimetr�y offers the advantages of simplicity ,  
speed of op ration, low cost , eas of control of the subject , 
1 ck of psychological upset , lack of complication from the heat 
of fermentation and adaptability to a variety of studies. Howev r ,  
indirect calorimetry (which for th e  above r aeons was the method 
adopted in this investigation) also has its difficulties which 
must be understood or surmounted . e .g . : -
Brody ( 1945 c )  argues that i t  i s  not necessar.y and often not 
even advisable to employ the R. • for estimating the metabolism 
of ruminants by indirect calorimetr.y because it does not always 
have the fundamental significance given it in human medicine . 
Thus cattle and other ruminants produce huge quantities of C02 in 
the digestive tract by anaerobic bacterial fermentation and by the 
liberation of co2 from bicarbonates .  Thi extra metabolic co2 can ­
not be distinguished from the respiratory metabolism C02 • Under 
these conditions the R.Q. has no metabolic significance and of 
course the quantity of co2 production cannot be taken as a measure 
of metabolism.  
Nevertheless Brody (1945 d)  points out that although the 
range in caloric equivalent of C02 is relatively wide , from 5 .0 to 
6 .7  G al s . per litre , the range of caloric equivalent of o2 is 
relatively narrow , from 4.7 to 5 . 0  Cals . per litre , and is  within 
the limits of experimental error in met bolis measurements . 
Furthermore , since the average R.Q. of protein is 0 .82 which 
corresponds to the average c lorio value of 02 of 4 .825 Gals . per 
litre no correction need be ade for protein metabolism when measur­
ing energy metabolism from oxygen consumption. 
Within indirect c lorimetry the easur ent of o�gen consump­
tion is more easily accomplished than the me surement of C02 
production . In view of the above , the measure of metabolism 
adopted for this investigation was oxygen consumption . 
B-II (b)  Technique Development 
37 
For the pUrpose of connecting ani!.Jals to oxygen spirometers 
Brody ( 19)0 a )  developed a face mask and others including Dougherty 
� .!!• (195.5 ) have used cuffed rubber or metal endotracheal tubes . 
The face mask suffers from the disadvantage that although 
eructated and exhaled C02 is easily removed from a closed circui t• 
by a reagent such as soda. lime methane is not . In addition , the 
production of methane varies within wide limits and is particu-
larly voluminous when the ruminant is on full feed (Pilgrim 1948 ) .  
Brody (1930 b )  overcame part of this problem by firstly starving 
the animals for twelve hours,  when methane production had fallen 
to a low level , and sidestepped the remainder of the problem by 
simp� discarding those graphic records which showed the libera-
tion of such gases .. In this project it was desired to measure 
the 02 consumption of sheep on full feed. The face mask was there­
fore ruled out because of the methane probl m .  Later events were 
to indicate that this was a correct d oision as ; when the sheep 
were on full feed; eructation took place ver,y frequently during 
etaboli r cordings • In add1 tion the length of tim required 
for dealing with a number of animals made it practically impossible 
to carry ut many repeat runs .  
•As oppos d to an open circuit where the expired gases do not 
re-enter the spirometer . 
Connection of the pulmonary system direct to an oxygen spiro­
meter overcomes the methane problem in that it prevents the con­
tamination of pulmonar.y by rumen gases, and it was therefore 
decided to approach the tecnnique problems from this angle. The 
use of cuffed rubber or me t �:l  tubes was , however, rejected 
because the author was advised that over long periods they were 
most inconvenient pieces of equipuent to manage . ( Colonna 19.56 )  • 
It was therefore decided that before any long term experi­
ments could b� pl�nnP.� it would be advisable to devise a new tech­
nique for making direct contact between an oxygen spirometer and 
the pulmonary system via the trachea of an animal. 
Subsequently the experiments were started ( 27/7/56 )  by 
fitting two sheep* with 'Field' pattern tracheotomy tubes (Plate ?a ) 
and devising a piece of supplementary quipment to make this type 
of tube usable for such work (�late ?b ). 
The tubes were worn permanently and kept plugg except when 
r cordings were to be made. The solid plug was the replaced by a 
hollow (half inch lumen ) , soft brass, machine tapered plug linked 
to the spiro eter. The hollow plug carried with it a 2mm . ,  hard 
brass, by-pass tuQe which, extending some 6cm. beyond the plug 
itself, was bent in an arc corresponding �the curvature of the 
upper tongu of the tracheotomy tub so that the free end of the 
by-pass located within the lUl'llen of the upp r tracbe when the 
plug was in rted into the tracheotomy tube. 
• Under a general anaesth tic. 
Plate ? .  ' Field ' pattern tracheotomy tube ( a )1 with introducer (b ) 
and rubber balloon ( c ) .  
--· ------
0.. 
w � 
A deflated rubber balloon securely attached to the free end 
of the by-pass tube (Plate 7c ) was then gently inflated and clamped 
off after the insertion of the hollow plu • 
By this device (quickly and with a minimum of disturbance to 
the animal ) it was possible to complet ly occlude the upper trachea. 
Plate 8, an X-ray print in which the inflated balloon has been 
visualised in situ by means of radiopaque contrast �dium cle ly 
demonstrates the effective manner in which the upper trachea is 
blocked. To reduce irritation to a minimum the deflated balloon 
was lightly smeared with a non-oily analgesic jelly prior to 
insertion. Throughout the cour e of the investigatio the anal­
gesic factor used was a one p rcent mixture of ethocaine in a 
mucila e jelly . The insertion of the hollow plu with its 
attached by-pass tube and b loon was never in any way resented 
by the sheep . Its placing took only a few seconds and caused no 
more than an occasional cough due to irritation of the tracheal 
mucos,a. Plates 9 and 10 illustrate the complete i="ldiff rence of 
the sheep to t is procedure . Note that in plate 1 0  the pilot 
balloon ' a ' is inflated indicating that the animal is breathing 
through the connection tube 1b 1  which was us d to link the subject 
with a spirometer. As soon as the balloon was inflated the animals 
resumed feeding or rumination if these processes were inter��t�to� . 
Various types of balloon were tried . The most satisfactory proved 
to be the upper two inches of an ordinary latex condom which waG 
41 
Plate 8.  
X-ray plate showin 
inflated balloon 
occluding the upper 
trachea of a sheep . 
rlate 9 .  �heep before the insertion of a tracheal balloon � a ) .  
Plate 10. Sheep after the insertion and inflation of  a tracheal 
balloon . Note - pilot balloon ( a )  connection tube (b ) .  
42 
43 
gathered and bound to t h e  tip of the capillary wi th an elastic 
band . A condo.rn has an advantage over toy balloons in that its 
exceedingly thin but tough latex is easily inflated and it im.me­
diately bonds itself to the mucous membranes of the trachea , 
thus forming a perfect seal . 
However. it  was not sufficient to produce an air tight stopper 
�dthin the trachea for the design of the ' Field' pattern tracheo tomy 
tube meant that there was no ai r  tight joint between the stem of 
the tube itself and the sl , c e  1 s ne ck . rrohe problem of obtainin ,. :m 
a i r  tight seal in this location Ttlas only overcome after a con­
siderable period of experimentation . 
The wounds originally healed snugly round the t r acheotomy 
tubes and thus made a firm joint which in the case of the first 
two anLnalr- was effec ti v e ly supplemented by a supporting sponge 
rubber pad having a rubber ring behind it vrhich pressed onto the 
sheep' s sha.ven neck . 't.'o o l  grouth 3.11 the relaxin of the walls 
of the wound a\<�ay f r om the cannula, 1oweyer, very shortly brought 
a great deal of leakage trouble . Pads coated with temperature 
resistant grease were almost ineffective and their use possibly 
led to one case of pneumonia from grease contamination of the lungs . 
Three rines cut from one inch inte diameter corrugated 
rubber tubing ( Plate 11 ) and sprung between the cannula and the 
neck of tl e animal were most effective in preventing leaks for a 
n ber of weeks . Unfortunately this seal eventually caused more 
44 
45 
trouble than it was worth for it led to physiological pressure 
adaptations which sooner or later caused failure of the seal. 
Finallyi a long-wearing air tight seal was produced bet een 
the base plate of the tracheotomy tube and the neck by using a 
commercial rubberised putty smeared onto a sponge rubber pad. 
The rubberised putty made an air tight bond between the sponge 
rubber pad and the sheep ' s neck ( Plate 12a) . A sheet of rubber 
cut from a car tyre inner tube assisted the sealing process . 
The sponge rubber pad t ended to slacken its grip on the s tem of 
the cannula but t e inner tube rubber (placed a ainst the sponge 
rubber ) fatigued in its grip only after comparatively ve� long 
periods . 'I'he inner tube rubber l-Jas an indi spensible part of the 
seal . The oxygen s irometer used (Plate 12b ) as a }1cKesson* 
closed circu · t Hetabalor designed for human usage , which \�as 
secured on loan from the Palmerston North General Hospital . 
' ..Ji th this instrument a measurement is  taken of the amount 
of oxygen consume during a five or six minute period. The essen­
tial feature of the apparatus is a four litre oxygen container ( a  
rubber bellows ) geared to a kymograph . As the oxygen in the reser­
voir is  exhausted t e kymograph automatically records its consump­
tion on a quantity - time chart . Thus on one graph are recorded 
the 02 consumption in litres and tenths of litres and the time 
period in minutes and tenths of minutes . 1espiration rate and 
depth are also recorded . Deing a closed circuit apparatus it 
* McKesson Appliance Go . ,  Toledo , Ohio , U . S . A . 
46 
P�ate 12 . A spirometer recording in progress using the modified ' Field ' 
pattern tracheal cannula . (a )  External sealing pads , (b ) 
Spirometer , ( c )  Two way valve corresponding to the animal ' s  
nostrils . 
47 
included a soda lime container. \·Jhen the machine was in oper tion 
the char e of soda lime absorbed t h e  exhaled co2 for approximately 
forty runs before it needed changing . Re-charging Has carried out 
at this  point even thour;h the sheep may not have indicated the 
need for this by overbreathing . The metabalor had a built-j n  ther­
mometer and barometer . 
The efficiency of the seal between a tracheotomy tube and a 
sheep ' s neck ,J"as checked before and after each r.pirometer cecord­
ing by placing the thumb over the open end of the tube used to 
conne c t the tracheotomy tube tdth the machine . If the seal was 
perfect the animal "Jas unable to draw breath . A leak manifested 
itself as a hiss of inrus ing or outrushing air. The test was , 
by its very severity, able to detect a weakening seal , i . e . ,  one 
�•hich had given a p e r f e c tly nonnaJ. run but w.1ich would almost 
certainly l e ak  the next time if the rubberised putty was not 
changed. 
Good service was obtained fro� this arrangement and, in fact , 
almost all of the calorimetry work to be reported was obtained by 
its use . 
A weakness of the 'Field ' tube which only became apparent 
after the subject had been intubated for several weeks was a ten­
dency for the cartilagenous rings of the trachea to collapse . 
l eril1at ( 1921 ) gives a description of this type of occurrence in 
intubated horses . 
'l'he trach ototn,y tubes e r e  t er''!  'ora Mod i f i ed  bt �; ld c ri n 
onto t ei r tongue s h ·  f s u e t o n s  of copp.r tubing ( Platt. lJ a , b) . 
These alf secti 1ns com�letoo the ring sha1Je of the tr eh ·o tomy 
tube tongues 
the liOun • 
gave support to t. h e  upper nnd lower edg s of 
It  . l s o  tr>1ns�,ired th a t , after the ' F '  eld ' pl-\ t te rn t bes 
tad be n f i t ted fo r some woek s , tho tr· chea ... s h owed a. m11 rked 
ter�en y to cloo u e  t o  a n  e d e M � t o u s  condition o f  the mucou 
brane Juc:.t below the lo.rer tonbue o f  the tracheoto tube. 
1'his w.� undoubtedly brought a bo u t  by the i rri tatin[ f e e t of 
the un� rot cted e d g e  o f  t h e  t. r a o l  otoey tube working on t wall 
of the t ra c h  a when ver tho? shee , >  n t  i t s  he ad . . contributin · 
f a c t  r was i..>O S  ib y tho in c r  �ased n .g'!tive ;;res,; r s e r ted 
thin the tra.che;t to onable t.h :\n · 1'1a to draw ai r r: s t h e  
p r t i  o b s truction f o n  ed � the ��emat us ti s Gue s . 
T o  ov re e thia , �rob l urn  th e  lo•<� e r  a e c t i o n  o f  the tr eh o tom 
tube wa · sh athec t A. half i n c h  i n t c rn a  · dimueter -.oft rubber 
tub ( Plate 14 ) • T t e  ru b r e e s t. i l  g a ve troubl o t only 
after Ol"le l o rv' tirlle had e ap ea. • lo• •e ve r . 1en the o o::..  ng o f  
b c �  e. s  obvious to t h e  operator , it is � simp l 
t t e r  to lip on · 811ghtly lo ncc r ' i ec . of rubb r tubi nr . I t  
of sone ' v n t  e to fea t r t h  lo1·1er end or the rubb r tubin � 
to pr uc e  a. so f te r ed e. 
Plat. .. l J . ' ou '  f i e o  1 F : "' l <' ' pattern tr:::ch<i::otom tube with tracheal 
S'.J.t;)porL (a)  �ne: (b ) . 
Plate 14. Modified ' Fi ld ' pattern traoh oto y tube with rubber­
sheath (a)  and introducer cones (b) and ( c ) .  
49 
50 
Again i n  connection •i. th the ' l"'i e ld ' pattern cannula, the 
tee ni ue o r  connectin th shee to t h e  o x.yg@"...n s p i rorn e t  r 
ultimately l ed  to an ther mo fio�tion of thi s  piec of uipr'lOn • 
The re u · rements were for a r o th and rapi connecti n so 
that the sheep \�e re istur s l i  t. t l e  as r.ossible , normal 
breathir!g w�s p ·rrni tted an d  the ti e f r d alin . �J i  th ach 1 al 
; as no More than ten minutes whic ino udoo the spirom ter recording. 
1 s t-•rev o u s ly s c r ibed , t e smooth conn ction was ri.,inally 
ob 1 • ined by uo · ng a !rtach.tne ta ; r , hollo , h r a .. s pl g ilh i c h  as 
easily insertec i n to the lumen o£ the cannula. he t. per and th 
soft me ta l or th e �1lub 1:, v 1n �J. r tit•ht junction wi thout t..he 
exertion of mor than lir; t i'in,'4er pr ssure on th matin r of the 
two 1 .1icc • \Jnfortun.'\tel ' . hOWtlVo r , the re as a dr!l back to this 
arranger. ent in that the t 1pc rec1 ! l l ( \vhon f' t t.<;JC! • nt•) the lumen 
of the cannula) severely reduced the d net r of the tube t h rough 
ion t s �ocp drew i t s  b rea th . 
T ' me de oted to this ;o1nt resultc in a. gre&.t, .i.y improv 
t chni• JUe . To the outer f c e  o the cannula ·•as solder pr -
oision turned • hollow , so t braos on� Cl a t �  14b ) . Th intro-
duo r cone ( f a tc 14c ) wa . ach in to fit  exactly over the 
c ·  ul cone so th t an ai r ti  ht joint w a �  obtain wi th only 
s ight twist of the fingers . e 1 en of the cannu la was ther -
b left free of all obstruct! n other th the f i  , hnrd , br ss 
c pill y c a r ryinff th tracheal lloon. 
51 
The rapidity w· th �1hich it  was possible to connect up sheep 
after sheep to the oxygen spirometer owed much to the simple 
linking devices . 
In linking up the sheep to the spirometer another factor had 
to be taken into account . This �;as the question of "dead" space 
between the trachea and the two-way valve ( Plate 12c j which corres­
ponded to the .J.nimal ' s nostrils . The problem of dead space is  
directly related to the necessity of  avoiding a build-up of  ex­
haled co2 in the tube between the sheep and the two-'tray valve . 
Such a build-up of C02 results in overbreathing . 
Soft rubber tube of half inch internal di�eter was origin­
ally used to ink the s h e e p  to the spirometer . The slight resis­
tance to breathing created by the na.! ·o•11 bore of the tube however 
�ended to produce an overbreathing eff et d th  some sheep no 
matter 1.1h a t length ·;as used as an estima te of the dead spac e . A 
most satisfactory solution to the prob�em was obtained through 
the use of one inch internal diameter , corrugated soft rubber 
tubing as used \.Jith anaesthetic machines (Plate 15 ) . This tube 
�as 8 inches 1 ng an had an internal capacity of 80 ml . 
All of the indirect calorimetry work in this project was 
earrie out using the modified ' F i e ld ' pattern tracheotomy tube . 
iowev r ,  the fact th t a technique so potentially useful for 
indirect calorimetry work should be complicated by the likeli­
hood of leaks led to the final development of this  section of 
52 
53 
the project--a tracheotomy tube designed specifically for indirect 
calorimetry work . 
This tracheotomy tube (Plates 16 and 17 ) consists of three 
main parts, the upper and lower sections and a rubber extension 
with roll-on inflatable cuff. 
The lor,rer secti�n which is inserted first has five main 
features : -
A deeply grooved edge (Plate 16a)  and a capillary tube 
(Plate 16b) w hich allows the roll-on rubber cuff (Plate 16c ) to 
be inflated after the cannula has been inserted into t he trachea . 
The reason for making the inflatable cuff (Plate 16c )  of a roll-on 
type is one of economy, it being necessary to lengthen the rubber 
extensi :m of the tracheotomy tube (Pla 1 e 16d) whenever edematous 
thickening of the tracheal mucosa trclatens to  close the trachea . 
he rubber extension tube (Plate 16d) fits over and is welded to 
the metal tube by means of a commercial bonding paste . A pilot 
balloon ah1 ys within sight of the oper tor ( Plai e 16 e )  indic tes 
the state of inflation of the cu f hich forms a seal against the 
inner wall of the trachea . 
The upper section of the tracheotomy tube ha s two main 
features : deep cutting edge (Plate 16f)  which makes an air tight 
joint �lth the grooved edge of the lower section (Plate 16a) when 
the cannula is closed (Plate 17) ; and a chamfered tube (Plate 16g ) . 
Plate 1 6 .  "T " cannula open--showing 
( a )  Grooved edge 
(b ) Capillary tube 
( c )  Roll-on rubber cuff 
( d ) Rubber extension tube 
( e )  Pilot balloon 
( f )  Cutting edge to 
fit groove (a ) 
( g )  Chamfered tube . 
Plate 17 . "T " cannula closed . 
'-" .{::"" 
55 
The chamfer facilitates the fitting of the upper section of the 
cannula into position in the trachea. 
It  i s  proposed to refer to this tube as a "T" cannula and 
it is  intended to develop its use in future work for it  obviates 
the need for elaborate , costly, and time consuming seals between 
the base plate of the tracheotomy tube and the neck of the animal . 
Plates 12 and 1 illustrate the Field pattern and 11T"  
cannulae in use. Note the lack of external sealing p ad s  on the 
neck of the sheep in plate 18 . 
During the course of the metaboli sm experiments it  occasion­
ally became necessary to insert 1Magill 1 pattern cuffed rubber 
endotracheal tubes into the tracheas of some animals . At times 
a bent metal tube was used . The difficulties in the use of these 
tubes as forecast by Colonna ( 1956 )  were confirmed . \Vhen in use 
both tubes needed continual cleaning , there being no flow of 
tracheal mucus to keep the lumen clear . It was also difficult to 
prevent the open end of the tubes from becoming closed by a stopper 
of wool , hayseeds and ucus , especially as they had to be taped 
into position on the exterior of the sheep ' s neck . In addition , 
there was firstly a marked tendency for quantities of fluid to 
accumulate in the lmo�er half of the trachea >·rhen a tube had been 
in place mor than twelve hours and sec ndly the tracheal rings 
above the fistula collapsed within a few days . Long metal tubes 
had an additional disadvantage over the rubber tube . Continual 
Plate 18 . A spirometer recording in progress using the "T" cannula . 
Note the lack of external sealing pads on the sheep ' s  neck . 
57 
neck bending by the sheep rapidly lea s to abrasion of the wall 
of the trachea by the edge of the lower tongue of metal tracheotomy 
tubes . This results in edematous thickening of the tracheal mem­
branes followed by fibrosis and consequent narrowing of the l�men. 
Reference must now be made to the tracheotomy wounds . �Uite 
unnecessarily, as it turned out later, the first two tubes were 
retnoved and the wounds dressed daily until healing was complete : 
a period of about two weeks . 
The next group of animals had their tubes left in place 
until the stitches were removed . This proved a great success . 
Healing was rapid and no sloughing took place . he final policy 
wluch was adopted was to leave the tubes in place until the 
stitches were removed . A minimum of discomfort and. risk of in­
fection is thus obtained and this practice or preferably the use 
of dissolving sutures is recQ�ended to others who may contemplate 
work along the lines of this investigation. 
A point of note is that the first two patients had the corks 
omitted from their tubes for the first week or so to facilitate 
drainage of blood and any discharge from the trachea . l;.bat 
oc curred instead was a copious flow of slimy mucu"' whi ch coated 
the breasts of the animals with a noxious mess in about twelve hours . 
This flow of mucus is prob bly the trachea • s answer to th 
creation in it of an opening direct to the outside world. Pr sumably 
the flow is intended to prevent the entry of dust and other irritru1ts 
to the lungs and to provide material to close the wound until 
normal healing takes place . 
.58 
After all further operations the cork was inserted into 
the cannula within two hours of a sheep regaining consciousness 
and this proved to be a satisfactory procedure . 
It  was noticed at an early stage that if a tracheal cannula 
was removed for more than fifteen minutes or so it -vras very diffi­
cult indeed to replace owing to the powerful constriction of the 
muscles through which the fistula was made . vfuere a cannula was 
accidentally lost out during the night--as could happen if a 
fastening catch caught in the wire of the hay racks--the tracheal 
fistula �as completely closed by mornin and proved to be impos­
sible to reopen without the use of considerable instrument leverage . 
"'Ji th th e  facilities C�.Vailable in the shed allocated for this  
project it  w s difficult to maintain a high standard of  asepsis--
in fact the standard was quite low. Fortunately troubl. . s from 
infected tracheas were not met with until the last two weeks the 
bulk of the animals wer alive . A bloody mucus then formed in 
large quantities in the tracheas of six animals . This condition 
was unresponsive to treatment possibly due to the fact that it 
had long been the writer ' s practice to bolster the hygiene of the 
laboratory by greasin the rubber sheathing of the tracheotomy 
tubes with a suspension of antibiotics  in an aqueous base . 
Incidentally , this also overcame the need for lubricating the rubber 
tubes befor insertion , otherwise they would not sli e easily do\in 
the trache • 
B-III PR• -REQUISITE� FOR METABOLISM MEASUREMENTS 
59 
Not only the mechanics of calorimetry are of importance in metab-
olism studies , as for example , the psychological effects of unwonted 
interference vnth the subject may produce profound changes in the me-
tabolic rate . lso, when an animal is being regularly fed and the 
stimulating influence of food plays a large role , studies in metabolism 
must be carried out over a sufficient period of time so that the result 
corresponds to an entire representative day or succession of days of 
normal feed. 
itzman and Benedict ( 1938 a)  devote a great deal of space to 
dealing with the pre-requisites for metabolism measurements . In dis-
cussing the pre-requisites for energy metabolism studios i th ruminants , 
Ritzman and Benedict express their opinion that :-
"In spite of the large number of studies  nn other animals , in 
.Jhich comparable conditions of muscular �·epose and cesfiation of 
digestive activity were fairly readily obt ined , analysis of the 
literature on the metabolism of the ruminant, both early and 
late , shows that the pre-requisite conditions for measurements 
on the ruminant have been little understood by most investiga­
tors and the variable factors affecting such measurements have 
s e ldom been completely eliminated or standardised. 11 
In carrying out this project the vtri ter made every possible attempt 
to follow and to add to the principles laid down by these authors . 
These principles are aimed at eliminating or standardising : -
(1 ) 
( 2 ) 
( J ) 
�The activities connected with the consumption of food. I 
Extremes of environmeutal temperature . 
Huscular and nervous tension . 
60 
(For this pt·oject the following additional variables were considered ) : -
(4 )  Time of day. 
( 5 ) Position in the recording sequence . 
B-IV EXPERI�NTAL ARRANGEMENTS 
In the human, basal metabolism (otherwise standard metabolism or 
post-absorptive metabolism* ) is an expression used to designate the 
energy (heat ) output o f  the body at complete mental and physical rest , 
twelve to sixteen hours after the last meal . 
In the ruminant, hm.-ever , it  may be ar ed. that a similar post-
absorptive state possibly does not occur for days after the last f e ed  
(Forbes � �· 1926 ; R · tzman and Benedict , 1938 b; Marston , lj48 b) . 
In any case sheep on the hill or any type of normal pasture are not on 
a restricted diet--they have food before them at al i  tlours of the day 
and night . The oxygen requirements of the animal on full feed is  there-
fore one o f  some practical interest and so it  was decided that this 
project should be based on continuously fed animals . 
This project being concerned with hill country Cheviot and Romney 
Marsh ewes the experimental animals were selected at random from stock 
carried on Tua Paka, one of the hill farms attached to � assey Agricul-
tural College . 
Mature e es were chosen in order that growth effects could be 
eliminated from consideration . Pregnant animals were all that were 
* The state attained when the effects of the last meal have ceased to 
register in increased metabolic activity . 
61 
available but it  was hoped it would be possible to  pick up the effect 
of lambing and lactation on the oxygen consumption of these sheep . 
All of the ewes were in*lamb to the Southdown ram. 
'l' he first ewes were moved to Hassey f\gricul tural College on 5/? /56 . 
These sheep (five of each breed) wer� housed one to a pen (Plate 19 ) 
and were offered a diet of good quality hay � libitum and one pound 
per head of a concentrate in nut form. The Cheviots settled down i . e­
diately and ate hay and cake voraciously, but of the Romneys one starved 
herself to death while the other four came only slowly to eat their 
rations . Romney No . 5 did not eat cake until the t ' t i rd day of Block II 
energy metabolism experiments �1d this was possibly reflected in her 
records which were below the general run of all the other sheep to this 
point (Appendix A , tables 1 and 2 ) . 
The general management of these animals was quite simple , the aim 
being to interfere with them as little as possible . Fresh hay was put 
in the racks at 9 : 00 a.m . every morning and the cake ' as fed at the 
san1e time . Feeding of a concentrate sheep cake was soon abandoned. 
The sheep ate it well for three or four weeks an then grew tired of 
it  and left the major portion of their rati on . 
In place of the cake it was found that one pound of c ru s h ed  oats 
with a handful of kibbled pe as were eaten avidly. The latter were 
particularly welcome to the animals who would refuse to touch their 
oats without the sprinkling of peas . This diet of hay � libitum, 
crushed oats and kibbled peas was maintained throughout the investigation . 
Plc:.te 19 . Interior of sheep shed . 
62 
6.3 
At no time did the animals show lack of interest in it until the after-
math of a ten d� period of complete starvation . Water was alw�s 
available in each pen . No attempt was made to measure water consump-
tion but hay was consumed at the rate of approximately three to four 
pounds per head per day. 
Environmental variations within the shed due to the sun rising 
on one side and setting on the other , a leaky roof and wind-driven 
rain were compensated for by randomising the sheep within the shed. 
The ewes all lambed without assistance between 24/8/ and 13/9/56 
and both they and their lambs took quite kindly to thei:i.' living 
conditions . 
Foot rot presented virtually no problem during the entire period 
of confinement which in some instances extended to eleven months . 
Attention was paid to each foot individually as and when i t  was o 
served to be balled up with a mixture of hay and dung. 
From the ver.y beginning the writer set out to win the confidence 
of the animals in order that they might be completely rel , d when 
being handled for recordings ui th the oxygen spirometer . All figures • 
graphs and statistical analyses in this division of the project depend 
for their final accuracy on the degree to which success was achieved 
in this matter. The sheep soon came to accept the necessary handling 
as part of the routine of living . 
On 27/7/56 , one Cheviot {No . 10 ) and one Romney {No . 5 ) ere each 
u n 
fitted with a Field pattern tracheal cannula. E eh wound healed snugly 
64 
round the stem of the cannula, the stitches being removed about the 
fifth day after the oper tion . During the second post-operative week 
the technique of using a tracheal balloon and the method for connect­
ing the .sheep to the oxygen spirometer were tested . 
The first tests were successful , neither of the sheep seriously 
objected to being handled and ever,y indication was given that the 
technique was likely to prove worthy of dev, �.'J .... ment. The first two 
sheep (which survived until they were slaughtered on 8/5/57 )  were used 
for a period of approximately three weeks before any further sheep were 
cannulated. Technique development having proceeded to a point at which 
it could be decided to continue ¥it.h the project , four mol·� each of 
Cheviot and Romn ... y ewes were cannulated on J0/8/56 . 
�ecordings on the or3gen spiro�eter were made in a small room at 
one end of the sheep barn. Against one wall of this room small pens 
were constructed as illustrated in plate 12. These recording pens 
were of approximate dimen8ions 20 " x 4 2 "  and held one sheep comfortably. 
Later experience showed that they would have been improved by being 
sev 'ral inches narrower to prevent the sheep from turning their heads 
away from the operator. Each pen was closed by a sliding gate which 
could be operated smoothly with one hand while the other was busy with 
the oxygen spirometer. 
The sheep were quickly trained to >-Talk quietly out of their pens 
and to the recording room where they were backed into the small pens . 
So used to thiE procedure did the sheep become that they soon turned 
themselves round and backed into the recording pens with only slight 
guidance. This  cooperation was regarded as being of the utmost impor­
tance to the work . 
From 5 to 19/9/56, the ten sheep were subjected to a period of 
training to the oxygen spirometer. This lengthy training was aban­
doned for all future animals brought into the experiment as it  was 
found that four dav � at the most were needed before the animals were 
completely relaxed in the recording pens . They all regularly started 
chewing the cud immediately after the gate \·Tas dropped to fasten them 
in the recording pen. Sometimes they would continue cud chewing 
throughout the recording runs . In the case of these records i t  was 
found that a second run could be obtained free of the interruption of 
cud chewing if given immediately after the first . Although no clai 
is made that all aniffials behaved themselves at all times , it is sub­
mitted on the basis of the cud chewing phenomenon that recordings were 
obtained with the animals in the maximum attainable state of relaxation . 
As will b� seen from plate 12 , during the recording period the gate 
was in the raised positiryn . 
Plates 20 and 21 illustrate typical spirometer recordings , their 
straight line nature , repeatability , and eruct tion blips . 
The recording period was originally fixed to commence at 1 : 00 p.m . 
every d� but after the first four d�s of Block I experimental record­
ings ( Appendix A , table 1 )  the policy was altered to one of making all 
records before 9 : 00 a.m . , commencing as nearly as possible at d�break. 
Plate 20. 
s::: 
%'a-
$( 0 
:J) 
D .... � •M rl 
Ct-1 0 
(f) .� 1-
c Q) +' 
�· 
� 
1-:n t--Q) t-H +' 
·M I I J.ll 
.._._ 
Typical oxygen spirometer graph . Note the downward blips "X" produced during 
eructation. These blips are due to cessation of breathing and not to the 
entry of eructated gas into the spirometer . 
I TLIJ 
-· " I+ t - ! ' 
- 1---t 
- 1---t t-H -
H 
I} 
t--
L• 
� 
m I -1-l 
Hinu t e s  ;:.nd tenths of minut e s  
"' "' 
A 
B 
Plate 21 . Metabolism charts illustrating : -
n 
u: 
+ 
� 
t-
ttt+RsttBtttm:ttffifEH=t�mTI I I I I I I I I I I I I I I I I I \- .-
I -
.f..l 
·r:j mf i 1Hff'� - ---=fFITI I lfFR+T +It��ttl1 1 �fH � ; I ' 
. .  n s  of minutes 
l .  The lack of effect of prolonged met2bolism recordings on the respiratory pattern and rate 
of oxygen consumption . 
2 .  The repeatability of recordings ; chart B was made 8 minutes after chart A . 
3 .  Eructation blips " X " . 
0\ ---.) 
68 
At this time of d� temperature and barometric pressure were almost 
static and except during midsummer there was no accelerated bre:.1thing 
due to the effects of heat . 
B-V EX P ERIMENTAL DESIGN 
In choosing the experimental design for the energy metabolism 
experiments account had to be taken of the aim to study the animals 
on full feed at different peniods during the year and the effect on 
them of prolonged fasting . 
It was also necessary to recognise the possible existence of 
fluctuations in ruminant metabolism such as were indicated by the work 
of l itzman and Benedict ( 1938 c )  and Benedict (1938 c ) . Further , 
since the total time required to record 10 sheep was between two and 
three hours , a consistent bias could be introduced if each animal 
o ccupied the same position in the recording sequence from day to day. 
In addition , the sheep were moved to the recording pens in batches of 
three ( except the tenth which was brought in immediately after the 
seventh had been recorded ) ; hence there was a possibility that the 
recent muscular exercise might be an unbalancing factor if the record­
ing plan was not designed to even out any such influence . 
The above considerations suggested the use of a Latin square 
design in which the oxygen consumption of each sheep wqs recorded 
daily in the early morning , each of the sheep appearing once in each 
of the ten possible positions in the recording sequence over a ten day 
period, these ten day periods occuring over the year. 
B-V I CHRONOLOG ICAL EVE T S  19/9/56 - 8/8/57 
Table 2 presents in chronological order the events from the 
commencement of the first routine energy metabolism recordings on 
19/9/56 to the slaughter of the last sheep on 8/8/57 . lso in 
chronological order, a detailed discussion of the listed events 
follows below : -
19/9/56 
69 
Block I of routine energy metabolism recordings was commenced 
(Appendix A ,  table 1 ) . The ewes h d all lambed and the sheep were in 
the early stages of lactation . Three of the Cheviots were unavoidably 
five to six d�s closer to their parturition dates than the Romneys. 
This block of work went reasonably smoothly being complicated only by 
the main defects in the technique at this  stage--leaks between the 
cannula and the neck of the animal. These occurred in fourteen ( eight 
Rornneys and six Cheviots)  out of the total of one hundred recordings . 
To rule out the effect of separation of ewes from lambs , during 
the recording period the lambs were placed in the recording pens th 
their mothers . Having b en previously trained to this routine the 
lambs would in many cases voluntarily follow their respective dams into 
the recording room and then into the small pens. 
7/10/56 
Block 11 of routine energy metabolism recordings was commenced 
(Appendix A , table 2 ) . 
Table 2. Chronological events--energy m tab0llsm sheep-19/9/56 to 8/8/57. 
19/9 - 4/10/56 
7 - 17/10/56 
19/10/56 
2/ll/56 
2)/11 - 28/12/56 
5/1/57 
2)/1 - J/2/57 
28/1/57 
8/2 - 12/2/57 
14/J/57 
29/J - 7/4/57 
8/4 - 17/4/57 
18/4 - 27/4/57 
7/5/57 
8/5/57 
8/S - 8/8/57 
8/8/57 
Experiment (Block (I ) :  Appendix A;  table 1)  to eo:mpa th en rgy 
etabolism of R:omney Marsh and Cheviot ewes on f\111 rations shortly 
after lambing in the spring of the year. 
Experiment (Block (llh  Appendix A; table 2 )  to compare the energy 
metabolism of lactating Romney Marsh and Cheviot ewes fUll rations. 
Also to aso rtain the effect of 72 hours starvation on their n rgy 
metabolism. 
All ewes turned out to grass in an attempt to improv their condition. 
All ewes returned to sheep barn - lambs weaned. 
Experiment (Addendum a) to investi ate the ef.tect of sh arin and 
drenching with simulated rain on the respiratory excban e and 
r spiratory pattern of sheep. 
Experiment (A<idendwn b)  to ascertain the effect on their metabolic 
rates of feeding one pound oats to each o£ three h ep after twelve 
hours starvation. 
Experiment (Block (Ill );  Appendix A; table J )  to compare the nergy 
metabolism of Romney Marsh and Cheviot ewes on tnll rations in Jdd­
SWIIIler. 
Experiment (Addendum c )  to ascertain the effect ot 1 thyroxin on 
the oxygen consumption, respiratory rate and bOdy ight of on 
Romn y l�arsh ewe. 
Experiment (Block (Ilia ) ;  Appendix A;  table 4) to compare the energy 
metabolism of Ro:mney Marsh and Cheviot ewes on f'u.l.l rations in mid ... 
er. 
Experiment (Addendum d) to investigate thane exhalation in sheep. 
Experiment (Block (IV) ; Appendix A,  tab1e 5 )  to c par the energy 
met.aboli.s of Romney Marsh and Cheviot ewes on .f"Ull. rations in 
autumn.. 
Experiment (Block (V) ;  Appendix A ,  table 6) to ec:u:pare th en rgy 
etabolism of Romney Marsh and Cheviot ewes over a ten day p riod 
of total starvation in autumn. 
Experiment (Block (VI ) ;  Appendix A, table 7 )  to compare the energy 
metabolism of Romney 1-!arsh and Cheviot ewes with f'u.ll. rations 
offered after a ten day period of total starvation. 
All ewes photographed. Blood samples taken for haemato1ogical studies. 
Six ewes slaughtered. Carcass data and skin areas obtained.. 
Remaining two sheep used for tracheal cannula development {one died 
4/6/57) .  
Last ew slaughtered and th n cessary carcass and skin 
recorded. 
'J 
a data 
?l 
This consisted of seven daily records taken under normal conditions . 
The eighth and ninth d� records were obtained after twenty-four and 
forty-eight hours of complete starvation, respectively. Tenth day 
records (72 hours starvation ) were not taken due to a breakdown in 
the writer' s  health . Of Block II , seven records (two Romneys and five 
Cheviots ) were spoiled by leaks.  
19/l0/56 
The ewes were turned out to grass.  They had all recently lost 
condition . It was believed that their lambs were bearing heavily on 
them in the comparatively confined space of the pens.  
During the period at grass one Cheviot (No . 9 )  died of maggot 
fly strike in the trachea and one Romney (No . l )  died of suffocation 
following the loss of her cannula which sprung its fastening catch. 
One Cheviot (No . 10) was very sick due to the thickening of the mucous 
membranes below the lower tongue of her cannula. It was at this stage 
that a method for dealing with this type of situation was discovered, 
i . e .  by simply lengthening the cannula with a rubber tube . The Ch viot 
(No . 10 ) made a splendid recovery. 
2/11/56 
The ewes were returned to the sheep shed, the lambs being weaned 
at this time . 
72 
23/11 to 28/12/56 
The opportunity was taken at this stage of investigating the 
effect of shearing and drenching with simulated rain on the respira­
tory exchange and res1 iratory pattern of the animal&•  (Addendum a) 
5/1/57 
Three of the ewes , two Cheviots and one Romney, were us d to 
ascertain whether or not the technique could be used for plotting the 
effects on their metabolic rates resulting from feeding each sheep one 
pound of oats after twelve hours starvation (Addendum b ) . 
Unfortunately towards mid-January hot weather with shed tempera­
tures rising to almost 90or . brought great discomfort to the sheep 
which , showing the symptoms of a pasteurella type infection, began to 
respire very rapi�·. !t  was then found to be eseential to leave the 
corks  out of the tracheotomy tubes to compensate for the slight resis­
tance offered to rapid breathing by the presence of a tube within the 
trachea. Rapid passage of warm air in and out of the stem of a tra­
cheal cannula leads to a build-up of dried mucus on its lower tongue . 
Under these conditions it was necessary to remove the cannulae for 
scraping twice or thrice daily. An inordinate amount of time and atten­
tion was required at this stage . Only rapid breathing produced the 
effect for at all other times the cork could be left out with impunity. 
23/1/57 
Block III metabolism recordings commenced (Appendix A ,  table J } .  
7J 
At this time the early morning temperatures were moderate and in the 
ten day period only two recordings were spoiled by leaks.  
28/l/57 
One sheep (Romney No . 2 )  was allocated to a cooperative study of 
the effects of 1 thyroxine on its metabolic rate , respiration rate , 
and live weight ( Addendum c ) .  
8/2/57 
A further block of recordings , Block Ilia (Appendix A ,  table 4) , 
was brought to a close after the fifth d� because high temperatures 
and humidity were maintained throughout this d� and continued for 
some weeks after. One recording was spoiled by leaks. 
During the hot w ather period it was found of benefit to the 
animals to thoroughly spray down the shed about noon hour. 
More clement conditions arrived towards the end of March when 
night temperatures started to fall and mild hoar fro ts occurred. 
14/3/57 
In view of the statements made by Brody ( l945)f  Blaxter (l954)t 
and Ritzman and Benedict (19J8)•to the effect that the ruminant exhales 
methane in considerable quantities , it was deemed necessar,y to inves-
tigate the amount of methane , if any , which accumulated in the oxygen 
spirometer during recordings . This was done as soon as appropriate 
arrangements could be made for gas analyses to be carried out by the 
Dominion Physical Laboratory at Wellington (Addendum d) . 
• These references appear in the bibliography for Addendum d--Methane 
exhalation in sheep. 
74 
29/3/57 
Experiment Blocka IV, V, and VI followed a prel.im1nary two or 
three dqa re accustoming the animals to the routine. Block IV 
(Appendix A,  t ble 5) was nm oft w1 th the sheep under normal tee4 
conditions and Block V (Appendix A, ta'ble 6) followed with the 
animals completeq deprived ot tood although water was alloHd � 
libitum. For thct starvation period all bedding wae removed and 
onl¥ a hanc:l.tul of scattered sawdust was used to aoak up the ur�. 
The sheep made no attempt to eat the sawdust. 
From the manag nt viewpoint the ten d8i.Vs of complete starvation 
were ot great interest. At no time did the sheep tall to hunt tor 
food on the journey trom the recording room to their pens, or taU 
to shw 1nwrest when the teed room door was opened. The animals 
show'ed no signa ot wealme a and spent no undue length ot tS.. 
lying down. Water consumption, though not measured, fell quite 
appreciably. Respiratien r tea fell most markedq (Plate 22) . 
For Block VI (Appendix A,  table 7) the animals were again 
offered. tull rations . Init1�. they were in all cases � 
part1al.ly oon8Uitl8d. In tact, it took thre to tour daya before 
the sheep regaiMd their appetites. Two weeks af't.er the completion 
of Block VI it was decided to abandon t\lrther energy a..taboll. m ork 
owing to the onset of a tracheal infection in six aniroals. 
7/5/SZ 
T� experimental animale were photograpbecl. The ae  photogr phs 
� 
l ., 
I 
I 
.. s:: g>n � -X u 
U) (!) 
!1· .,..., ,..-j 
� 0 
U) .s:= 
1:: I Nil (!) 
+' .  . 
"d ' 
� 
U) (!) � 
,. 
+> 
j I • 1 r 
Nf; ..., .. +.e� <>nr t en t l: s  of minutes 
-t-
-. 
;..... 
- . 
-- :it' tt��  
I I . I :l� .. � ,1 -
- - _ . .J --1 " _; lW' :.:r l �-- -·· · - ·  .. : + 
�-!  ; -+ - +--
I -
-�. ' .  I J -+---- ' ' I I -t-t· I 
.. 
' -
7.� 
Plate 22. The effect of starvation on the respiratory pattern of a Cheviot ewe . 
' 
A 24 hour starvation 
point 
B 240 hour 
starvation 
point 
-.,J \.)'\ 
7'6 
are presented in plate 23 as evidence of the general well being of 
the sheep despite the tracheal infection and the fact that they had 
been subjected to many months of constant handling. 
An a�tempt was made to obtain estimates of blood volume. packed 
red cell volume (P .C.V . ) and haemoglobin values .  This work was 
incompleted through poor artificial lighting and the extraordinar,y 
elusiveness of the jugular in some animals . 
8/5/57 l 
Eight of the metabolism sheep were slaughtered. At slaughtering 
the animals were most carefully skinned and the following weights 
recorded :- dressed carcass ,  heart , adrenals , thyroids , and ovaries . 
The dressed carcasses were put into cold storage for future reference 
and the ovaries were examined. A permanent record of each skin 
area was obtained by laying the skins on sacking which was then 
carefully cut to shape . 
8/5 to 8/8/57 
The remaining two animals (one Romney and one Cheviot ) were 
used for cannula development purposes . The Romney died of pneumonia 
on 4/6/57 .  The skin, heart , adrenals , thyroid, and ovaries were 
removed immediately following death . So fresh was the carcass that 
it bled quite freely under the knife . This animal was in a ver,y fat 
condition . The lone Cheviot was used for further de·-relo::omental 
experiments until 8/8/57 when it was slaughtered and the necessary 
measurements recorded . 
Plate 2J . The energy metabolism experiment sheep 7/5/57 (prior 
to slaughter on 8/5/57 ) .  
y 5 
15 
10  16 
77 
X 20 
7 17 
Plate 23 . ( cont . ) The energy metabolism experiment sheep 
7/5/57 (prior to slaughter on 8/5/57 ) .  
78 
B-VII BASES FOR THE PRESENTATION OF 
THE RESt�TS OF ENERGY METABOLISM EXPERIMENTS 
79 
The bases for the presentation of the results of energy metabolism 
studies have apparently been argued over since such studies were 
initiated. So complicated have some authors made their Qomputations 
and theories that Benedict (1938 d)  felt obliged to hope that the 
progress o f  studies in animal metabolism would not be deterred by 
such complicated techniques and methods of calculation. 8enedict 
(1938 e )  and Brody (1945 e )  devote a large section of their published 
works to the possible relationships existing between heat production 
and the following biologic factors : -
( 1 }  Surface Area 
The method of expressing energy metabolism as a function of 
surface area was introduced by Rubner (188) ) . Based upon a series 
of observations on dogs of greatly differing weight• Rubner came t o  
the conclusion that the surface area calculation equalised the heat 
production of these animals .  This concept was subsequently applied 
by Rubner and notably Voit (Benedict 1938 f) to animal species in 
general and is today generally accepted as applicable to all warm 
blooded animals with few exceptions . Benedict and Brody, in general, 
criticise the use of this method of presentation as follows t -
Representing metabolism as a function of external surface area 
would imply that the surface area is the cause of the met bolic rate . 
whereas heat production is merely incidental to metabolism as the 
anllnal ' s body does not , within the zone of thernml neutrality. 
met�bolise because it must produc heat but produces heat because 
it metabolises ; the immediate causat ive mechanism being resid nt 
80 
in the neuroendocrine system and not ln the external surface area . 
In addition, the surface area of a living anim2.l is not constant and 
cannot be measured in such a manner that the results can be eh eked 
by different investigators . Further, an animal ' s  surface area , as 
it rel�tes to heat loss , changes with environmental temperatur 
not only by its skin contracting in cold weather and spreading out 
in hot weather but also by its developing heat conserving and 
heat dissip ting devises , . g .  wool in sheep . 
( 2 )  Body Weight 
Reporting energy metabolism upon the basis of heat production 
per unit of bot� weight is according to Benedict , a time honoured 
custom: however,  the same author argues that there is little basis 
for the belief that the expression of the heat production per 
kilogram of body weight can equalise either intraspecifically or 
interspecifically animals of varying sizes . 
(.3 )  Body weight'> o.z; 
Brody argues that basal energy metabolism does not vary directly 
with simple body weight or surface area but varies directly with 
what can be called metabolioa.lly- effective body weight. Brody found 
that this value was about W\0.7J for animals ranging in weight 
£rom,;mice to elephants . In 193.5 the Committee on Animal Nutrition 
of the National Research Council* ( U.S .A . )  had recommended that 
this figure w\0.7J be adopted for metabolism work . Benediet ex-
presses some dissatisfaction with it . 1n considering th fUtility 
8l 
of attempts to discover a unifYing principle in metabolism. Ben e di ct  
reaches a conclusion that : - uThe more one thinks over the factors 
known today as influencing basal metabolism, the more one realises 
that basal metabolism in many species is not a constant and that 
with many races there are differences in m tabolism. It seem 
therefore unjustifiable to apply mathematics to the pooled end 
result of the activities of millions of cells , each highly di!-
ferentiated with different energy potentialities and actuated by 
di£ferent stimuli . "  
However , Kleiber (1947 a )  in a review of body size and metabolic 
rate remarks that , "If this i s  the way Benedict feels , one cannot 
help but wonder how he ever became interested in conducting a 
respiration trial and why, furthermore , he even calculated the 
means o£ groups of several of these pooled end results , which indeed 
is applying mathematics t "  Kleiber {1947 b )  goes on to eventually 
put forward his own recommendation that the three-fourths power of 
the body weight is representative of metabolic body size for com-
parison purposes . 
• Report of co:r..ferenoe in energ-3 metabolisr4 held at State College , 
Pa. ,  under auspices of Committee on Animal Nutrition , Nat . Research 
Council , J\.ine 1935 � 7 { cited from Benedict , F.  O .  (19J8) Cam .  
Inst . Wash . Bul . 50J : 177) . 
82 
Both Benedict {19)8 g)  and Kleiber {1947 e )  support the possi­
bility of blood measurem nts being o f  value as an alternative t o  
car c a s s  :factors in expressing energy metabolism measurements . 
r1arston (1948 o) adopted the funetion ·w 0 •73 as a parameter to 
rel te the heat productions of sheep of different body weights 
and as a basis for comparing their behaviour with cattle . In 
eonsid.orin Harstons work, however , Hellberg (1949 a ) comments 
that "The correction for the very wide variation in weight ��s 
carried out in a manner that is probably not correct . It also 
s eems as though certain systema ti c errors are inherent in the 
ma terial . " ll.elJ_b'erg ( 1949 b )  put s- forward an argument that so far 
as possible the variation in body weight ought to be analysed with 
reference to its causes and that the different items ought to be 
treated differently in the way of correction . 
For this investigation the problems of reporting are to a 
certain extent sidestepped by the presentation of the oxygen con­
sumption data corrected to Standard Temperature and Pressure and 
the data as related to : -
( 1 )  Animal .; 
{ 2 ) Kilogra.lJls body weight • 
( J )  S q . metr�s computed surface area. •  
(4 )  Kilograms body weight 0 •73 
The presentation of the basio data in litres o2/5 minutes is 
a departure from the usual practice of expressing metabolism in 
* Brody 1945 f 
Cals ./5 mins . or Cals ./24 hours . Benedict ( 1938 h )  puts forward 
a powerful argument against the use of the 24 hour period in so 
much as it seems illogical to him to report results on the basis 
83 
of the 24 hour period when the measurements are almost n ver continued 
for 24 hours . Benedict states that the results could be expressed 
as cubic centimetres consumption of 02 per minuta . 
The writer considers that since his measurements were made as 
02 consumption per five minute periods , th�n the results should be 
r eporte on that ba is . For purposes of comparison the multipli­
cation of th figures by the appropriate factors will give Cals . 
pe� 5 minutes , 1 hour , 12 hours , or 24 hours . There is some risk 
in doing this , however, for the multiplication by K. results in 
increasing the rror variance by K2; also in starvation periods 
the caloric value of the o�gen consumed may change drastically. 
Simply expressing the metabolic rates in o�gen consumption 
per five minutes avoids all argument about caloric Values and time 
periods ; therefore , that is how the data are presented• although 
on one occasion the Galories per 24 hours presentation is used 
when it i s  desired to compare standards . 
3-VIII RES LTS 
(a )  Statistical Proc�dures . 
Missing values for Block I ,  Block �I and Block IIIa (Appendix 
A,  tablee 1,  2 ,  4)  were calculated by standard statistical pro­
cedures for randomised blocks (Cochran and Cox 1957 ) and the tables 
were analysed on a randomised block basis . Block III (Appendix 
84 
A. table 3 )  was completed by the standard procedure for two missing 
values in Latin squares (Snedeoor 1946) .  Block-s III , IV, V ,  VI 
(Appendix A ,  tables 3 ,  5 ,  6 ,  and 7)  were analys d as Latin squares .  
B-VIII (b) Int;rodugtion of New _tmimals 
The introdu ction of new animals during the period of this 
proj et makes it necessary to interpret all results ��th caution. 
However , it is a striking fact that whenever replace.rr.ent animals 
were introduced they r sponded in the same general manner a th 
remaining animals in eaoh group (Tables 3 and 4,  appendix A•  tables 
3 to 7 inclusive ) .  
For the changes b,y way of the introduction of new animals 
(four Romneys and three Cheviots ) it should be noted that no sheep 
were brought 1n during an energy metabolism experiment. • Of the 
ones that fell by the wayside , No . 1 died of suffocation following 
the loss of her can ula ·when out at pasture J No. 2 failed to regain 
flesh after rearing a ver,y good lamb and so was relegated to other 
work ; No . 3 died of pneumonia follol-1ing on probable grease contam­
ination of the lungs ; No . 9 died of blow fly strike in the trachea 
while out at pasture ; Nos . 6 and 4 died of pneumonia prob�bly 
connected with a pasteurella infection for which No . 2 was suspected 
of being the carrier ; No . 21--a very good and fit animal--died for 
no readily discernible reason . 
It is notewo rthy that the fi.rst two sheep to be c annulated 
Table ) .  Showing for the energy metabolism experiment ewes : 
R 1 
R 2 
R 3 
lt- 4 
R 5 
c 6 
c 1 
c 8 
c 9 
e 1 0  
R 1 5  
R 1. 6  
R 17 
c 20 
C 2l 
R X  
e x  
• 
•• 
•••  
+ 
# 
+. 
a . Mean livewei.ghts in ki.logr ( experiment Blocks I- VI) .  
b.  Square metres computed surface area'! ( experim nt Blocks I - IV).  
c .  Final 11. veweights in kilograms. 
d.  Square metr s measured surface areas. 
19/ 9 - 4/10/.56 7 - 17/10/56 23/1 - 3 / 2 /57 8 - 12/2/57 29/3 - 7/4/57 8 - 27/4/57 
Block I Block n Block III Block IIIa Block IV Bl.ock 
V VI 
b* * a b* * a b* *  a b* *  a b* * *  a &' c d 
53 · 5  1 . 61  .54 . 0  1 . 17 
42 . 6  1 . 02 40 . 4  0 . 99 
44 . 0  1 . 04  )6 . 7  0 . 93 
49 . 0  1 . 1 0  46 . 3  1 . 07 49 . 0  1 . 10 5 0 . 6  1 . 12 
40 . 8  0 . 99 39 . 5  0 . 91 ,54 . 1  1 . 1.7 51 . 5  1 . 13 50 . 8  1 . 12  4) . 5  44 . 4  5l.r 1 . 1) 
4) • .5 1 . 03 3 9 . 0  0 . 91 
46 . 3  1 . 07 42 . 6  1 . 02 41 . 1  1 . 00 42 . 9 1 . 02 ) 6 . 7  Ch98 )4 . 0  )4 .. 0 31 . -r 0 . 91 
48 . 1  1 . 09 46 . 7  1 . 07 44 . 1  1 . 04 45 . 6  1 . 06 45 . )  1 . 04  39 .. 5 40 . 8  40.4+ l.o6 
so.) 1 . 12 47 .2 1 . oa 
4) • .5 1 . 0) . . 3 9 . 4  0 . 97 4.5 . 0  1 . 05 47 . 2  1 . 08 49 . 4  1 . 09 44 . 0  45 . )  44 . 4  ... 1 . 02 
,54 . 6  1 . 17 53 · 5  1 . 16 5 7 . 6  1 . 18 49 . 9  so.a 51 . r+  1.22 
47 . 7  1 . 09 46 . 0  1 . 06 .51 . 0  1 . 10 4,5 . )  47 . 2  44 + . o 1 .1 6  
4 5  •. 5 1 . 06 46 . 0  J.o6 47 . 6  1 .07 41 . 3  42 . 2  40.8+ 1 . 02 
42 . )  1 . 01 44 . 0  1 . 04  44.4 l.o4 41 • .3 42 . 2  41 . 7"" l.f)� 
45 . 1  1 . 09 5 0 . 1  1 . 12 
sa.o  1 . 21 54 . 6  5 4 . 4  ++ 1 . 17 
6o . s  1 . 24 .5 4 . 4  5 6 . 2  56 . 6+• 1 . 20 
Brody ( 1945 f) 
Computed from the weights of the animals in the particular period 
Computed from the weights of the animals o·trer the period 15/1/57 - 7/4/57 excepting for no. ' s  X and. Y 
whose surface areas were eOliif)uted from their weights over t.he period 4/2/57 - 7/4/57 . 
Slaughtered 8/5/57 
Died 4/6/57 -
Slaughtered 8/8/57 
Table 4. Showing (1 ) The breed means and St. D1 s per experiment for oxygen consumption* in litres per .five 
minutes . 
(2 ) The F. values derived from. between breed anaJ.ysee of variance (Appendi.x A t  tabl 
8, 9, 10, 11) carried out on the oxygen consumption data related to the following 
bases . 
(a)  Animal (b) Kg. body weight (e )  Sq. metres (d) • body w ight to th �r 
su:cl'aoe area o.n 
{ co!ilp�ted ) 
(3 ) Signi.f'icanc ot dif.f'erenee betw en breeds . 
(4) Mean body weight for periods in kilograms . ,  
(a ) (b) (c )  (d) 
Dates Mean OXYgen Consumption in litres Animal Kg. body Sq. metres Kg. body 
per five minute weight sur.fac w igbt 
area 
( corn uted) 
O.'n 
Cheviot Romney F.  F. F. F. 
Mean St. D Mean St. .  D 
19/9-
4/10/56 
Block I 1.57 0.16 1.79 0 .29 2 .70 2.50 3.71++ 3.18* 
Breed Di£.f'erence 0 .22 
7 - Block II 1.45 0 .20 1.44 0 .25 0.02 o .oo 0 .04 0 .02 
17/10/56 
Breed Difference 
ZJ/1- Block III 1 .40 0.29 1 .53 
J/2/57 
0 .21 1 .18 0 .32 0. 02 o.oo 
Breed Differen�e o .13 
8 - Block Ilia 
12/2/57 
1.37 o.rel 1.56 0 .22 2.)4 o.s3 1 .o6 0 .90 
reed Difference 0.19 
29/3-
7/4/57 
Block IV 1.18 0 .12 1 .39 o.u 10.50** o.os 4.68++ 1.81 
Breed Di.f'ference 0.21. 
8 - Block V 0 .86 0 .14 1 .05 0 .13 19.12+ 2.42 18.61+ 8 • .53** 
17/4/57 
I · 
Breed Difference 0 .19 
18 - Block VI 1.06 0 .20 1.18 
27/4/57 
0•19 3 • .57* o.oo le45 0.37 
Breed Dif.ferenc o .12 
• Appendix A ;  t.ables 1-7 give the data from which these figures were calculated 
++ Approaches significance 
•• Significant at 5� level 
+ Significant at 1� level 
Mean body weights 
for perlods in Kgs. 
(Group · ) 
Romney Cheviot 
46. 0  46.3 
' 43.4 4).0 
50.2  44.1 
49.5 46.o 
5) •. 0 47.o3 
(Nos . 5 and 10 ) survived in good condition until the end of the 
project--a period of about ten months . Of the original animals 
87 
in Block I (Appendix A , table 1 ) ,  Nos . 5 ,  7 , 8 ,  and 10 made the whole 
journey. 
B-VIII ( c )  Bo<iv Weights and Surf'ac$ Areas 
Table 3 shows that the body weights of all animals introduced 
into the experiments were maintained approximately in equllibrium. 
The main rea on for th J drop in ewe weights betw en Block I and 
Block II was probabl the rearL�g of the lambs wh i ah  ·chems lves did 
very well gaining at an ratas of 0 .34 lb s . and 0.46 lbs . per day 
for the Romneys and Cheviots , respeotiv ly (Table 5 ) .  
Surface area s for Blocks I ,  n ,  I I I , and IIIa wer computed 
from the weights of the sheep in each particular period (Table J ) .  
Surface areas for Block IV were cal culated from the mean weights 
of the anlinals ov r the period 15/1-7/4/57 exoep ing for Nos . X 
and Y whose areas �ere calculated on their weights fr�m 4/2-7/4/57 • 
There is a good level of agreement between the surfac areas computed 
at various times fer indi vi .ual sheep . The computed and the measured 
surface area s for each sheep also agreed closely. OWing to the 
difficulties involved in obtaining a true measurement of actual 
surface area (3enedict 1938 i )  the actual measuremen s were us ed  
only as a check on t h o  computeci figures . A stan ' ard surfac area 
was adopted for a eh £ue in Blocks IV ,  V and VI , normal , starvation 
and rehab i:i. i tat. ion periods resp cti v ly. The alidi ty of thi 
88 
Table 5 .  Showing {1 ) The lamb production data of ewes 1-10 1n the 
energy metabolism experiments. 
( 2 )  The analysis of variance between breeds 
for lamb rate of gain. 
Ewe Parturition No. of Birth Weaning Lamb 
No . Date Lambs Weight Weight Liveweight gain 
lbs . lbs . in lbs . /day 
1 6/9/56 1 7 20 0.29 )  
R ) 
0 2 24/8/56 1 8 29 0.35 ) 
M ) 
N 3 7/9/56 1 8f 23 0.33 ) mean 
E ) 0.34 
y 4 6/9/56 1 10 39 0 • .39) 
s ) 
5 30/8/56 l 7 27 0 • .35) 
6 12/9/56 twins 17! 34 0.53/2 0.26 )  
c ) 
H 7 27/8/56 1 8 33 0.41 ) 
E ) 
V 8 13/9/56 1 st 28 0.44 ) mean 
I ) 0.46 
0 9 28/8/56 1 8 26 0.33 ) 
T ) 
s 10 1.3/9/56 twins 15 36 o.57/2 o.2s ) 
Apalysis of varianqe between breeds for lamb rate of gain 
D.F. Variance F. Nec.F 
5'1> 1'1> 
Breed 1 .0325 6,25 5.32 11.30 
Error 8 .0052 
procedure may be questioned ; however, it is relt that no useful 
purpose would be served by introducing more mathematical analyses 
until some investigation has been made of the effect of improving 
condition and st•n·vetion on surface areas . 
B-VIII (d )  Position in Recording Sequence 
89 
Analysis of Blocks IVand V (Appendix A , table 10 ) showed that 
position in the recording sequence was exerting no significant effect. 
These analyses confirmed indications g iven by other blocks , whose 
analysis was complicated by missing values . 
B-VIII (e )  Lability 
Some concern was originally felt about the fluctuations in the 
metabolism of each animal (Appendix A , tables 1-7 ) ,  particularly 
when so far as the operator couJ.d a s certain there was nothing 
technically wrong with the apparatus . riowever , Ritzman and 
Benedict ( 1938 d)  remark that variations of' up to 30 percent or 
more were commonly met with in cows and that extraordinar.r variations 
in the basal metabolism may o c cur in the same individual and within 
a relatively short period of time . The stan 'ard deviations of the 
recordings of oxygen consumption in this project were as follows 
(Table 6) when expressed as a percentage of their particular m an. 
The figures in 11 cases out of 14 are within or approximate 
to the ;,t 1.5 percent experi..1ental errdr for metabolism recordings 
f, Ut f'orward by Wiggers (1949 a )  and are well below Ritzman and 
90 
Benedict ' s  :30 percent figure. Furthermore the analyses for day 
effects (Appendix A•  table 10) revealoo no statistica.ll.y s1gn1t1cant 
differences bet een days for Bl o c k s  II a nd  III t�hile th�t� significant 
day effect in Block l may be attributable to t h e  effect of tailing 
lactation in Ina to rising t .peratures. 1n Block IV to a season effect 
and in Blocl s V and VI to t he d eliberate s t arv ;. t i on a nd reb&bUitatiO'n 
of the animals. 
Tabl.e 6.  The standard deviations of the breed aeana for 02 consumption• 
within blocks expressed as percentages of the means (to 
nearest% ) . 
Bloek Cheviots Ramrteys 
I l.O .. O 16.0  
II 14 .0 17 .0 
m ( 21.0)  Hot weat her (14 . 0� Hot weather 
m a (2J.O) period (14 . 0 ' periOd 
IV 10. 0  8 . 0  
V 16.0 12.0 
VI 19. 0  16.0 
• Corrected to Standard Temperature and Pressure 
B-vm (t) Breed c� 
Table 4 includes the ean oxygen consumption figures per five 
minutee for each breed in tbe different experimental blocks . The 
--ou;o._�table shows th s1gnif1canc of the difforenoe s betw n these 
1Mintif·, a · derived :from �� s of Vill'1anca (Ap}mldix A. table• 
· -u 1nelua1Y � ) carried out on the �en oo.nsumpttoo r c ord s  on the 
previously discussed bases of presentation, i.e • •  animalJ Kg. body 
wt . • J computed surface area 1 Kg. body wt .  0.1). 
91 
It will be seen from table 4 that on a per animal basis except 
for Block II there apparently were breed differences between the 
mean figures for oxygen consumption, tbe Cheviots having an oxygen 
consumption rate ranging approximately from 10 to 15 percent below 
that of the Romneys . 
The absence of an apparent breed difference in Block II may 
have been due -to the fact that the Cheviots were in some cases quite 
close to their lambing dates and the Cheviot group were, as a whole , 
rearing their lambs statistically significantly better than the 
Romneys (Table 5 ) . High milk production probably necessitates a 
high metabolism (Ri tzman and Benediot 19.38 e ) .  Block II consisted 
of only nine days on the last two or which the sheep were starved. 
Tenth day (72 hours starvation ) records were not obtained due to 
a breakdown in the writer' s  health . 
However, statistical analysis (Table 4 and appencU.x A,  table 
8 )  showed that except for Blocks IV and V the differences betw en 
the br ed means for oxygen consumption p r animal were not significant 
although the F. values obtained are quite interesting. They indicate 
that the odds against the differences being significant rose for 
Block II and then fell steadily to Block IV where the difference 
was significant at almost the one percent leve�. Block V was a 
t n day starvation period and at this time the differences were 
very highly significant. For Block VI the gap clo ed and the odds 
92 
against the differences being signifi�ant rose quite considerRbly. 
Relating the oxygen consumption records to Kg. body weight , 
computed surface area , and Kg . body weight 0 .73 (Table 4 and appendix 
A , tables 9 .  10 , 11 ) gives a rather different picture , for the 
Romneys in the experiment Blocks III ,  IIIa, IV ,  V ,  and VI weighed 
more ths� the Cheviots (Table 3 )  and thus in general the higher 
oxygen consumption figures of the Romneys were treated with larger 
divisors than the Chevi ot s . Thls tended to equalise the breeds 
and it is recognised that the breed difference in oxygen cons��ption 
may be due to the greater Height of the Ro.mneys . However , it is 
noteworthy that the Cheviots hao the lower ox,ygen consurr�tion, 
firstly, in B l o c k  I when the Romneys paired with Cheviots of almost 
�qual weights , &nd secondly, 1n five cases out of eight 1n the other 
exper�ent blocks when individual Cheviots paired ��tb Romneys of 
equal weights . In tw� cases out of the remaining three the Romney 
(No.  5 )  was not eating cnncAntrf tss �nd this was possibly refl.ected 
i n  h e r  metaboli�m records (Appendix A , tables 1 ,  2 ) .  
On a per kilogram basis (Table 4 and appendix A ,  table 9 )  
non o f  the differences in oxygen consumption between the breeds 
ere statistically significant . Calculations b a s ec  on computed 
surface are s (Table 4 and appendix A ,  table 10 ) raised the F• value 
for Block I while lock IV approached significance at the five 
percent level and Block V was highly significant .  '!'he difference 
between th� breed me� s for Block V were again significant when 
.I 
9.3 
tr..e oxygen consumption was rel•�ted to the body weight in kilograms 0 • 7.3 
(Table 4 and appendix A ,  table ll ) .  
It i s  not a main object of this project to discuss or criticise 
the various metrods of reporting energy metabolism studies , but in 
view of the above results it is impossible not to recall the 
w-ords of Ritzman and Benedict (19.38 f) : -
"The general picture of such a complex combination of biological 
forces indicates that the basic need for heat production is not 
dictated primarily by the size of the body, its surface area 
and consequent heat loss , but by other factors that are essential 
to actuate the inherited function to wtich the organism is 
adapted . " 
In this respect tables 3 and 4 are most enlightening . From 
experiment Blocks II to IV inclusive the trend is for the mean 
weights of the groups of animals to be rising (Table � )  and so the 
oxygen consumption should also have shown a tendency to rise in the 
raw data as greater total body weight is supposedly associated with 
higher total oxygen requirements . What is  quite obvious , however, 
is  that the oxygen consumption was , in fact , falling (Table 4) . 
This fall in metaboli�1 to rev ch a low point towards the end 
of March is in complete agreement with Brody (1945 g )  who found in 
goats that the minimum metabolism occurred in the autumn or breeding 
season. OVulation in the human is notable for its effect in increasing 
metabolic activity (Wiggers 1949 b )  so that the effect of season 
would appear to counteract this possibility in the sheep . However, 
Brody, in showing that the metabolic peak in goats occurs in early 
spring, states that the weight gains in gro��ng goats are also 
maximum in early spring . Although data could not be obtained, 
th writer would venture that under normal spring conditions , 
at .}.east in the Cheviot hills , lactating ewes do not put on weight 
but rather produce milk "off their backs" .  
Ritzman and Benedict (1938 g )  concluded that some factors 
connected with season exert a dominant role in stimulating the 
metaboli&� of the tissues . In experiments on steers f d to maintain 
body weight they found a low metabolic level occurred in all cases 
between the end of January and the middle of March. In the period 
from May to November a high level of metaboliam occurred in all 
cases during late May, Ju..'le , or July. This was followed in all 
eases by a general declina in September with a relatively precipitat 
drop in October (March in New Zealand corresponds to Octob r in 
the Northern Hemisphere . ) .  Comparison of their cow data with their 
steer d . . t.a suggested a striking parallelism in the reaction of the 
metabolism to change in sttason. Pe�·iods of high and periods of 
low stimulus not attributa.ble to the ingestion of food w-ere definitely 
indicated. These authors considered these seasonal variations to be 
so pronounced that special experimen.ts wer designed to inv st1gat 
them using six cows . At that time no other exp riments on the effects 
of season on the energy metabolism of ruminants existed in the lit­
erature. Ritzman and Benedict. (1938 h )  summaris their work with the 
statement , "This influene of season presents a perplexing paradox 
to the theory that heat production is governed by heat loss ,  for th 
beat production of their cows was in all cases greater in summer 
when they least needed. 1 t.  • 
95 
Br� (1945 h)  in commenting on seasonal metabolic rhythms 
writes to the eftect that th,yroid actiVity increases in spring , 
reaches a l2l8JtiJinUl in llid..SWJUiter , and th•reafter decline a to a 
minimum in late winter. He believes that this variation is possibly 
related to seaeonal tood supply. Coop (195))  refers to tbpoid 
activity as a possibly intluence on wel production. The energ 
met.abolia of the sheep in this project would appear to have fol­
lowed a si.m1lar rhythm. 
So tar as the present vr1 ter is aware after Ri tzman and 
Benedict·• s work (19J:S g) no further energy metabolls experillente 
deliberately designed to explore the effects ot the seasons have 
been reported for large farm animals .  It does seem to  the writer , 
however ,  that the autumn fall in energy metaboli mq pr -relate 
the animal to food shortages in the cOJiing hunger season. This 
tra1 t is doubl¥ interesting when 1 t is considered tro the standpoint 
that the sheep in this projec.t and R1 tsman and. Bened.ict • a cattle. 
were being adequately ted. without a hint of a time of ahortage to 
cae. It should also be COJ.lsidered from the standpoint that in the 
aut\11111 the sheep is lqing up re•erve tor the winter and a high 
ae�abolic rate wou� perhaps defeat the �ction of th• &niaal which 
is , at that tiae , food storage. Remembering also the words ot JCqes 
96 
n .£!!.. (1950) ,  that to the starving individual the reduced metabolic 
rate means that his rate of loss of strength and endurance is 
d1minished and that , to carr,y it to the limit, he will survive longer, 
it is ,.,orthy of note that the differential in oxygen consumption 
( per animal)between the Romney Marsh and Cheviot breeds (Table 4 )  
had widened quite considerably between J/2/ and ?/4/S?.  The 
latter date corresponds to autumn in the Northern Hemisphere . 
Under natural conditions the Cheviot is the breed most likely to 
be exposed to food shortages , and this may be what is refiected 
in the comparatively low metabolism of the breed at this time . 
Emphasis is given to this viewpoint by the reactions of the 
breeds to complete starvation over a considerable period (Table 4, 
Block V and appendix A, table 6) . Under these conditions the 
difference in energy metabolism between the breeds was reduced 
below the point of statistical significance only wh en  expressed 
as oxygen consumption per 5 minutes per Kg . body weight. 
It is not intended to digress far from the main subject ,  but 
as this period of complete starvation is (so far as the writer is 
aware ) the longest reported in which metabolism records have be en  
taken, the opportunity cannot be missed for drawing attention to 
two points (Appendix A, table 6 ) .  
( 1 )  That a point of equUibri appeared to have been reached 
in the Cheviots at the 96th hour of complete starvation 
which corresponds v ry well with the figures given by other 
workers for the post absorptive stage (Forbes � �. 1926, 
97 
Ritzman and Benedi-..t 1938 b:  Marston 1948 b ) .  
( 2 )  The Romneys on the other hand show no obvious metabolism 
plateau, the trend being downward but in a ra"t,her erratic 
fashion. In the rehabilitation period (Appendix A , table 
7)  the Romneys ' metabolism rose faster than that of the 
Cheviots . 
For comparison purposes table 7 lists a number of references 
to the metabolism in Cals . /24 hours for sheep together with those 
derived from this investigation. 
It will be seen that the caloric values calculated from the 
oxygen consumption records obtained in this investigation are in 
general higher than those of the other authors . As has been mentioned, 
the error in calculating a 24 hour figur from a five minute recording 
may be considerable . Also, the animals were standing; and except 
for one experiment, Block V,  they were on fUll feed. Nevertheless , 
the fact of large seasonal and breed variations in energy metabolism 
may indicate a need for a revision of many of the figures put forward 
for the nutritional requirements of sheep. 
B-VIII ( g )  Carcass Data 
Despite the ten day period of starvation to which they had 
recently been subjected, of the eight sheep slaughtered on 8/5/57 
�1 but No. 7 showecl ovulation activity (Table 8) • and their packed 
reel cell volumes did not suggest anaemia (Table 9 ) .  
Carcass analysis (Table 8 )  carried out on the slaughtered 
98 
Table 7 .  Showing -- figures in the 11 terature and those calculated 
from the data in this project for the Calories/24 hours 
heat production of sheep . 
Author Cal.s/ 24 hours Metabolism classification 
Benediet (1938 a)  1200 Basal ( 24 hour fal!t, 45 
Kgs . sheep) 
Blaxter (1954) 1.557 Adult sheep on 862 gms . 
feed per day 
Brody (1945 1)  1580 Resting at 30 months 
" " 1760 Resting at 24 months 
Lines & Pierce (1931) 1270 Standard (48 hour fast ) 
Wood (1927) 1340 Maintenance requirement 
This Project 
Cals ./24 hours* 
Block Cheviots Romneys l1etabolism classification 
Mean St. D.  Mean St . D.  
I 2182 222 2487 403 Group Av rage (on full 
rations ) 
II 2015 278 2001 347 " .. 
III 194.5 403 2126 292 u " 
Ilia 1904 431 2168 306 " " 
IV 1640 16? 19Jl l.SJ • .. 
V 1195 194 14.59 181 Ten day tarvation period 
VI 1473 278 1640 264 Full r tions 
• Using 4.82.5 Cals . per litre as cal.orific value of oxygen 
!: 3:  
m >  
;;v �  
l> fTI  � -<  
"'(! )>  l> :.i) 
r· :.o 
: :') !.:'. C: .... v ,-l : -4 -· c � :;o  - ,...,.., Z r·-0 ()  
� 0  
:::e r 
• r Z fTI  • G)  !'l iT!  
Table 8. Carcass and organ data , energy metabolism experiment ewe s . 
t Significance 
r-.. r-.. r-.. r-.. r-.. r-.. r-.. r-.. r-.. r-.. of di£ference V"\ � V"\ V"\ V"\ V"\ V"\ 1.1"\ V"\ V"\ -..... -..... -..... -..... -..... -..... -..... -..... -........ between the V"\ "' V"\ V"\ '-0 V"\ V"\ V"\ V"\ � breeds -..... ........ -... -..... -..... -..... -..... -..... -..... CX) CO a) CO .::t CO CO a:> a:> 
Romneys Cheviots 
17 15 5 16 X 7 10 8 20 y 
Ovaries gms • 2.7  2 .8  3 • 0 2.5  2.7  1 .2  3 .2  2 .9  2.8  3 .2 
Adrenals " 4.9 6.0 5 .3  * 6.2  5 .0  6.4 6.9 6.4 7 .5 
Thyroids " 5.4 3 .7  5 .2  6 .7  7 .8  3 .8 6 .9  4.1 6.1 9.5 
Heart " 231 221 194 234 273 248 266 208 235 265 
Live wt .  lbse· 90 114 105 97 + 70 98 89 92 125 App. 5� level 
Dressed wt . - lbs.**  45 56 48 49 + 36 49 45 50 63 
Dressing �++ 46. 0  45.4 42 . 0  46.4 + 46.o 46.1 46. 0  46 . 0  45 .5 A pp. 5'f, level 
Carcass lean good fat fat + thin good lean v.good good 
Classification 
Heart/body wt .  2.57 1 .·:1ll· l .-85 2 . �1 + 3 ·54 2e 'll 2.)4 2.55 2 .12 AFP• 5� level 
Ovaries lsl  2 : 0  0 :1 1 : 0 0 : 0  0 :1 0 : 2  2 : 0  
( Corpus lutea ) 
'-0 '-0 
• Accidentally unweighed ** Hot carcass weight plus ++ Hot carcass weight minus head and cannons + Dead head and cannons liveweight 
Table 9.  Haematological data of  energy metabolism experiment ewes 7/5/57. 
Romneys 
17 15 5 16 X 7 8 
Blood vol. 
(litres per animal) 2 .03 2.82 2.10 2.39 2.13 1.87 
Plasma vol. 
(litres per animal)  1.52 1.96 1.60 1�79 1.56 1 .33 
Red cell vol. 
(litres per animal) 0.51 0.86 o .so o. 6o 0 .57 0.54 
Packed red cell volumes 
(mJ.,/100 mL blood) 39 26 32 25 26 28 30 
-- -
Cheviots 
10 20 
27 -
y 
41 
f-1 0 0 
animals and Nos . X and Y, which died and were killed respectively 
on subsequent dates , indicated a heavier heart in the Cheviot . 
Analys�s of the carcass data for the eight animals slaughtered on 
8/5/57 did not confirm this difference to be statistically sig­
nificant , but when the heart weight was related to body weight 
the difference approached the five percent level of significance. 
101 
Of the other weights recorded , only total body weight and dressing 
percentage indicated a breed difference .  the Cheviot dressing out 
two percent (up to five percent if the head and cannons arc included ) 
better than the heavier Romney. 
B-IX ADDEt'DUM 
The development o£ Divisior1 "B" o£ the overall 
project led to several incidental studies which have 
no part in the main b� o£ the results but being 
derived directly £rom the work are there£ore pre­
sented as an Addendum to it. 
102 
CONTENTS 
{a)  The effect of shearing and drenching with simulated 
rain on the o�gen consumption and respiratory pattern 
of sheep . 
{b)  The effect on their metabolic rate of feeding l lb. 
of oats to each of three sheep after twelve hours 
starvation. 
10) 
Page 
104 
{ c )  The effect of 1 thyroxine on the o�gen consumption, 115 
respiration rate , and body weight of one Romney Harsh 
ewe . 
{d) Methane exhalation in sheep. 119 
(e )  The regurgitation process in sheep . 121 
{ a )  The effect of shearing and dfenohing with simulated rain on 
the oxrgen consumption and respiratory pattern of sheep. 
Introduction 
At the beginning of November 1956 some of the traoheotomised 
sheep began to oast their wool. It was therefore decided to dis-
1� 
continue , for the time being , regular metabolism studies , as it was 
unknown at that juncture whether or not variations 1n fleece cover 
affected oxygen consumption. 
Ritzman and Benedict (1938) report the effect of temperature 
changes on the energy metabolism of sheared sheep but not the 
effect of shearing itself which is surprising as they put forward 
an argument that sheep offer a striking illustration of the ffeot 
of covering on their adaptability to changes in environmental 
temperature . The opportunity was therefore taken to study the 
effect of shearing on the energy metabolism of as many animals as 
was possible . 
Procedure and Results 
Starting 23/11/56 normal oxygen spirometer recordings were 
taken using eleven sheep for five d� follo�� several days of 
preliminary recordings . On the fifth day even of the sheep w re 
shear d following arly morning oxygen spirometer recordings . Of 
the remaining sh ep thr e Cheviot had east their wool prior to the 
co encement of this work and the fourth , a Romney. was left un• 
sh ar d to act s a control . 
The metabolism charts (Plate 24) revealed an abrupt change 
in the respirator.y pattern of the sheep which were sheared. In 
105 
all animals the respiration rate fell by approximately two thirds 
and the depth o£ inspiration was almost doubled. The seven sheared 
sheep showed slight increases in oxygen consumption after shearing 
(Table 10) .  
On lJ/12/56 the sheep were utilised for a further aspect of 
climatologieal work . A routine oxygen consumption record was taken 
from each animal. Each animal in turn was then subjected to soaking 
with simulated rain for three minutes . This period resulted in the 
collection of approximately one inch of water on the floor of each 
small pen. The sheep were subsequent� reconnected to the oxygen 
spirometer and reeordin s taken (Table 10 ) .. It was found that 
soaking with water had a profound effect on the respirator.y pattern 
and in every case (except for one Romney which still carried a 
full fleece) the oxygen consumption rose J plate 25 illustrates the 
respiratory pattern of a shorn sheep before and after wetting and 
plate 26 th respiratory pattern of the control also before and 
after wetting. 
One week later, 19/12/56 , the sheep were recorded twice , with 
an eight minute gap per sh ep b tween recordings . This eight minute 
gap was estimated as being equivalent to th period bet we n recording 
before and aft r oaking. No change occurred in any of the records 
(Table 10, plate 21) which continued early exploratory work ( Plate 21) 
'f 
(I) Q) J... .+) ..-f r-i 
<H 0 
] 
� ....., 
""0 
§ 
(I) 
f .+) ·rl H 
Plate 24. The respiratory pattern of a sheep before and after shearing. 
Minutes and tenths of minutes 
p 0 0'-
Table 10 . Showing the oxygen consumption records (litres 02/5  minutes ) * of eleven sheep ; 
( 1 )  Pre and post shearing , 
( 2 ) Pre and post wetting after shearing , 
( 3 )  Routine recordings with an eight minute time interval , 
(4)  One final routine recording . 
Cheviots Romneys 
Date 8 10 20 6 7 21 16 5 4 2 15 
23/11/56 Normal 1 . 13 1.41 1 . 32 1 .33 1 .41 1.60  1 . 33 1 .55 1 . 14 1 . 32 
26/11/56 fl, 1 . 52 1 .69 1.23 1 . 71 1.43 1 .43 1 .43 1 . 71 1 .14 1 .43 
27/11/56 11 1 . 32 1 .52 1.43 1 . 81 1 . 32 1 . 13 1 .60 1 . 14 1.41 1 . 13 1 .41 
28/11/56 11 1 .23 1 .52 1.22 1 .41 1 . 51 1 . 23 1 . 13 1 . 22 1.41 1 . 13 1.42 
29/11/56 Pre shearing 1 . 04 1 .43 1 . 23 1 .32 1 . 32 1 .41 1 .42 1 .33 1 . 13 1 .51 
sheared no wool no wool no wool sheared sheared sheared sheared control sheared sheared 
29/11/56 Post shearing 1 . 31 1 .40 1 . 73 1 .49 1 .59 1 . 12 1 .49 
30/11/56 1 . 38 1 . 34 1 . 29 1 .43 1 . 38 1 .58 1 . 52 1 . 33 1 . 53 1 . 34 
1/12/56 1 .43 1.43 1 .42 1 .42 1 . 57 1 . 37 1 .32 1 . 57 1 .43 
2/12/56 1 .43 1 .52 1 . 66 1.42 1 . 72 1 .62 1 .57 1 . 38 1 .42 1 . 57 
13/12/56 Pre wetting 1 .48 1 .50 1.41 1 . 54 1 .59 1 . 59 1 . 31 1.45 1 .13 1 . 50 
13/12/56 Post wetting 1 .57 1.69 1.50 1 .87 1 .87 1.97 1 .97 1.45 2 .44 1.68 
19/12/56 Normal 1 .28 1 . 38 1.49 1 . 28 1 .65 1 . 58 1 . 28 1.47 1 . 37 1 .63 
Eight minutes interval 
1.28 1 . 38 1.49 1 .28 1 .65 1 . 58 1 . 28 1.47 1 . 37 1.63 
28/12/56 Normal 1 . 39 1 . 39 1 .30 1. 21 1 . 30 1 . 39 1 . 30 0 . 93 1 . 11 1 .53 
* Corrected to Standard Temperature and Pressure 
Before 
wetting 
1-+...j- H--1--+--t-t-t - + +-t --t--1 --t-i 
-- - -· •-! 
-1-++-+ -+-++- 1-H-H 
�++44-� 14-rr-I�T r n�t-HHH-r 
flllturM::ttr!::tt f"'" 
� 11111 1 1 1 1 1 1 1 1  m 1 1 1 1 1 1  i 1 1 1 1 1 1 mtil,.ll l l l lll l l l l l l l  r 
Ul T·l� • r• · fT t , 1 ..c l llll l  lfl I I . - T +' t- .... , . 1--. - -� • 1111111 1. , ' ' T \1) +' 
ll� IIIU -
� 11 
1 1 ""H - +r. · - I  �-
:;5 ., 'J . -, - �-
§ -� ; After .b t -1 4 -t- · .J_ . 
wetting j I' I I I I TT I I I I I I I I I T-1 I I I I I I I I I I II TT I I I I I I I I I I 
1�1 ; r1 u. t·- i.  �nrl tenths o f  minutes 
Plate 25 . The respiratory pattern of a shorn sheep before and after wetting . 
f--1 0 CO 
Before 
wetting 
After 
wetting 
' 
Minutes and tenths of minutes 
Plate 26. The respiratory pattern of an unshorn sheep before and after wetting. 
I-' 0 \,() 
�) <D � +> -� r-i 
<t-t 0 
[/) ..c: 
� <D +" 
-u � <l1 
(fJ (J) � +> 
-� 1---l 
Minutes and tenths o f  minutes 
Plate 27 . Oxygen spirometer graphs made with an eight minute gap between 
recordings using the same sheep . ...... ...... 0 
lll 
into the repeatab1lity o£ records  made within a £ew minutes o£ each 
other . 
On 28/12/56 a £1nal recording (Table 10) was taken £rom all 
animals . This revealed that one month a.fter shearing and in summ r 
temperatures the respiration rhythm was still slower and deeper 
than it was before shearing. 
(b ) The effect on their metabolic rate of feeding 1 lb . of oats 
to ach of �hree sheep after twelye hours starvation. 
Introduction 
ll2 
It has been found that shortly after the talcing of food there 
is  always a marked rise in the rate of heat evolution as measured 
directly or indirectly. 
Armsby (1906) dealt very ful ly  with the subject of the increase 
in metabolism due to feeding and remarks that this increased x-
penditure is often, although rather loosely, spoken of as the 
"work of digestion" . Modern authors refer to it as the Specific 
Dyn��c Action of the foodstuffs . 
This isolated experintent was carried out to asoartain whether 
or not the technique was sui table for this type of nutrition study. 
The advantage of the technique is that such studies could be carried 
out on large numbers of animals and this would enable between animal 
and between breed reactions to various rations to be ascertained. 
Procedure and Results 
On the afternoon of 4/1/57 three of the tracheotomised sheep , 
two Cheviots and one Romney, were confined in the small recording 
pens . They were left without food but with water available until 
early the following morning. Oxygen spirometer records were then 
made after a fasting period of about twelve hours . The sheep w re 
subsequently fed one pound of crushed oats each. The oats w r 
consumed voraciously, the animals havin been selected partly on 
llJ 
their liking for concentrates . 
Shortly after feeding was completed oxygen spirometer records 
were again made , the animals being taken in the sequence in which 
they finished their oats (Table 11 ) .  In every case the rate of 
oxygen consumption had risen quite markedly when compared with the 
fsst�g level . No further food was offered and metabolism records 
were taken approximately every hour until about 4 :15 p.m. when 
each animal was again fed one pound of oats . Two further oxygen 
spiromet r records were then taken . 
It will be seen from table 11 that the increase in oxygen 
consumption due to feeding one pound of oats was not at all regular 
but that the fall which followed the peak readings was uninterrupted 
except in the case of w1�vi0� B where the 5 :�0 p.m. reading was 
higher than that of 4:55 p.m. 
Table ll. The oxygen consumption records in litres consumed/five minutes* of three sheep 
each fed 1 lb. oats after twelve hours starvation and again eight hours later. 
Cheviot A 
Time 
7.4Sa.m. 
8.40 
10 .05 
ll .25 
1,25p.m. 
3 ·35 
4.15 
4.45 
.5 .30 
�*consumption 
litres/ 5 mins . 
1.03 
1.41 
1.32 
1 .21 
1.11 
0 .93 
1 .25 
1.25 
Cheviot B 
Time 
7•25a .m. 
8.55 
10.20 
11.35 
1.40p .m. 
3 ·45 
4.5.5 
.5 .40 
o2•consumption 
litres/.5 mina .  
1.17 
1 .32 
1 .)2 
1 .16 
1.11 
1 .02 
1 .11 
1.21 
• Corrected to Standard Temperature and Pressure 
Romney 
Time 
7•35a .m.. 
8.30 
9·55 
11.15 
1.10p .m. 
3·25 
4.30 
.5 ·20 
o2•consumption 
litres/5 mins .  
1 lb. 
1 .  03 crushed 
oats fed 
1 .27 
1.27 
1.2.5 
1 .22 
1 .02 
1 lb. 
crushed 
oats fed 
1.40 
1 .35 s 
ll.S 
( c )  The effect of 1 thyroxine on the oxygen consumption. respiratgry 
rate, and bogy weight of one Rompey Marsh ewe. 
Introduction 
In connection w1 th studies which were being carried out at 
Massey Agricultural College (Kirton 195?) on the effects of thyroxine 
on body weight and its components , the writer was asked to co-operate 
in ascertaining the result of a certain dosage rate of 1 thyroxine 
on the energy metabolism of ewes . A review of literature has been 
compiled by Kirton and the following references are cited from it. 
Cited Literature 
Although in low doses , thyroxin as such or as iodinated casein, 
has been used as a growth stimulant . it is well kno�n that hyper-
thyroid animals receiving thyroxine at levels above physiological 
normality, lose weight (Blaxter .!! y. 1949 ) .  In sheep , weight 
losses under thyroxine treatment are common , (Hart 1955 ; Jordan 
19.54; Warwick n .@l. 1948) .  Turner and Reineke (1946) used weight 
loss in this speci-es as an assay technique for the thyroxine potency 
of thyroprotein. Blaxter (1948)  working with wethers recorded lo ses 
ot up to 28 percent of th original body weight in 24 days. In 
ruminants it appear that one of the ways that thyroxine reduces 
li veweight may be by lowering the volume of gastro-1ntestinal 
contents . This could be caused by an increased rate ot peristalsis 
or in ore severe cases of hyperthyroidism by lowered food intake 
or by starvation. In cattle Blaxter !1 il• (1949) report d that 
ll6 
7 .7 percent sho�ed signs of scouring or digestive upset as compared 
with 1.7 percent of the controle . 
As it was no part of this project to develop a detailed study 
of the eff�ct of thyroxine on the metabolic rates of sheep , no 
original review of literature was attempted although Brody (1945 ) 
is cited for the following reference . 
Magnus-Levy ( circa 1895 ) discovered that loss of the thyroid 
may reduce the basal energy metabolism by nearly half, and that 
suitable thyroid feeding may aln.ost double the metabolic rate. 
Metabolic rate was therefore adopted as a measure of thyroid function . 
For the present work starting 28/1/57 measurements were taken 
on the influence of thyroxine injected daily on the metabolic rate 
(oxygen consumption) of a five year old Romney ewe , fed hay � 
libitum. This animal had previously been involved in metabolism 
studies (R.  No. 2 ) .  Her lamb was weaned at 29 lbs .  having gained 
0 .35 lb. per day which compar d very favourably with those of other 
ewes (Table 5 ) .  The results of the thyroxine treatment on oxygen 
consumption, respiration rate and body weight are presented 1n table 
12 . 
This sheep was quite thin and it was thought unlikely for 
any change to occur in the surfac area of the animal. The oxygen 
consumption figures are therefore presented corrected to Standard 
Temperature and Pressure only� The results show that for this 
animal the thyroxine treatment raised its oxygen consumption b,y as 
, . 
.. .. ' 
Table 12 .  The effect of  thyroxine treatment on the oxygen consumption. respiration rate and 
li veweight of one Romney Marsh ewe. 
Oxygen consumption+ 
litres o2/5 mina • 
Respiration Rate 
-
28th Jan. 1957 1 .0  55 
29th " 1.0  50 
JOth " 0 .9 50 
Jlst " 0.9  50 
1st February 1.0 60 
2nd lt 1.4 70 
Jrd tl 1.4 lOO 
4t.h " 1.8 110 
5th • 1 .. 6 110 
6th .. 1.1 80 
8th 11 1 .. 1 40 
20th February 1.,2  50 
24th 11 1.,0  40 
26th • • 60 
27th • 1 .. 1 45 
28th • l.J 50 
lst March 1.5 90 
2nd " 1.6 80 
Jrd If 1.5 lOO 
4th 11 1.J lOO 
5th If 1.2 90 
6th tl 1.2 80 
7th "' 1.0  60 
8th " 1.0 60 
+ Corrected to Standard Temperature and Pressure 
Li.veweight lbs . Treatment 
- 5 mg thyroxine 
62 10 
59 5 
- 5 
59 ' 5 
59 
58 
60 
61 
75 
70.5 5 mg thyroxine 
68 5 
65 .. 5 5 
- 5 
60 5 
57 
60 
60.5 
64 
67 
66 
• An air leak developed and so this reading was discarded. This did not effect respiration 
rate. 
B 
ll8 
much as 80 percent (it was increasing when the treatment was stopped 
in both cases ) .  The respiration rate was doubled at the peak point 
and live weight was reduced. A lag of about two days occurred between 
the first injection and the change in the pattern of oxygen consumption, 
respiration rate and live weigh� There was a distinct carry over 
effect of several �s duration for both experimental periods . 
It was noted that the output of faeces was quite considerable 
on about the second day after the commencement of thyroxine admin­
istration and that there was a tendena,y to scouring on the fifth 
day in both exper�.:-=-EI�tal periods . On both occasions the animal 
suffered a very considerable decline in appetite whi ch did not 
return to normal until some d� after the cessation of treatment. 
Since these results agree substantially with the known effects 
of thyroxine administration to humans and farm animals it is concluded 
that the technique here used is suitable for such investigations . 
119 
(d)  �ethane exhalation in sheep. 
Introduction 
In view o:f the statements made Brody (1945) , Ritzman and 
Benedict (1938) , and Blaxter (1954) to the e:f:feot that the ruminant 
exhales methane in considerable quantities it was deemed necessar.y 
to investig�te the amount o:f methane which accumulated in the 
oxygen spirometer during metabolism recordings .  Accordingly, 
appropriate arrangements were made :for gas anglyses to be carried 
out by the Dominion Physical Laboratory• at Wellington. 
Procedure and Results 
On 14/3/57 seven animals were chosen at random and each in 
turn was atta ched to th oxygen s irometer in the no:nnal manner 
until it had exhausted one half the reservoir capacity--a period 
of about ten minutes . 
The gas remaining in the spirometer was then expell d and 
collected over water (Dominion Physical Laboratory instruction ) .  
The samples , taken 1n one litre Winchester bottles ,were promptly 
despatched to Wellington. The results of the analysis of these 
amples are presented 1n table 13 . 
• Particular thanks are due to the Director of the Dominion Physical 
Laboratory, Wellington, for offering the necessary laboratory 
staff and :facilities for this work. 
120 
Table 13 . The results of analys s of gas samples drawn after the 
sheep had exhausted half the resevoir capacity of a 
closed circuit oxygen spirometer . 
Sheep No . 
Methane 
10 20 
0 .15� 0 . 2� 
2 
0 .15% 
5 
0 .15% 
X 
Hydrogen ( 0 . 02% or less )  ( ------------not found--------------- ) 
There was no possible lvay in which the methane could have 
entered the spirometer other than from the lungs of the sheep. 
However , in view of the fact that in eructation some eructated gas 
does flow back down the trachea--this is believed to be an original 
observation--the presence of traces of methane in the pulmonary 
system may be due to residues from previous eructations . When 
applied to the total gas volume of approximately two litres remaining 
in the spirometer , the methane percentages give maximum, minimum 
and mean values of 6 ,  1 ,  and 3 ml.  respectively of methane exhaled 
in approximately ten minutes . 
As the spirometer graphs cannot be ruled to a greater accuracy 
than 0 . 05 of a litre , a possible volume error of at the most 3 ml.  
of methane per five minutes is considered to  be low enough to  be 
ignored . 
121 
( e )  Regurgitation in sheep. 
Based on the work of Bergman � nd  Dukes ( 1926) , who acknowledged 
the investigations of Toussaint (1875 ) ,  Colin ( 1871 ) and Fluorens 
(�844) , the generally accepted current theory of regurgitation is 
that a quick inspiratory effo�' with a closed glottis leads to a 
sharp fall in intrapulmonary pressure with the lungs tending to 
contract towards the rigid walls of the " costal box" . This in turn 
is  believed to create a negative pressure within the thoracic 
oesophagus causing it to dilate an d  so allow the highly fluid 
rumen contents to siphon rapidly through the relaxed cardia when 
it is transported to the fauces by its own momentum aided by an 
anti-peristaltic wave in the oesophageal wall. 
Bergman and Dukes (1926) found that breathing and mastication 
are interrupted for several seconds at the time of regurgitation and 
that there was no increase in chest movement or rectal pressure and 
only a small and apparently insignificant momentary increase in 
intra-ruminal pressure . On the other hand a small cannula inserted 
into the trachea and attached to a recording manometer shot-Ted a 
sharp fall in intra-tracheal pressure at the moment of regurgitation. 
By using an animal with an established �en fistula they were able 
to demonstrate , by direct examination, that a negative pressure 
occurred at the cardia at the instant of regurgitation. Duk es 
( 1955 ) states ,  " • • •  in regurgitation the entrance of food into the 
oesopha s is  brought about by intra-oesophageal negative pres sure 
due to an inspiratory effort with a closed glottis . " He also 
quotes Kryzwanek (1934) as an authority for the statement that 
ruminants with pneumothorax or an opening into the trachea are 
unable to regurgitate or do so only with difficulty. 
122 
Another modern worker. Stigler (1931 ) .  also investigated the 
mechanism of rumination and reviewed earlier theori�s paying par­
ticular attention to those of Toussaint . Colin 1 Wester and Mangold. 
According to the above worker. Chauveau and Toussaint took 
the view that the entrance of the ingesta into the oesophagus is 
accomplished by an inspiratory effort with a closed glottis .  
Colin is credited with the belief that the oesophagus relaxes with 
regurgitation and forms a funnel shaped widening into which the 
half fluid forestomach contents stream . The same author writes . 
Wester' s theory was that the oesophagus sucks up the regurgitate 
by its active widening without the inspiratory fall in pressure 
being necessary. He believed that the cardial opening occurs when 
the diaphragm first contracts while simultaneously the circular 
muscle ring in the caudal part of the oesophagus is relaxed and that 
the oesophagus contracts in its entire length. Thus the posterior 
end of the oesophagus must be widened . The opening of the cardia 
is. according to Wester. assisted by a tretching of the stomach 
wall round the cardia. 
Stigler lists Mangold as the originator of the theory that 
primarily no significant activity of � organ may be necessary 
12j 
to cause the ruminating mass to go from the forestomach to the 
oesophagus as soon as the cardia opens . According to 1angold the 
greatest quantity of water giving sufficient difference in hydrostatic 
pressure to bring food into the oesophagus on opening of the cardia 
is 10 to 15 cm.  in the goat , and as one can see clearly in X-rays , 
the opening of the oesophagus into the stomach (in the standing 
animal ) lies deeply under the level of the rumen contents and under 
the air bubble in the food . Stigler comments that transport of the 
regurgitate by hydrostatic pressure alone is comparatively slow 
whereas in X-rays one sees that the ruminating mass is conveyed 
at great speed from the cardia through the cervical oesophagus to 
the neck . Stigler considered that this speed is brought about by 
a force derived from : -
(1 ) Greater pressure in the forestomach following its active 
contraction , or by abdominal pressure . 
(2 ) Decrease of the pressure in the oesophagus either by 
inspiratory lowering of the intra-thoracic pressure, or 
through active widenin of the oesophagus . 
After a lengthy investigation, Stigler ives the following 
sequence of events as his explanation of how regurgitation takes 
place and comments that the mechanism of insuction described cor­
responds to the theory of Chaveau and Toussaint. 
(1 )  Rumination begin as a rule with the swallowing of saliva. 
(2 ) Immediately after this appears a quick contraction of the 
, 
oral parts of the thoracic oesophagus , apparently in order to 
press out swallowed air.  
(J ) The glottis closes and the insuction phase begins . 
(4 ) The diaphragm makes a backward inspiratory movement. 
( 5 )  At  the same time the head and neck are flexed more or less 
dorsally. This has the object of improving the closure of 
the cervical oesophagus and hindering the jnsuction of air 
from the mouth into the thoracic oesophagus . 
( 6 )  Following the fall in the intra-thoracic pressure brought 
about by the inspiratory movement with a closed glottis , the 
thoracic oesophagus is sucked open and a decrease of pressure 
occurs inside it . 
( 7 )  S�muJtanoously the cardia opens . 
( 8 )  Some of the rumen and reticulum contents , mostly as a fluid 
broth , are sucked up into th� oesophagus by the negative pressure 
in the thorax and thoracic oesophagus and are carried by an 
anti-peristaltic wave to the mouth . 
As objections against the acceptance of this theory, Stigler 
notes that it is said , rumination can still take place after: ­
(1 ) The cutting of both phrenic nerves 
( 2 )  Tracheotomy 
( J )  Pneumothorax 
He states however, that these objections are not acceptable for 
the following reasons :-
( 1 )  According to experiments on the cat , one quarter to one 
half of the usual inspiratory force remains after the 
paralysis of the diaphragm through the cutting of the 
phrenic nerves . Such a remaining force would be suf­
ficient to suck up the regurgitate . 
( 2 )  Tracheotomised animals ruminate only with a very strong 
respiratory movement and the respiratory variations of 
l2.S 
the chest circumference are much larger than with normal 
rumination because the checking of the diaphragm movement 
by closure of the glottis does not take place . The breath 
rush is very much louder than under normal circumstances . 
(J )  Experiment showed that a goat with pneumothorax could not 
ruminate in spite of strong endeavours on its part , but 
sixteen days after the operation, when the wound had 
healed , the animal ruminated again . Stigler argues from 
this that it is obvious that the oesophagus , diaphragm 
and abdominal muscles together are not sufficient to 
produce rumination, since an intrathoracic fall in pressure 
was not produced by inspiration, and that this is a very 
strong objection to Wester ' s theory that the oesophagus 
may suck up regurgitate without the respiratory fall 1n 
pressure being necessary . 
However, observations on sheep tracheotomised as described in 
Division "B" , this project , made it abundantly clear that the 
126 
st2tements that regurgitation occurs only with difficulty in trach­
eotomised ruminants could not be borne out . 
Close daily study of seventeen sheep tracheotomised for periods 
of up to ten months and of a simila.rly intubated Jersey bullock calf 
for four months (Plate 28 ) revealed no interruption of the rumination 
pattern, irrespective of whether the tracheal opening was open or 
closed. 
Spirometer recordings (Plate 29 ) frequently showed a transient 
increase in rate and decrease in depth of respiration followed by 
a much greater than normal inspiratory intake coincident with each 
regurgitation . However, direct observation of the animal gave no 
outward indication of a deeper than ordinary intake of brerth at this 
moment although the extra intake was generally of the order of 150 
to 250 ml .  The greater than normal inspiratory effort was not 
always made , for some animals appeared to be able to regurgitate with 
very little or no effort at all and with no dorsal flexing of the 
head and neck as described by Stigler. These observations were 
made in many cases when the animals were conn�cted to the spirometer 
and the writer was sitting virtually at their feet. It was therefore 
thought that the ct .... .phragm must contract much more forcibly at the 
moment of regurgitation whereas the remainder of the muscular 
apparatus involved in the respiratory act maintains its normal 
t.bythm. Quite an appreciable force must be required to draw an 
extra 200 ml .  or thereabouts of air into the lungs , and such a 
force applied with a closed glottis in the intact animal would cause 
. � rl C1l tJ 
'0 (I) {/) ·ri 
5 +l 0 (I) ..c tJ C1l � +l 
� 
. 
CO N 
(I) +l 
cu rl p... 
127 
s:: <1l 
� 0 
Ul (I) � +' 
·r-i rl 
<H 0 
Ul ..s:: 
� <ll +' 
§ 
(/) <ll � +l 
·r-i ....:! 
H-
=t= 
-+-
T 
1. 
' 
I . 
-t -
I 
··- _t 
-� 
I - � ·  
'it- -�-1· 
' - - __;}-'ll . 
'a f-- � l--
+ -
T T • I 
..., 
' ' + ]_--1-� . !--,_. _! 1 _1 +. - +· . 
tt��- I . I I 
i_ H 1 - _I + �  -� -+ 
_:-..!...Ll..l L.�-I _L__i_ _L 
Hinu t e s  and tenths of minu t e s  
-
·-
- I -
·m�-I++ -
-
j 1--1_ � 
+ -
t-q --
r ' - �r-
Plate 29 . The respiratory pattern of a ruminating sheep . The points of regurgitation 
are marked by the large upward blips 11 X11 • f-' l\.) ()) 
129 
a marked intratracheal and intrapulmonary negative pressure at the 
critical moment . However, the belief that closure of the glottis 
and the resultant negative pressure are essential to the regurgitation 
reflex is clearly invalidated by the behaviour of the tracheotomised 
sheep in which the presence or absence of additional negative 
intratracheal and intrapulmonary pressures appeared to be entirely 
irrelevant to the operation of the reflex. It is thus perhaps just­
ifiable to allow that Wester ' s hypothesis may have s��e merit despite 
the attack that has b en made on it . 
Stigler' s observations that there is a closing contraction of 
the oesophagus to re�ove swallowed air and that the strength with 
which the oesophagus of a cow can contract is considerable , lend 
some support to the opinion which 1..ras formed as a result of the 
present work. This opinion was that at the moment of regurgitation 
the cardia and posterior portion of the oesophagus dilate and 
simultaneously the oesophagus is jerked into a state of relative 
tension by a sudden forcible contraction of itself and the diapl1ragm. 
This would create a negative pressure within the oesophagus sufficient 
to cause the stomach contents to siphon rapidly forwards , although 
the inspiratory effort must also produce intrathoracic negative 
pr ssure which may hav a primary or s condary effect . Th supposed 
jerkin of the oesophagus into a state of tension would possibly 
be aided by the skeletal muscle which runs throughout its entir 
length . In this latter respect the ruminant oesophagus differs from 
) 
lJO 
that of most other species . After the entrance of the regurgitate 
into the oesophagus the cardia closes and the fluid bolus is carried 
to the mouth by an anti-p ristaltic wave in the wall of the oesoph-
agus . 
It is nevertheless recognised that all inspiratory efforts , 
glottis closed or not , are made as the result of the creation of 
intrathoracic negat1 ve pressure and so any "Oesophagus-I:;iaphragm" 
11 
hypothesis cannot but be extended to include Inspiration� 
Indirect evidence in support of an "oesophagus diaphragm 'j 
hypothesis is supplied by some recent work of Dougherty and 
Meredith (1955 ) and Dougherty and Habel (1955 ) .  In the cours 
of a study of the bloat problem these workers investigated the 
eructation mechanism by means of an X·· r"l.y and cinefluorographic 
technique and were thus able to obtain a continuous visualisation 
of the eructation process and to demonstrate the presence of three 
oesophageal sphincters . These were : a very efficient cranial 
sphincte r normally keeping the entrance to the oesophagus closed; 
a second situated a short distance anterior to the diaphragm; and 
thirdly the cardia itself . Their films also show that the walls 
of the oe�ophagus , particularly in the rumen terminal position, can 
be remarkably active . Dougherty, in a personal communication (1956) ,  
stated that in similar unpublished cinefluorographic studies he 
observed a sudden sharp diaphragmatic  contraction at the moment 
of regurgitation. 
DIVISION "C" 
RANGING BEHAVIOUR STUDIES 
CON!ENTS 
Page 
C·I Introduction and review of literature 133 
C-II Technique development 140 
C-III Experimental procedure 149 
C-IV Results 159 
C-I INTRODUCTION AND REVIEW OF LITERATURE 
lJJ 
The aim of this Division of the overall project was to compare 
the mileages traveled by New Zealand Romney Marsh and Cheviot sh ep 
grazing on hill and lowland pastures . It was also hoped that it 
would be possible to study :-
(1 )  The effect on activity of different sized pastures within 
the above two locations . 
(2 )  The result of switching the two breeds from hill to flat 
and from flat to hill land . 
Literature d aling specifically with the distances traveled 
by grazing she p is very s oanty. 
Louw .!1 al. (1948) in South Af'rica used a checker-board technique 
in which they divided pastures into 20 acre squares by means of 
stakes . Hourly observations made by means of binoculars traced the 
movements of the sheep in relation to the takes . Table 14 presents 
the results of the above work . 
Tribe (1949) in North Scotland studied the general features 
of the grazing behaviour of Cheviot sheep on one acre of lowland 
pasture over continuous periods of 24 hours during 12 consecutive 
months . The one acre pasture was divided by stakes at ten yard 
intervals following the method of Wallaoe and Kennedy (1944) .  He 
reports that th sheep walked an average of 2.60 miles per 24 hours : 
2 .06 miles being between 7 : 00 a.m. and 7 a 00 p.m. and 0 .55 miles 
between 7 r00 p.m. and 7 : 00 a .m. Most traveling was done in the 
sprin and autumn, and least in s er. h animals walked propor­
tionat ly mor at night in summer . than in winter. (Tribe does not 
Table 14. The daily distances walked by different sheep breeds (Louw et al 1948) singly and 
in various combinations . 
Black Headed Dorset X Persian Karakul Merino Persians 
Daily Distance Traveled 
Yards Yards Yards Yards 
lst day together 5229 6738 4868 4836 
2nd " tt 4580 4053 3764 2795 
3rd " " 6405 4845 5075 5945 
1st day separately 4555 5505 3325 4515 
2nd " lt 6240 5645 4260 3350 
Merino & Karakul Dorset Black Headed Black Headed & Karakul (Together) X & Persian Persian Persian (Together)  (Together) 
Daily Distance Traveled 
Yards Yards Yards Yards Yards Yards 
2nd day 7285 6380 9685 8660 7995 6035 
3rd daJr 6160 7410 10575 8275 8970 6860 
2nd day 456o 5l10 7780 7430 7125 4290 
3rd day 4680 4575 4575 9385 7055 5685 � 
lJS 
ention the po eible r lationehip 'bet-w en t.he short night of the 
Northern er and th se resul \a) . The method he adopted vu t.o 
follow the b haviour of only one abeep a t  a time . Tribe report.e 
"iult 1n con.f'i Uon of th work or Stapledon and Jonea (1926) 
and. Jones (1926) , it was found that the record from a inale animal , 
on each eoc i on  Nflected. t.he actions of the expel'i ntal group . 
and (1954) , working in t les , produced the follo\dn& nsul ta 
from rour or di ffe rent breed t,ocethttr w1 th their lamb on 
two plots each of 1.9 acres (Table 15) .  
Table 1.5. The 'total distance (in 7ar.d ) t. veled by e eh ehe p 
durin twenty faur hour periods (En and 1954). 
Eve at Day Second Day 
Blaokt oe 15.58 2888 
Cl.un 2268 3112 
Spaniah 2400 )876 
Suffolk 1922 42 
Diatance (11'1 yards) travel by the she p trom 10. p • •  to 8 •••• 
n . a . s . T: 
Eve 
Bl oktace 
Clun 
p h 
Suffolk 
• ble Bri t1 h S er 
Firs t  Dq 
216 
2� 
292 
,98 
.Althoup ho vea no diet.ances, 
Second Dlq 
.sso 
644 
4)6 
8.52 
ter (1954) • J'V88 that hill 
ehe p are ore aoti e than lo land he p duritlc the hours f4 ciayllpt. 
o ta wer avaUable on the nipt 'b · viour ot bUl beep but Bun r 
136 
(who worked in Scotland) believes it quite probable that they are 
more active than lowland sheep . The .following data (Table 16) are 
presented by this author : 
Table 16. Percentage time spent resting in the hours 7 : 00 a.m. -
7 : 00 p .m. by lowland and bill sheep, April - September 
(Huntet" 19.54) . 
Apr.  May June July Aug. Sept . 
Lowland 
(Tribe ' s  Cheviots 1949) 32 32 40 45 45 47 
Hill 
(Hunter• s Scotch Black 
.faced sheep 1954) 13 13 21 26 
Hunter overlooks the fact that he is comparing not types of 
sheep but locations , !or the Cheviot is not a lowland type . This 
kind of contusion of the sheep breeds can only be deplored. 
23 
In a comparative study of the grazing habits of sheep carried 
out by Van Rensburg in South A1'rica (1956) the .following results 
were obtained (Table 17) :  the technique is not specified. 
T ble 17. 
Breed 
Merino 
The average daily distance walked by four she p breeds 
(Van Rensburg 1956) .  
Averag daily distance walked 
(yards) 
G r.man Merino X Merino Cross 
449.5 
6532 
4845 
4980 
Blackhead Persian 
Dorper 
137 
Tribe {Hammond 1955 ) reviews the literature on grazing behaviour 
studies . Excerpts from the review are now presented in order that the 
present work can be assessed in the light of this modern summary. 
Tribe writes , that valid and standardised techniques are the 
prerequisite of any scientific investigation . and it is an important 
criticism of grazing behaviour studies in all parts of the world 
that there is no standardised technique of observation , and no 
oomparisbn has yet been made of the accuracy of the many different 
methods used. There is , for instanc , no uniformity concerning 
even the period during which observations should be made. Some 
workers have mistakenly assumed that during the hours o! darkness 
animals will remain at rest and have therefore only recorded their 
observations during th hours of daylight. 
Tribe continu a--similarly the methods of recording vary 
considerably . Some workers record the behaviour of individual ani.Dlals 
while others record the behaviour of flocks or herds . Some workers 
make a continuous recording while others record only ob ervations 
taken at regular and frequent intervals .  Some records are the work 
of singl observ rs while others are ade by several observers 
working on a shift syst • Tribe further remarks that in an effort 
to increase the accuracy o! behaviour records a number of automatic 
and semi-automatic r oorders hav been volved• and 1n use with 
anim.als penned or in stalls the e auto-r co:rding ebtnes have 
proved very suce s f'ul .  However, · according to  Trib , the difficulties 
ot u ing such machines to r cord gr zing b haviour are of course 
much greater and have not yet been overcome . He mentions that 
Burton and Castle (1950 ) have described the construction and u e 
of a portable infra-red ray equipment for animal observation in · 
lJ8 
the field . For watching from a stationary position not more than 
four animals at a rang of ten �o fifteen yards on a clear dark 
night the equipment was ideal . \1/hen using it to observe a large 
number of animals either in the field or in the cowshed the obs rver 
was liable to miscount the animals owing to the limited field of 
the screen. Also, the constant wearing of the equipment which 
weighed 21 pound was very tiring and, when the weather was misty, 
the lense quickly clouded over. 
Tribe considers that the differences between the animal behaviour 
patterns describ d by various workers are not surprising • In addition 
to the differences in observational technique , there is a complex 
of environmental factors which must be expect d to influence a 
normal pattern of behaviour. For example , such variable factors 
as th climate. the density and quality of the sward, the ize of 
paddock, th system of grazing managem nt .and of cours , the in­
dividuality of the grazing animal may v ry significantly influ nee 
the times devoted to different acti vi ti s .  
Tribe discusses these and other factors a they affect many 
functions of th grazing animal . A thi project is primarily 
concerned w1 th activity • further diseu si on will be limited to the 
stated effect of various factors on this item, 
139 
l, Cli.Jnate 
Bot weather causes animals both to graze for a shorter 
time than normal and to increase the proportion of time that is 
spent grazin at night , When idling under these conditions 
the animals app ar agitated and in consequence the distances 
traveled tend to increas e . 
During cold windy or rainy weather grazing times are again 
shortened. and the normal overall pattern may be radically 
altered, When a storm begins animals will usually cease 
grazing and walk to shelter if such is c l o s e  at hand. otherwise 
they will stand with their heads down,..wind , Under such conditions 
hiU sheep and cattle will move to higher ground, 
2, Size of Pasture 
It has been shown on several occasions that the size of 
the pasture available to the grazing animal will influence 
its: behaviour and in particular the daily cftstance it tr v ls , 
In general one can say, the bigger the area of pasture the 
farth r will an a.ni.mal walk, Shepherd (1921) reported that 
beef cattle on a JO acre pasture walked one and fiv eighth 
miles between 4 : 00 a.m. and 8:00 p . m . whereas simUar cattle 
throughout the same p riod traveled three and on sixteenth 
mil s on a lOO acre pastur and ti v and a halt miles on a 
640 acre pasture . There must obviously be limits to thi 
tendency at both ends of the seal , but so tar they have not 
been established, 
140 
3, Indi viduali t.y 
Perhaps the greatest and most imponderable problem which 
besets the animal watcher is the degree to which recorded 
behaviour may be due to environmental factors and how far 
merely to the personal whim of the animal . 
Hancock (1950) using six sets of monozygotic cattle twins 
in a one acre paddock showed that inherited variability in 
razing behaviour was by far the largest source of variation 
due to individuality. A part of thes inherited differenc s 
could be explained by relating them to differences between the 
physiological requirements of individual animals. It seems 
reasonable to argue that increased nutritional requirements 
during growth, pregnancy, or lactation, result in an increased 
food intake and therefore in increased grazing and ruminating 
times . 
C-II TECHNI\:UE IEVEWPHENT 
In approaching the problem of measuring sheep activity, as 
expressed in miles t�aveled per w ek, certain difficulties had to 
be con idered ' 
(1) Work on the other divisions of the overall project would 
limit tb amount or time available . 
(2)  It was n oes ary to  measure th ranging mil age of  th 
sheep und r natural conditions on hill and lowland arazings . 
For this reason it was impossible to use the checker-board 
techn1.que (1valla.ce and Kennedy 1944) .  Even where the 
terrain would permit subdivision, no time was available 
£or 24 hour observation shi£ts . 
141 
( ) )  Radio control , pedometers and other such devices £or 
tracking movement were all too complicated or impossible 
to use satisfactorily on the grazing sheep . 
The restrictions thus imposed on the work subsequently led to 
the development of an entirely new device. This device took the 
i"orm of a machine which it has been proposed shall b known as a 
Rangemeter. 
The Rangemeter consists of a light-w ight harness , two metal 
shafts , and a chassis wP�oh sprung onto the ground carries a small 
land wheel to motivate an automatic mileage recorder--a Lucas 
oyclometer*-•as used on bic,ycles . The total weight of the machine 
was about 8 pounds , but the sheep only carried ) pounds or half 
the ' spring load d' weight of the shafts which at one end are 
pivoted on the shoulder harness and at the other skidded on the 
ground . The draught of the machine was exceedingly light, a pull 
of only approximately one pound being required to move it along 
the flat and on the typical hill slope . 
The detail d construction of the apparatus is giv n in plat s JO , 
Jl, )2, )j , and their ke,y. 
Development or th first machine took place at Ma s y Agri• 
• Joseph Lucas & Co. , Ltd. , Birmingham• Engl.and 
a :: 
b = 
c = 
d = 
e = 
r = 
g = 
b = 
i = 
j = 
k = 
1 = 
m == 
n = 
0 c: 
p = 
q = 
r = 
8 = 
t = 
u = 
KEY TO "RANGEMETER" PLATES JO, 31, 32. 33 
34" adjustable sha.fts 
13" 
11" 
2" 
3" 
6t" 
2t" 
9'' adjustable belts 
28" adjustable belts 
1211 adjustabl belts 
6t" diameter wheel 
harness clamp 
chest strap 
5" bolt 
leather strap ( spring) 
oyclometer shield 
cy cl o  eter (26" whe 1 )  
striker 
hard steel shaft 
hara steel bush 
142 
14J 
design and construction 
1� 
Plate 31 . Showing details of the design and construction 
of the Rangemeter. 
Plate 32 . Showing details of the design and construction 
of the Rangemeter.  
145 

147 
cultural College using one Romney ewe. Further machines were 
constructed as soon as it was obvious that the principle was sound 
and mechanical defects were remediable ; e .g . , the land wheels were 
originally fitted with one inch wide brass bushings which theoretically 
should not have worn to any marked degree . However , the bushings 
ground themselves loose on the steel shafts and the subsequent 
lash in the movements of the wheels was sufficient to result in 
bent strikers and cyclometer s ha ft s . Eventually the brass bushings 
were chru1ged for others made from hard steel tubing which revolved 
on the full length of the shafts (Plate )) ) .  This type of bushing 
gave very efficient service . fhe greatest problem was the tendency 
for long grass stalks to wind themselves round the revolving shafts 
and thus cause an obstruction to the working of the strikers . 
A ball bearing axle and a plate set just below the shaft to 
fend off grass , will probably overcame all of the technical dif­
ficulties of the mileage recordeq and these are to be fitted in the 
future ma chines . 
The final and most satisfactory type of harness developed was 
that shown in plate )0, The machines complete with harness could 
be removed from, or replaced on• the sheep in a matter of seconds 
merely by unfastening one buckle . This simplicity of arrang ent 
was beyond value to the success of the work. 
It is e timated that some of these machines must have trav led 
approximately 500 iles , which is a testimony to their d sign and 
construction. 
1� 
There was no tendency for wheel slip to occur. As can be seen 
from plates JO and 31 the pivoting of the wheel chas s i s  means that 
its whole weight bears down onto the ground . In addition the pivot 
is fitted with a leather spring (Plate 31) wh i ch allows the wheel 
to rise and fall •4ith the terrain but nevertheless keeps it fi� 
pressed to the ground . This spring is essential to the success of 
these machines . 
149 
c-III EXPERIMENTAL PROCEDURE 
On 17/7/56 rangemeters were attached to five each of mature e 
pregnant, Romney and Cheviot ewes from hill stocks c�rried on Tua 
Paka , one of the h1ll fs.nns of Massey Agricrultural College . 
The sheep became accustomed to being in harness in a very 
short time ; after an hour or so they were grazing normally and 
were then left over night. 'I'he following morning the sheep were 
turned onto an 18 acre hill paddock. 
This paddock is extremely steep and rough , contains a small 
stream and is well littered with stumps . logs , old fencing posts , 
etc . Plate )4 illustrates a view of this paddock from one side of 
the central ridge . The height of the central ridge is about 500 
feet and the gradient from the sheep pens at its base direct to the 
top in places approaches ' one in one ' . No better testing ground 
could have been found for both ma chines and sheep . Although of a 
precipitous nature , the hill carri d an abundance of grazing through­
out the year. 
The rangemeters in no way hindered the movements of  the sheep 
which walked , galloped, lay down and got up quite normally. Exp ri­
mental animals wer accepted by the other sheep, which evinced o� 
curiosity about their changed brethren. After parturition the wee 
were quite ble to suckle th ir lambs while in harness . Only twice 
during the whole course of the investigation did a sheep become 
hung up on a log or fence post . 
Plate 34. A view of the holding paddock , Tua Paka , taken from 
the central ridge . 
f--1 \.;\ 0 
151 
A number of illustrations are presented to demonstrate the machine 
at work . (Plates 35. 36 .  37, 38 , 39 . ) 
Mileage recording conh�enced on 28/7/56 and from then until 
24/8/51 the machines were removed only for repairs . lambing . tupping, 
s h e a ring and foot dressing. It was soon di s covered that once a 
sheep was broken to harness  it never forgot and could be left for 
weeks between work periods without it becoming excited when harnessed 
up again. 
Mileage records were read every Saturday morning starting at 
8 : 00 a .m. , wh n the sheep were mustered from their night camping 
spot to a nearby yard or field pen (Plate 40) .  Thus gathering errors 
were reduced to a minimum .  
During the whole period of the Rangemeter project the ewes were 
run as normal sheep • The sheep gave no indication of lack of thrift 
and this was amply borne out by the weights of the ewes on 24/8/57 
(Table 18) .  It will be seen that both Cheviots and Romn ys were 
above the mean figures established for th Haematology flock• 
approximately four weeks earlier. Some of this extr weight was 
no doubt due to the growth of the foetus • but even allowing for this 
the sheep were in very good condition. As a part of the investigation 
it was decided to measure the foreleg lengths of the Haematology and 
Rangemeter sheep. The Haematology sh ep were measured on 13/6/57 
and the Rang eter sheep on 24/8/57 . 
The mea urements taken were height at withers • height from 
• c .t. Division "D" ;  table 27 
Plate 35 · Rangemeter flock sheep mustered for mileage recordings 
to oe noted . 
1-' V\ N 
Ul :s: Q) 
·� 
1.5) 
Plate 37 . Romney Marsh ewe nursing lamb while in harness . 
..... 
"$ 
Plate 38.  Rangemeter flock ewes on their way out to the hill 
after mustering. 
.... \1\ \J\ 
156 
Plate 40 . Reading the Rangemeters . 
Plate 41 . Measuring the height at withers . 
157 
Table 18. Rangemeter sheep weights 24/8/57 . 
Romneys 
No . lbs . wt .  
37 166 
32 131 
36 135 
59 (lamb at foot ) 108 
60 1)2 
No . 
53 
42 
35 
48 
54 
Cheviots 
158 
lbs . wt .  
137 
148 
1)4 
120 
13) 
Comparison of Haematology !lock* and Rangemeter sheep weights . 
Haematolgoy Flock (30/7/57) 
Cheviots 
Romney 
lbs . wt .  St . D.  
125 .4 
120 .1 
10 .9 
� 
ll.6 
* c.t. Division D:  Table 27, 
Rangemeter Sheep (24/8/57) 
lbs . wt .  St • .. D. 
1)4.4  �o.o 
141. 0  lG.l 
(4 sheep ) 
1.59 
ground to brisket and the distance from the point of the elbow 
to the coronet . Measurements were taken with sliding calipers 
(Plate 41 ) and a flexible linen tape . All measurements were taken 
four times to the nearest half inch, the sheep being moved and 
resettled in a normal standing position between the measurements . 
C•IV RESutTS 
The mileage records obtained are presented in table 19 and in 
figure III which is constructed from the t-reekly mean mileages of 
the breeds (Table 19) .  
Statistical analysis of the results was complicated by missing 
records . It was finally decided to analyse the material between 
breeds using the figures that make up figure n r , i.e  • •  the weekly 
breed nleans (Table 19 ) and also to analyse ea ch breed on a bett-veen 
period within breed basis using sheep means . The l'lri ter fully 
realises that analyses based on means derived from unequal numbers 
may oontaL� error which should be compensated for, but having regard 
to the data and after consultation it was felt that little or no 
greater accuracy of interpretation would be obtained by complicating 
the stati ofJti e :t.1 methods used. 
Only when paddocks were being compared wa s  there any cause to 
consider mustering distance adjustments to the records ; but as these 
distances w r all of more or less the same value and at the most 
would give an error of lOO - 200 yards , or about o .l miles per week 
Table 19, ¥Lileage records , Rangemeter sheep, 28/7/.56 - 24/8/57• 
Cheviot: No. 
48 55 35 53 
9.8 8,0 
11,1 7.0 
10.2 7..2 
2.& 
7·5 8,6 5.6 � 
8.7 M 
5.4 6,4 8.5 6.8 
10,8  
6.1 
5•5 
8.5 
7.2  
8.2  
8.3 
6,.3 
6 .•
 3 
4.0 
7o7 7.7 
3-7 
9.0  8.6 
8.1 11.2 ' 
u,2 
.9..1 
.9..1 
J....1 
6.5 8.2 
M 
8,8 
.2.& 
10,3 -
7.4 6,6 1.& 
a,o 9,9 .z.& 
7.2 B. 0 2.!2. 
7.1 9•3 
7.1 7.1 ' 6,4 10,1 ?;.8 
7 .1 8.o 6,2 9·.1 :z..:z. 
7.1 
s.6 6,7 9,6 10.1 � 
10.2 9·9 14.1 10.9 
15.0  7•9  n.o 
Romney - No. 
50 59 36 60 
4.9 5.0  
4.8 4.6 4,1 
l1achines Removed fo.,.. Lambing 
s.o 
4.2 3.9 4,0 
. 3 .? 6.3 4.3 6.1 
3.8 6.7 4.3 }.8 4.6 
3.5 2.4 5.5 3 -5 5•0 
4,0 
5.5 3 .0  
Shearing 
1.5 2,J 
4.2 
7·4 6.7 9.4 
6.o z.1 9.5 
7•9 ?.8  9 -5 
5,6  6.6 6.6 
7.1 
3·1 
5 .7  
6,1 
4.2 
5·7 5·3 4.3 
r•!ac:bines Removed for Repairs 
7.4 8,8 3•9 
5.8 7-9  3•8 
4.6 5.6 4.3 
6.2 
1o.o  5.3 7.8 
13·5 9•3 7·1 
!1achines Removed for Tupping Time 
6,9 
6.0 
11.0 
14.0 
12.0  14.7 15. 0  . 13 ·9  
7·1 9 ·8  6.5 8,2 1.;2 
6,4 14.5 13.8 11.6 
2,4 
6.3 14.1 
9o0  
6.5 8,4 
8,3 7.8 
12,0 11.2 9.1 
4.4 11.9 11.6 12.? 10.9 
5 .1 8,2 10.5 7.8 1.& 
7-3 8d 7.2 8,0 
8.3 3 ·9 
3·1 3·2  
6.o  1o,o 8.1 
6.o 9·7 - 7•3 
6.3 _ 8 .4 7•9 
3.1 5 .1 5·2 
3 .2 5,4 4,4 
37 
5.0  
7•4 
11.4 
32 
1...2. 
4.4 
z.s 
hl 
9·1 
13.0 12,0  8 .8  
7 .1  6 • .8 
4.5 3.3 4.5 
6,2 4.5 .i4 
7•3 6.2 ..2.& 
6,6 ?.4 6.5 
9,4 n.o  10.6 
6,5 2.5 J....§. 
10,3 4, 7 :z.& 
4,8 3,0  4.6 - -
7.1 1..!2 
1o,o 7.7  8,1 · 
5·8 4.5 4.7 
5·2 
Machines . Removed for Foot Dressing Eto, 
8.2 
9•1 13.0 9.5 � 
n,z 14.4 11.3 11.8 
9·8 10•0 _9.1 .2i:i 
9·1 9•9 9-3 9.4 
8, 9 8, 7 8.2 7.4 ' 8(3 
9.1 8.8 11.1 
4.4 5•0 
4,0  6.0 3•7  
3.7  6,o  
2.9  5.3 
6.7 
6.8 
7·8  
7.5 hl 
..2..5. 
5.5 :id 
8,4 7.1 ..!illt 
8,7 
8.o 
6 - 0  =..:. 
Dates. 
1956 
28 Jul. - 4 Aug. 
4 - 11 Aug. 
22 - 29 Sept. 
29 Sept. - 6 Oct. 
6 - 13 Oct • 
13 - 20 Oct. 
20 - 27 Oct. 
27 Oct. - 3 Nov • 
10 - 17 Nov. 
17 - 24 Nov. 
24 Nov. - 1 Dec. 
1 - 8 Dec. 
8 - 15 DeC'o 
22 - 29 Dec, 
29 Dec, 1956 
5th Jan• 1957 
5 ... 12 Jan. 
12 - 19 Jan, 
9 - 16 Feb. 
16 - 23 Feb; 
23 Feb, - 2 J.Tarch 
2 - 9 .March 
9 - 16 Hareh 
16 - 23 March 
20 - 27 April 
27 Ap1, - l1- May 
4 - 11 !iay 
11 - · 18 Hay 
18 - 25 May 
· 25 May - 1 June 
1 - 8 June 
!? . .. 15 June 
15 - 22 June 
29 June ... 6 July 
6 - 13 July 
13 - 20 July 
20 - 27 July 
J - 10 Aug. 
10 - 17 Aug, 
.,: 
ll! � -t-�- J i 
. ,, i 
r • E .. 
! l2 ..a -' 
Figure I I I . Showi n E  ( 1 )  weather data , ( 2 )  average distance traveled in mi�es per 
week f o r  Chevi o t  and Romney t•iarsh ewes , ( J )  p e r i o d s  and loca ticr:s , 
( 4 )  dates of recording r eriods . 
� ::-;oo:.• •• • �U. • :a2•.::l .. • , .... ,"/':.. ac • •• ·�I'� tS 1 ,. ':a-1 ao •• • •• � •• u • •• 2'i1ao �� .. _ • , ..... ;- :;  •• ! !. . .  
i ' 
' I : j .  ... . : . : . . 
. . .  · ·�· . .  . .: . . . . ! _ ' ... j .. 
. l ·  . -·l·.. . . . . .  i . . . .  ; . . . .  � . . .  
! . ; i : 
. . . . .  · . : . . . . . . . :. . . . . .. 1 . . .  ; .... , .. . L .. L � . . . . .  : ... : .... : . . ' 
! : : ' : : . :  ' � . ' 
... i;.:.;;,' . . . ·j.� •. 61.1·� ,;; 
�lia.J· .. �: l ! .. 
, .. .. .  o ... ., .... . 
--- -- -----
' • •• ao aC 
•"'"' 
) 
. .. ... .. ., .. " ...  " 
1 t was decided to make no theor t1cal compensation re., them. 
It is desired to draw attention to the following pointss •  
( l) Tbe difference 1n the position ot the means for the breeds 
(Table 19 and Figure III) . In general., on the hill, the 
Cheviot ranged. about ? .,5 miles per week and the Romney 
about .5 .0 miles per week. ()). flat land the Cheviot 
cavered bout 9 • .5 miles per week, &� the Romney about 
&.o.  In eight periods out of eleven (Table 20) the 
difference between the breed means was statistioaJ..l¥ 
s1gnit1oant. 
(2) The � in which the breeds raised or lowered their mileages 
togeth r • within th same paddock, as in periods A and F 
(Figure Ill) • 
( 3) The wa;y in which the Romneys rdsed their mlleages when 
transferrecl from the hill to flat land and then droppe4 
m � when returned to the hill (Figur III) . 
For exmBple, in the change from the l8 acre hill to the. 
6 acre flat paddock (Figure Ill, perioda A - B) tbe 
R011JD18Ya raised their mileage from 4.4 .. 7 .8 miles per week 
while after the fir t week the Cheviots raised their 
mileage 0� from a hill mileage or 8.2  to flat l.an4 
mileage ot 8.8.  In the transference from flat to bW. 
(FigUre In, periocls B .. C) the Roaleys teU from 6 .8 -
4.1 U s while the Cheviot• actual.l.T raised. their level 
r 
Table 20 . (l)  Average � greater weekly mileage traveled by Cheviot over Romney sheep, within 
periods .* 
Period 
X 
A 
B 
c 
D 
E 
F 
G 
Il 
I 
J 
( 2 )  Significance of difference in yreekly mileages between breeds within periods 
using week means . 
(3 )  Significance of difference between weeks within periods within breeds . 
Average � greater Level of significance Significance of difference 
Location mileage in favour of difference between between weeks within 
or Cheviots breeds periods within breeds 
R c 
18 acre hill) 1% 
) 69� 
18 acre hill) 1% 
6 acre flat 29% no significant difference 
18 acre hill) l� 
) 51� 
18 acre hill) 5% 
3 acre flat 30% 5'� 5� � 
18 acre hill 75'f, approaches � level+ 
l acre flat 2n no significant difference 
18 acre hill 72� 5' 
49 acre bill 82,� 5j, 
18 acre hill 45'f, 1� 
* For the identification of the periods see Fig . III 
+ High variation between weeks within breeds 
- ---------------
� 
{!) 
164 
of activity. The changes in activity between different 
periods were statistically significant in 7 cases out of 
10 for the Romneys and in only 2 instances out of 10 for 
the Cheviots (Table 21 ) .  
(4)  The general lack of varic;.tion in the mileagss travc:""..ed b:; 
both breeds within periods . There w·ere no statistically 
significant differences between weeks within periods except 
for period E in both breeds (Figure Ill� table 20 ) .  
( .5 )  The small value of the rise in the Romney' s mileages when 
transferred from an 18 to a 49 acre hill paddock , whereas 
the Cheviots increased their mileages significantly. 
(Table 19: figure III; table 21 , 29/6-lJ/?/57 . ) 
(6)  The respective mileage levels for both breeds were ap­
proximately the same on 27/7/.5? as they were on 4/8/.56. 
( Figur III . ) 
(? )  The extraordinary steep rise in the activity of both 
breeds before tupping (Figure I II , period E )  and the fact 
that after tupping , which took only three weeks , the ,high 
level of activity was repeated on the one week befor the 
sheep were sw1 tched back to the hill . The difference 
between the w eks within period E was statistically 
significant for both breeds (Table 20. )  
(8 )  With only on exception (Cheviots on larger hill pasture 
Figure III, table 21, periods H to I )  no statistically 
Table 21. Levels of significance of difference in weekly mileages between consecutive periods 
within breeds using sheep means . 
Locations• Periods• Cheviots Romneys 
18 acre hill - 18 acre hill X - A 
18 acre hill - 6 acre flat A - B 1� 
6 acre flat - 18 .acre � B - C 1• 
18 acre hill - 18 acre hill C - D  5� 
18 acre hill - 3 acre flat D - E 1:� 1% 
J acre flat - 18 acre hill E - F approaches 5% 5� 
18 acre hill - 1 acre flat F - G 1% 
1 acre flat - 18 acre hill G - H  1� 
18 acre hill - 49 acre hill H - I  5� 
49 acre h1ll - 18 acre hill I - J 
• For the identification of the locations and periods referred to in this table see Fig. III � 
166 
signi£1cant effect on mileages could be found for dif­
ferences in pasture size within the same pasture location 
(hill or flat ) .  
(9 )  The frequenc.y with which identical or aL�ost identical 
mileage figures occur with individual sheep for conte cutive 
weeks (Table 19 ) .  
( 10 )  Admittedly , weekly mileages may not be an entirely satis� 
f a c t o r,y  way of examining the effect o£ weather en distances 
grazed , but except £or one period, 24/11-1/12/56 , no 
regular relationshi1) a;,n be traced between weekly climate 
and the distances traveled. However, the weather records 
do give a possible explanation why the Ch�viot group 
mileage fell below that of the Romneys in the week 24/11-
1/12/56 (Figure III ) .  The week in question saw a drop of 
10 to 14 degrees in mean temperatures , almost lOO percent 
more wind and half an inch of rain spread o·var five rainy 
days . The recently shorn Cheviots would appear to have 
taken badly to these weather conditions as their mileage 
dropped severely despite the fact of their transf r to 
lowland country. It is interestin� to speoulate whether 
or not the effeet o£ weather conditions on the Romney was 
neg ted by their favourable re etion to lowland pa turage-­
their natural environment. 
(11) Grazing distances did not increas during lactation 
although they did tend to rise during the latter stages 
of pregnana,y in 1957. 
167 
Comparison of the results with Tribe ' s review is of some interest, 
for they tend to contradict some of the ideas put forward by this 
author regarding the influence of pasture size , weather and lactation 
on th� activity of grazing animS:. s . 
An�ysis of the leg measurement records (Tables 22 and 23 ) 
show·ed that the Romneys were on the average three quarters of an 
inch greater in h-eight at the withers , half an inch greater in 
height to the brisket and approximately one to two tenths of an 
inch greater in elbow-coronet measurement than the Cheviots . 
T his was , of course ,  to be expected, �for the Cheviot is , generally 
speaking, a smaller sheep than the Romney although the measurements 
do not altogether agree with those of von Borstel (1951) taken on 
ten a.ni.mals of each br ed (Table 2) ) . von Borstal found his unshorn 
Cheviots to be about one quarter of an inch higher at the withers 
than the un horn Romneys and one twentieth of an inch higher when 
both breeds were shorn. 
It would nevertheless appear that the greater distances traveled 
by th Cheviots (Figure III: table 19) cannot be related to a greater 
leg length (Table 22 ) .  
Table 22. Compa.riscm of foreleg mea�ements,  lfaematology and 
Rangemeter sheep. 
Cheyj.Ot! 
B!atolm Flock 
Height at withers 22.S" 
Height to brisket 11.4• 
Elbow to coronet l.J,)" 
Rgnnve 
Height at withers 2).2" 
Height to brisket 11.9" 
Elbow to eoronet 1) .5" 
§t. p. 
0 .9• 
0.9" 
o.s" 
Bfngemeter Sheep 
&w1 St. ]), 
2). 0" 0 .6• 
11.9" 0 .2• 
1).4• 0 . 2" 
12.5" 
Table 2).  Comparison of height at withers measurements (unshorn sheep) 
Yon Borstal (19.5'1 )  and HaematolOgy flock this project . 
M an het.ght 
at witheri 
NUIIlber of sheep 
Qb!Yiqts Bomn!ll 
JOD Boretel This froject t.QD jprsttl Tbit Project 
22.77" 
10 J7 10 37 
DIVIS ION " D " 
HAE� TALOG IC.A L S T UDIES 
170 
CONTENTS 
Page 
D-1 Introduction and review of literature 171 
D-II Standards 173 
D-III Techniques 175 
D-IV Experimental arrangements and procedures 180 
D-V Results 184 
D-I Il�TRODUCTION AND RE IEW OF LITERATURE 
The aim of this Division of the project was to carry out 
comparative haematological studies with Romney Marsh and Cheviot 
sheep. 
171 
A survey of the literature revealed that haematological studies 
have been little utilised as a means of pinpointing or explaining 
differences between breeds of sheep even though Kerr {19)7 ) ,  working 
with about twenty species of vertebrates , observed that the most 
strikin feature revealed by the investigation was the wide variation 
in the blood composition of various sheep. Kerr found that although 
indi.vicluals of the same species as a rule have similar chemical 
composition of erythrocytes , enormous differences were found among 
sheep in their content of potassium. For fat tailed sheep the high 
values were about 64.2 m .  equiv. 1 .  and the low values about 18.4 
m .  equiv. 1.  
Kushner and Kitaieva (19)8) investigated the physiological 
differences which distinguished hybrids fr�, local Kirghiz and 
Merino sheep. They found that the blood indices , haemoglobin, 
erythrocyte count and alkali reserve of the hybrids in all age and 
sex groups were consistently higher than for local fat tailed sheep 
and the Merinos of the New Caucasian type . Karpov {1941) showed 
that crosses of the Romney Marsh and the Tsigai were higher than 
their parent breeds in regard to ha oglobin• number of erythroa,ytes , 
total surface area of the erythrocytes and functional capacity of 
the ha oglobin. 
172 
Mcl.eroy � al. (19.52) endeavoured to ascertain some of the 
pnysiological processes involved in the greater heat tolerance of 
Rambouillet sheep as contrasted with that of the Suffolk under the 
extreme heat of Central Arizona.· The pa.cked red cell volume ( P.C .V . )  
of eight ewes and their lambs for each breed was investig&ted. The 
breed differences were not statistically significant. 
Todd et al. ( 19.53) comment on the fact that information on the 
cellular elements of sheep' s blood largezy rests on a report by 
Holman (1944) on Scottish sheep. In reporting a comparative haemato­
logical study of .5.5 healt� pure-bred Southdown and 3.5 pure-bred 
Hampshire ewes, all within two months of lambing , they attached no 
significance to aQY of the differences between the breed means. No 
statistical ana:l.Jrsis is reported. 
Evans (19.54) found approximately 50 percent of 93 Scotch 
Blackfaced sheep belonged to a type with a high concentration of 
potassium in the whole blood and red blood cells . Only 16 percent 
of 43 Cheviot sheep belonged to the H .K. type. 
Evans and Mounib ( 19.57)  report a survey of the potassium con­
centration in the red blood cells of British breeds of sheep. They 
pres nt data as follows (Table 24) showing that the mountain and 
hill breeds on the avera contain a greater percentage of high 
potassium animals than do the lowland breeds. 
Table 24. The distrlbut.ion . of high potassium values in the red 
blood cells of British breeds of sheep (Evans and 
Mounib 19.57 ) .  
Sheep subtyp 
Black!ace Mountain 
Other hill breeds 
Lowland Longwools 
Lowland Shortwools 
Percentage H.K.  type 
51 
37 
11 
13 
173 
These authors find it difficult to assess the significance of 
their findings but. believe that the predominance of high potassium 
animals living in less favourable climates suggests that. this factor 
may have soc;•r. '1:- e:. a riug on survival in mountain and upland regions in 
the temperate zone . The br eds can be classified on whole blood 
estimations . 
Gregerson (1951 ) reviews the subject of blood volume and states 
that normal values have been reported on a number of the common 
laboratory animals, but beyond this little has been done in the 
comparative physiology of blood volume. 
No literatur could be located which reported blood volume 
studies on a comparative basis between sheep breeds. 
D-II STANDARDS 
To facilitate comparison, table 25 presents a number of sheep 
' bl 2,5. tolo cal • 'lN tor ame.p. 
Jo land (193)) 
il r ( 19)3) 
Hamer (19 • )  
Bennett and Ch pun ( 19)7 ) 
X:err ( 1937) 
· ston !i • (19)8) 
Und rwo ...1 !!· ( 19)9) 
U1erott (1941 ) 
Karpov ( 1941) • 
•• 
Ho lllan ( 1944) 
(194.5) 
An m1 
Mild 
od.er te 
Ho an ( 1941) 
a (194? a) 
d tltout (19,1) 
• (19SJ)+ 
++ 
( 195)) 
Me • (19:54) 
8 (1954) 
� 8 (19.54) 
Hb. P . O • •  f\ . B . C . 
lo.4 ! o.?s 10 • .5 � .97 
15.0 
10.0 - 21.0 
u.o  - 12.0 u.o 
10.0 - l.S.O 
10. 8 
9 • .5 - 1).4 
9.7 10.0  z 0.)1 
9.9 10.1.5 ! 0 .2) 
8.0 - 10.? 26 - )8 6 - u 
u . .s 
9·? 23 ... 1? 
?. s 17 ... u 
12.4 + 1.4 - 30.5,! 3.5 n.s 1 1.s 
11.18 8.1 
.6 
13.41 ! .1.6 .tto.)8 : o. u.6s 1 o.21 
12. ! 0.19 .)S 1 0.66 12.47 1 o .,2 
l.S.l5 (Sierra ot Peru h t bt 11000' • lSO 
?.0  • 
• 6 9.S 
. •  per 100 • 
cooytea (thou 
M . C . l • 
++ ,. 
w • •  c . 
?.0 ! 1.15 
10 .9? !. 0.61 
6.91 !  0.)6 
s ... 14 
9.2 1 ).1 
?.lt4 
8.0 
?. !. 0.26 
7.74 + ·" 
t ) 
- -
Pot.ua1 
M . c . v . M . C . fl . M . C . • cone. • equi •./L r; 
Alsaad aau�a 
gh Lov 
64.2 18.4 con•er-
· ion· 
34 - .50  12 - 20  
27 41 
21·4 ± 3 . 6  41, 2 + 4.0 -
)2.2) ! 0.44 
)1.07 .: o • .so 
.s 
�.1 
. o 16.8 
9.0 2 .o 
17.5 
blood values reported by various workers . 
Cited from Ransard � 2!• (19.53 ) table 26 gives a summary of 
their findings for sheep blood volumes together with those in the 
literature at that time. To these must be added , the results of 
Barcrort (1939 ) and Sehambye (19.52 ) .  The fonuer reported the average 
blood volume of three hysterectomised sheep as being 60 ml . /kg. The 
latter author obtained mean values of 2.94 litres per animal and 
60 ml.. /kg. body weight for six sheep. 
D-III TECHNIQUES 
Estimations of haemoglobins , packed red cell volumes , cell 
counts and potassium values are very frequently carried out in 
medical , veterinary and agricultural investigations ; and the 
techniques are well standardised. In this project, cell counts 
were carried out according to the methods of Dacie (1951 ) ,  potas­
sium values w r estimated using an EEL name photometer and haemo­
globins were done on a Hilger Biochem-Absorptiometer .  However, 
blood volume estimations are not so commonly attempted , at least 
in Agricultural research : nor are the techniques standardised. 
In measuring blood volumes early investigators used the Welcker 
exsanguination method {Wiggers 1949a) .  This t chnique depends on 
bleeding the animals to death and measuring the volume of blood so 
obtained. The blood remaining in the animal is washed out bv po•t­
mortem perfusion and its volume caleulat.ed ·from the comparative 
Table 26. Summarising data in the literature on blood volumes of sheep (Hansard et al - 1953 ) .  
No.  of Average ml. Blood 
Animals Age Weight lOO gms. St . E . Ref .  
lbs . Body Weight 
-
4 1 day 7 16.9 - Gotsev 1939 
4 3-4 days 16 10.9  - ,, " 
8.0 - Dukes 1947 b 
5 4 months 27 8.0  0.2  Hansard 
et a1 1953 
4 6 11 92 6.3  0 .1 u " 
4 1 year 13.5 5 • .5 0 .3  " tl 
4 3 years 156 .5 .8  0.3 If 11 
3 7 years 11.5 8.0 0 .4 " If 
·-
..., -.,J 0\ 
177 
haemoglo'bin content. of the normal blood and of the washin s .  
Today. most determinations are made by measuring the dilution 
of a known quantity of material which is added and which remains 
confined to the blood stream. One of the commonest substances us ed  
is th dye Evans Blue (T-1824) .  This dye combines with plasma 
albumen in vertebrates and leaves the blood very slowly. Carbon 
monoxide and red cells containing radio-active iron or phosphorus, 
or protein with radio-iodine have also been used in determining 
blood volume. 
There is little information in the literPture on the minute 
details of estimation procedures for the blood volumes of farm 
animals . 
Turner and Herman (1931) state that the dye injection method> 
as with all blood volume methods , has been the subject of some criticiam. 
The proper time after injection for the drawing of the samples and 
the rate of disappearance of the dye from the blood stream immediately 
following the injection are most perplexing problems . The e workers 
eventually arrive at the conclusion that four to eight minutes would 
se to be the optimal time for withdrawing the samples after injection 
of the dye in the bovine . Miller (1932) using vital red dye estimated 
the blood volumes of 19 mature cattle several times each during an 
18 month period. The quantity of dye he used was 1 ml .  of a 1.5� 
aqueous solution for each ten pounds of body weight. The sampl 
of blood was taken from the opposite jugular vein ten minutes af"ter 
178 
the administration · o:f the dye . On the basis o:f 81 deterruinations 
the average quantity of blood per pound· of weight was 27 . 07 ml .  or 
6 .7  percent o:f b� weight . The results showed a considerable 
variation , i . e .  22�33 ml. /lb. but 75 percent o:f the values were in 
the narrower range 25-30 ml. lb. During pregnancy there was a 
marked increase in blood volume proportional to the weight increase 
until parturition occurred. Shortly afterwards <, .ere was a decrease 
in both blood volume P�d body weight . 
Barcro.f't .!i 1!1• (1939) give the blood volumes :for three hyster­
ectomised sheep • The technique appears not to have included a starv­
ation period and the sheep were transported singly about two miles 
to the laboratory anything :from half an hour to one day before the 
estLmation . E v an s  blue (T-1824) was used. 
Gotsev (1939 ) estimated the blood volumes o:f 5 lambs at birth 
by the injection o:f a 2 percent solution o:f Congo red dye. T h i s  
was injected into the jugular which was exposed by a skin incision 
under a local anaesthetic.  
Courtioe (1943 )  using Evans Blue (T-1824) on rabbits• dogs 
(including greyhounds ) ,  goats and horses , at about 0•8  mg . dye 
p r K • body weight first drew a ample o:f blood to act as a stand­
ard• The dye was then inject d into a jugular vein, a further sample 
being withdrawn from the opposite vein arter six minutes • The 
concentration of the dye in the pla was t.h n d termined by means 
of the ha tocrit spun for 45 minutes at 3000 r.p.m• A correction 
179 
f a ctor of 4 percent for the qye trapped between the corpuscles 
was u s e d  after the figure put forward by Gregerson and Sehiro (1938) .  
Reeve (1948) deals with the difficulties of the indirect 
method of blood volume determination as follows :- A sample may be 
cloudy "When drawn, or cloudiness may develop or increase sub­
seq<lently. The dye content of all plasma samples should be estimat d 
as soon as possible after withdrawal, preferably within two or three 
hours . and the temperature of the sample is best kept above 20°C. 
On standing , fat may separ�te from the plasma and float to the top ; 
and this process becomes more rapid as the temperature falls . �ben 
!at ri s e s  to the top either the plasma beneath may be removed 
uncontaminated by very slow pipetting or the superficial fat layer 
may be held in an added surface layer of mineral oil. Plasma 
cloudiness is reduced to a minimum b,y the starving of man or animal 
for at least twelve hours before the experiment. 
Prosser _1 �. (1950a) say that the various methods for estimating 
blood volumes give somewhat diff rent values , but there is a strik• 
ing gener�1 agreement. 
Schambye (1952 ) used both Evans Blue (T .. l824) and p34 tagg d 
erythrocytes , and Hansard ,!! !!• (1953 ) used a phosphoru labelled 
r d cell method for blood volume estimatione . 
The proc dures for blood volume estimations in this project 
were adopted after a con ide ation o£ the aforementioned technique • 
Evans Blue dye {T-1824) was the agent used. Possible complications 
180 
due to pregnancy, lactation and o strus were avoided by carrying 
out the operation when the sheep were approximately half-way between 
weaning and tupping, i . e .  w·eaning 12/12/56, t.upping 27/4/57 , blood 
volume estimations 19-20/2/57. 
D-IV EXPEP.IMENT.AL ARRANGEMEN'XS AND PROCEDURE 
A flock of 35 of each of Cheviot and Romney Marsh ewes was 
established at Tua Paka , one of Massey Agricultural College ' s h� 
farms . Both groups of sheep originated in similar hill country and 
consisted of mature ewes of equal ages in-lamb to the Southdown• 
During the entire period of the investigatio� this flock was maintained 
as a single unit on the same hill paddock . Age, growth and environ­
mental variations were therefore minimised as far as was possible . 
Hewi tt (1947) describes this area as second and third class � 
country of an easy nature and running up . to approximately 700 feet . 
The pasture is mostly browntop-dominant , capable of carrying one ewe 
to the acre at the most . 
The ewes were weighed at appropriate intervals 1  and account 
was taken of lambing records , lamb growth and wool production. 
Total worm egg counts carried out in July/56 indicated that the 
ewes were not heavily infested with worms . I� is recognised that 
low worm egg counts do not necessarily mean a low worm burden, but 
at no time during the project were aQY obvious symptoms of parasitic 
infestation shown. 
181 
The first •bleedi ngs ' took place on 7.S/1/S6, the Romneys 
being dealt with o n  the first day. More than 35 of each breed 
were included in the first bleedings.. '!'h i s  was because several 
of each breed were pr(Wided as spares for the over-all project , and 
it was decided to utU1se them in the establishing '3f the breed 
means for the haematologieal values il'l"'estig ted . 
Second and third bleedings took plaee in early Oetober/S6 
and mid�ebruary/S? . 
Ql all occasions bleeding t ook place 1n the late afternoon. 
The samples drawn frOl" the jugular v.er� eJCelftined 1.n the Haematology 
Department of the laboratory, Palmerston North Pub lie Hospital. • 
The mid-February operatian was different to the previous two 
as at this time procedures for blooa volume estimations were carried 
out 1n addition to routine blood sampling.  
At this time location of the jugular va facilit ted by shaving 
the necks of the sheep (Plate 42) . Also at tbis time the animals were 
hou ed 1n the evening before each of the bleeding dqs. This ga1'e 
a starvation period of approxim tel,y 24 hours , as the fir t d7e was 
not injected untU 6 t00 p.m. on the bleeding day. 
• Jfo ba .... tologtoal laboratory being situated at hssey Agricultural 
College, the Medical Superintendent, the Senio-r Pathologist and the 
Principal B cter1olog1st of tbi hospital generously offered the �·• 
of their .f'ao111tteo for llm1ted oount of wcwk. All equipaent was 
eal1brated and controlled by the Principal Bacteriol. 1 st  ot the 
Pa�eraton North Pltbl.lc Ro p1tal. 
Plate 42 . Blood dye injection . Note the shaven 
neck . 
182 
/" 
183 
Being too great a number to deal with on one night, the flock 
was divided into two groups , each. group consisting of an equal 
number of each breed. These groups were then dealt with in sub· 
groups of seven , alternating between the breeds . 
Th e  operation proceeded very smoothly on both nights . No 
difficulty was encQuntered in locating and entering the jugular 
because of tha close shav1ung of the animals ' necks . An initial 
sample of blood was drawn from the first Cheviot and Romney on each 
night, these samples being thereafter utilised as a standard. 
There was no breed difference in the opacity of the plasma samples . 
20 ml .  of 0.9 percent NaCl containing 15 mg . of Evans Blue (T-1824) 
was then injected into the right jugular (Plate 42) .  This is the 
concentration of dye recommended by Mollinson (1951) and is that 
used in routine blood volume estimations for hwnans from 100-200 
lbs . weight at the Palmerston North Public Hospital . Eight minutes 
later a sample of blood was withdrawn from the le.t't jugular into 
a heparinised bottle. Timing was arranged ro that the samples were 
centrifuged within two hours of the first injection of dye . The 
plasma dilution of the dye was then calculated using a VIC photo­
electric colorimeter. Packed red cell volumes were obtained and the 
total blood volumes etc . arri v d at by using a correction factor of 
4 percent (Gregerson and Schiro 19)8) for the dye trapped between 
the corpuscles . 
Owing to a miscalculation of the amount of dye available , 
the final score was Cheviots 27, Romneys 24. 
\ 
D-V RESULTS 
It was impossible to set up the three bleeding dates as a 
two-way table , there being numbers of differing animals missing 
1� 
from the second and third stages of the work. Statistical analysis 
was therefore restricted to analyses of variance between breeds and 
between dates within breeds (Appendix B, tables 1 to 5 inclusive) .  
Table 27 clearly demonstrates that the Cheviot ewes • started 
heavier and maintain d their weight lead over the Romneys throughout 
the project. The Cheviots lambing percentage (Table 27 ) was greater 
than the Romneys (140 rll5� ) .  Only three Cheviots and two Romneys 
lost their lambs . The rate of gain of the Cheviots ' lambs (Table 
27 ) was significantly superior to the Romneys when expressed in 
pounds of lamb per ewe between docking ( 27/9/56) and weaning (12/ 
12/56) ,  although the difference in the weaning weights of the lambs 
of the two breeds was not statistically significant (Table 27 ) .  
Romney wool production was 50 percent greater than that of the 
Cheviot (Table 27 ) .  
Comparison of the haematological data obtained in this inv stigation 
(Table 28) with those in table 25 indicates that there were no 
abnormalities 1n the general haematological picture of the flock. 
It will be seen from table 28 that in arly July/56 there w highly 
significant difference between the haemoglobin& , packed r d cell 
volumes {P. c.v . • ) and leucocyte counts for the two br eds , the 
Table 27. The general production data o.f the Haematology fiock . 
Factor Date Cheviot 
mean St. D . 
Ewe weight lbs . 6/7/56 128. 0  8. 8 
(Pregnant) 
Ewe weight lbs . 27/9/56 117 .5 9 .8  
Ewe weight lbs . 21/2/57 111.2  9 .4 
Ewe weight lbs . 30/7/57 125.4 10.9  
Lamb production lbs . 27/9/56 49 .1 14.7  
to 
12/12/56 
Weaning weights 12/12/56 74.6 20 .8  
of lambs lbs . 
Wool production lbs . 22/11/56 5.25  l . OJ 
Lambinv 14� 
Romnes 
mean t.  D .  
105 .3 8.4 
111.2  10.7  
10) .6  11.1 
120 .1 11 .6  
41 .6 8.5 
66.2 ll .J 
8.10 1 .27 
115� 
Level of significance 
of difference between 
breeds 
1� 
5� 
1� 
approaches significance 
5'1t 
1� 
� ()) \.1\ 
Table 28. Hae��tological data of the Haematology flock and the results of analyses or var1ance 
carried out on the same data . 
Levels of sig . of diff. 
Cheviots Romneys between between between 
Date breeds following Jul.y/56 
n.  mean St. :C n.  mean St . D dates Feb./  57 
c R. c .  R .  
Haemoglobin 
9/ 7/56 (Hb. ) *  42 13 .45 o . ao 43 12 • .58 0 .96 1% 
3/10/56 33 11 .84 1.10 30 ll.8l. 1.13 1" 1:' 
19/ 2/.57 30 13 .37 1•43 29 12 �27 1.80 1� ·s?h 5% 
9/ 7/56 
Leucocytes 
(I>J.B . C. ) *  41 8.4o 2 . 20 42 ll .09 2 .50 1� 
3/10/56 32 8.75 2 .30 30 ll . 24 2.44 1% 
19/ 2/.57 30 9 .42 2 .05 29 10.98 2.56 app.l% app • .5% 
Packed red cell 
volume 
9/ 7/56 (P. C.V . ) *  42 4.5 .3 3 .6 40 42.2 4 .4  l;:b 
3/10/.56 30 37 .6 4.1 30 37.4 4 . 0 1'% 1% 
19/ 2/57 30 1+2 . 0 6 . 8 29 38 .8 .5 . 2 5� app • .5'f/, 1� 5% 
Erythrocytes 
9/ 7/.56 (R.B . C. )* 8 13 .16 1 .58 8 11 .46 1 .04 5% 
3/10/56 13 10 .46 1 • .50 1.5 10 .72 1.39 1� 
19/ 2/57 14 ll .18 1.15 13 10 • .50 1.t8 1� 
!4 . C. V . *  
9/ 7/.56 8 34.6 4.3 8 36.5 4.4 
3/10/.56 12 36 .5 7 . 0  15 35.3 3 .4 
19/ 2/57 14 37.6 4.9 12 36.7 6.2 
-1' . C.H. * 
9/ 7/56 8 10 .4 1.2 8 ll .2 1.1 
J/10/.56 12 ll.6 2.3 15 ll.O  0.8  
19/ 2/57 14 12 .0 1 .6 12 ll.8 1.9 .5% 
n. C.H. cone . *  
9/ 7/56 42 29.8 2 .3 40 30.0 2.LJ. 
J/10/56 30 31.4 1 .8 28 31.5 1 .2  lJb J$ 
19/ 2/57 29 31.7 3 .1 29 32.1 5 ·2 1� 5% 
* See table 2.5 for definitions 
18? 
Cheviot being higher in the first two and lower for the last factor. 
Erythrocytes were significantly higher in the Cheviots {at the five 
percent level) but no significant breed differences existed for 
those indices of normality , the mean cell volumes (M.C .V . ) , mean 
cell haemoglobin (M. C .H . ) , and mean cell haemoglobin concentrations 
(M . C.H . cone . ) .  Just how much these figures were influenced by 
pregnancy and the recently differing locations of the breeds {brought 
together in early June/56 ) it was impossible to assess . However, 
in October/56 the flock had existed as a single unit for upwards 
of four months . 
At that time leucocyte counts were the only significant (very) 
difference between the breeds, these counts being very close to their 
July values . Haemoglobin and P. C.V. ' s  in both breeds , and erythrocyte 
count in the Cheviots , had fallen very significantly . There was 
virtually no difference between the breed means for these iterns . 
M . C.V . and M . C . H . were again not significantly different and corres­
ponded closely with the July values . There was no significant 
difference between the breed means for M . C. H. cone. but both values 
had risen very significantly from their July level . 
In February/5? leucocytes and P. c.v . • s  were the only two factors 
showing a significant breed difference . Haemoglobin levels had 
risen very significantly and significantly in the Cheviots and 
Romneys respectively since October/ 56. In the Cheviots the packed 
red cell volumes had risen almost significan�y. 
' 
On a comparison being made between the July/56 and February/57 
values it was found that in February the Cheviots had recovered 
to their July mean for Haemoglobin, but the Romneys were still 
significantly lower than their July level . A slightly larger 
difference between the breed means for haemoblogin in February 
(as compared with the July figures ) was not significant, possibly 
due to a higher variability within the groups . 
The July/56 and Februar,y/57 Romney leucoe,yte counts were 
similar,  but those of the Cheviots had risen almost significantly. 
Despite this rise , the Cheviot means for the three bleedings lay 
below 10, 000, and all of the Romney means lay above 10 , 000 . P . c.v. • s  
suffered very significant and significant falls in the Cheviots 
and Romneys respectively between July/56 and February/57 and er,yth� 
rocytes were very significantly lower in the Cheviots in February/57. 
The M. C .V. showed no change between the two dates , but the M. C.H . 
for the Cheviots was significantly higher in February/57 than July/ 
56. This latter was due to the fall in the Cheviot erythrocyte 
counts . The i� .C .H. cone . showed very significant and significant 
rises between the July and February records in the Cheviots and 
Romneys respectively. 
Although both sets of figures lay within the normal limits 
for mammals , of seven to ten percent of the body weight (Dukes 
1947 b; Wiggers 1949 b ;  Prosser � !!· 1950 b)  the results for the 
blood volume estimations (Table 29) revealed a very marked difference 
between the br eds . The Cheviots had an average of 1.06 litres ore 
blood p r she p than the Romneys . This difference remained statis-
Table 29. Blood volume and relevant data-Haematology flock, Febru.ary/57. 
Cheviots Romneys Level of significance 
Factor 1-!ean St . D Mean St. D of breed di.f.ferences 
Total blood volume 
in litres 4.64 0 .46 3 .58 0 .?2 1� 
Total plasma volume 
in litres 2 .?6 0 .28 2 .26 0 .39 1� 
Total erythrocytes vol. 
in litres 1.88 0 .12 1.32 0 .14 1% 
Body weight Kg .• 49.8  4.3 46.1 4.9 1� 
Blood ml./Kg. 93 .2  9.2 77 .7 14 .1 1:' 
Plasma al .  /Kg. 55 .6 6.9 49.3 8.5 1' 
Erythrocytes ml. .  /Kg. 37.5 6.9  28.4 9 .2  1� 
Average sheep boqy 
surf'ace area (sq. metres 
computed)* l.ll 0 .05 1 . 06 o .o6 � 
Blood litres/s�. metres 
surface area 4.17 0 .38 3.36 0 . 62 1� 
Plasma 11 tres/ sq . metres 
1% surface area 2.48 0 .26 2.1) 0 .35 
Erythrocytes 
11 tres/ sq. metres 
surface area 1.68 0 .32 1.27 o .43 1� 
• Brody (1945) 
190 
tically very significant when plotted against Kg. body weight* 
and sq. metres computed surface area. * The mean difference 
in blood volume is composed of approximately equal parts of plasma 
and erythrocytes . 
Estimations of whole blood and red blood corpuscle potassium 
(Table JO )  showed that approximately three fifths of the Cheviots 
and one fifth of the Romneys belonged to a high potassium type.+ 
Th@ mean values for whole blood and red blood cell potassium 
in the H.K.  and L.K. types agr ed very well with those obtained by 
Evans (1954) . However,  the distribution of the values for the 
Cheviot is different to Evans ' findings . He found only 16 percent 
of Ch viots belonged to the H.K. type. 
• The sheep were weighed the day aft r the blood volUIJle stimations 
w re complet d , i .e .  on 21/2/57 . 
+ As defined by Evans, 1954. 
Table )0. The mean concentrations of Potassium in the whole blood and red blood cells . 
Breed 
Cheviot 
Romne;y 
Scotch 
Blackfaced* 
Cheviot 
Romney 
Scotch 
Black£aced* 
HaEmatology fiock J/10/5"6, also data ex Evans• (1954 ) .  
n. 
Whole Blood 
17 
5 
47 
Red Blood Cells 
17 
5 
47 
High K .  values 
(m. equiv. /1itres ) 
mean St . D 
Jl.02 J .J9 
)0.34 6.)5 
)6.00 
77.0) 12 .05 
78.82 4.6) 
89. 00 
n. 
1J 
21 
46 
9 
21 
24 
Low K . values 
(m.equiv. /litre ) 
mean St. D 
10 .48 4.4) 
10.27 1 .70 
15.00 
21.14 12.98 
21.6) 4.6) 
24.00 
• These figures are the high and low mean values put forward by Evans (1954) for Scotch Black-
faced sheep. 
� 
DIVISION "E" 
DISCUSSION AND CONCLUSIONS 
OF THE OVERALL PROJE CI' 
(Divisions "A" , "B" , " C" ,  "D" )  
193 
DISCUSSION 
This investigation found its roots in the fact that 1n New Zealand 
the Romney Marsh sheep is being asked to live and perform on certain poor 
hill country on which it is giving disappointing results (Peren !!.!l• 
1951 )  whereas the Cheviot (and the cross between the two breeds ) 
thrives well under the same conditions . 
An examination of the Unit d Kin dam backgrounds of the breeds 
confirmed that the Romney is historically a sheep of fat lowland 
pastures and comparatively clement climatic conditions, �le the 
Cheviot was developed as a sheep for a hill region noted for its 
thin pasturage ·and severe climate.  
These backgrounds suggested three lines of investigation which 
would possibly pinpoint differing breed characteristics acquired in 
Great Britain and now related to the comparative success and failure 
of the Cheviot and Romney respectively in the particular New Zealand 
conditions . The studies taken up were Energy Metabolism, Ranging 
Behav1Pur· andHaematology. 
For en rgy metabolism studies (Di. vision "B') a new technique 1n 
closed circu1. t indirect calorimetry was vol v d .  This enabled 
com.parati veJ y long tenn energy metabolism exp riments to be carried 
out ; and was also used to demonstrate its usefulness tor otheJ· ·'-JP88 
ot study; ollmatological, physiological and nutritional. 
So far as the writer is awar , thi is the first inter-breed 
metabolism study with sheep set up as such both 1n its conception 
194 
and experimental plan. 
Interpretation of the results of this division of the project 
is complicated by the introduction of new animals and the fact that 
the Romneys (except tor the first block of energy metabolism 
experiments ) tended to be heavier than the Cheviots . It can thus 
be argued that for future work every effort should be made to 
maintain equal weights between the experimental groups and so avoid 
the necessity for mathematical equating . However , the policy 
adopted would depend upon the hypothesis being investigated, i .e .  
is  there a differential between the breeds at typical breed weights 
or between members of different breeds which happen to be , as 
individuals , at identical weights . 
Analysis of the records obtained over an eight to ten month 
period indicates that the Cheviot--a true hill breed-has a lower 
metabolic rate than the Romney--a true lowland sheep . The Cheviots 
(per animal) normally metabolised at a level 10-15 percent below the 
Romneys . However, when the oxygen consumption records were related 
to kilograms body weight , sq. metres surface area (computed) and 
kg . body weig�t to the power 0 .73 , the differences between the 
br eds wer not statistically significant, except under severe 
starvation condition for the latter two bases of presentation . 
When starved the Cheviots rapidly adjusted their metabolic 
rate in an orderly manner to a low level, while the Romneys appeared 
to be more erratic in their response to food shortage . 
195 
This reaction of the Cheviot to starvation is considered to be 
support for the hypothesis on which the energy metabolism division 
of the overall project was based , namely that the Cheviot in New 
Zealand should possess physiological mechanisms related to the 
ability of its forebears to survive on the barren and often snow 
bound pastures of the Cheviot hills ; physiological mechanisms which 
the fat-land Romney need not have developed . 
Some basis was found for doubting the validity of the presently 
used mathematical techniques designed specifically for making 
homeothe.rms of varying sizes fit into a general metabolism law. 
This finding was that the metabolism of both Cheviots and Romneys 
fell slowly but quite consistently from lambing time to the autumn 
despite the tendency for both groups to be maintaining or even 
increasing weight . Greater body weight is supposedly associated 
with greater oxygen consumption per animal. The fall in energy 
metabolism as autumn approached would appear to follow the seasonal 
rhythm in thyroid activity (Brody 1945a) .  This trend was unbroken 
despit the introduction of new animals (four Romneys , three Cheviots ) 
into each group during the experimental period. 
Carcass analysis carried out on experimental animals indicated 
that the Cheviot dr ssed out better then the heavier Romney and that 
the heart-body weight ratio may be higher 1n the Cheviot. 
Although th se differences were not statistically significant, 
they are interesting when considered from the vi w-point of relating 
196 
body weight or some portion of the body weight to oxygen consumption. 
If further work were to confirm that the Cheviot is a more 
muscular animal , has a higher dressing percentage , a greater heart 
weight and/or greater blood volume then it would be quite feasible 
to rate this breed with a higher physiologically effective body 
weight than the Romney. vlork is being undertaken at l-1assey Agricul.-
tural College on the ohemical and dissection analysis of the 
dressed carcasses . The indications are that the Cheviots wer� in 
fact , the more muscular group. This work is to be published inde-
pendently. 
In view of the above , the writer contends that no justification 
can be seen for the present methods of equating races within a 
species by mathematical techniques that are based on an assumption 
of constant ratios of weight and function• between the various parts 
of their bodies . Such methods may be very well for general studies 
on a comparative basis which show that homeotherms all obey basically 
the same laws , but it is certainly not a good enough approach for 
critical intraspecific studies . 
Support for this contention is found in Hellb�rgs argument (1949) 
that , so far as is possible , variations in body weight ought to be 
analysed with reference to its causes and that the different it s 
ought to be treated diff rently in the way of correction. 
• Behavior and haematological studies with Cheviot and Romney Marsh 
sheep in this project indicat d that there were large breed dif­
ferences in activity , blood factors and blood volume . 
197 
w'hen the o;,cygen consumption data were expressed as Calol"ies/24 
hours , the mean breed figures were in general higher than those in 
the literature . That the sheep in this project were standing and 
on full feed may be implicated , as well as the error of conversion; 
however, the fact of large seasonal and breed variations may indicate 
a need for a revision or many of the figures put forward for the 
nutrition requirements of sheep. 
As an Addendum to the energy metabolism studies , five incidental 
reports are presented :-
(a)  The ef'f'ect of shearing and drenching with simulated rain 
on the o�en consumption and respirator,y pattern of sheep. 
(b)  The effect on their metabolic rate of feeding 1 lb. of 
oats to each of three sheep after twelve hours 
starvation. 
( c )  The effect of 1 thyroxine on the o:x;ygen consumption, 
respiration rate, and body weight of one Romney Marsh ewe. 
(d)  Methane exhalation in sheep. 
(e)  The regurgitation process in sheep . 
These reports are self contained, but taken as a whole do indicate 
that there is a wide field or work to which the devised techniques 
can be adapted. 
It being also hypothesised that the Cheviot should be a more 
active animal than the Romney, an entirely new device-the Rang ter-
was developed to enable weekly grazing mileages to be recorded under 
completely normal conditions on all types of pasture 1n any location 
(Division " C" ) .  
The records show conclusively that of the animals under investi­
gation the Cheviot , the shorter legged breed, was 50 to lOO percent 
more active than the Romney under hill conditions . The Cheviots 
also responded to wider spaces on the hill . Under lowland conditions , 
however, the Romney increased its distance traveled an� in fact, 
very nearly equalled the Cheviot, only to fall back again quite 
decisively when returned to the hill . The fact that the Romney 
greatly increases its activity on flat lowland pastures contradicts 
the commonly held idea that the Romney Marsh sheep 1n New Zealand 
has adapted itself to hill grazing . 
The rangemeter was enlightening in other ways , for it allowed 
the first long term records derived from a number of sheep to be 
critically compared with the ideas put forward in recent publications 
on animal behaviour (Hammond 1955 ) .  Contrary to these ideas were 
the findings that :-
(1)  Within the breeds under lowland conditions , size of 
field had little or no effect on the distances traveled and that 
this applied to the Romney on the hill ; however , the Cheviots • 
increase in grazing distance in a change from an 18 to a 49 acre 
hill pasture may have been due to a pre-ordained grazing routine 
rather than nutritional necessity. 
(2 )  Weekly weather as such had no r&gular effect on the weekly 
distances traveled, although just after shearing there y have been 
a distinct tendency for the sheep to r uce activity when it rained. 
199 
The moving of the sheep to a lowland pasture at trds time complicates 
interpretation of the results . 
(J )  Lactation did not appear to result in increased grazing 
distances . Howev r ,  in 1957 when the records were much more complete , 
there was a rise in the activity of the pregnant animals as they 
approached parturition. This rather contradicts the shepherds ' 
traditional idea that ewes tend to spend more time lying down as 
lambing approache • 
Four other interesting findings were : 
(1 ) The frequency with which almost identical records turned 
up for individual sheep in consecutive weeks . rhis may possibly 
be due to the particular animal establishing a grazing beat for 
itself. 
In this respect further observational work in combination with 
the use of the rangemeter may vindicate the belief of many shepherds 
that sheep establish a :> o:--:: :: ld.thin a paddock (Firbanl.< 1940 ) .  Such 
work could also help to explain the working of hefted sheep on 
unfenced hills in Great Britain. 
(2 ) The way in which the activity of one anima1 t;eflected the 
activity of its breed group. 
(J ) The gr atly incr tSed activity of both breed groups in 
the br ding s ason. At that time (autumn) , according to the findin&s 
of the previously discussed en rgy metabolism experiments , sheep 
are possibly metabolising at a low rate--a contradiction which may 
MASSEY AGRICULTURAL  COLLEGE 
LIBRARY PAU•icKSTON NuRTH, N.Z. 
200 
repay further investigation . 
(4) Foreleg measurements showed that the more active Cheviot 
was the shorter legged of the two breeds--an interesting finding 
when considered alongside the idea that longer legs will impart 
to the New Zealand Romney greater ease of travel and better leverage 
for hill climbin (Barton 19.54) .  
It is recognised, however, that these results were obtained 
in a New Zealand environment and are thus n o t  strictly comparable 
with work in South Africa or in the North of Scotland . Nevertheless , 
it is suggested that further inqui on the lines of the present 
investigation may be worth considering as the rangemeter is at least 
a "valid and standardised technique" .  There is no reason why the 
rangemeter s h ould not be developed as a carriE>Y .for an arsenal of 
other devices which would thus automatise a deal of the work of 
grazing studies . For example , pressure tiJt;e switches can be incor­
porated in the girth strap so that lying down periods can be measured 
and mapped within a day or week. The machine can also be modified 
to enable a ram to serve while wearing it. 
Haematolo ioal studies (Division "D" ) carried out with a 
specially created flock on one of the Massey Agricultural College 
hill fanns indicat d br ed diff renoes for total leucoc.yte counts , 
distribution of potassium values and packed red cell and blood volumes . 
In other re peot ; haemoglobin , M. c.v . ,  M. C.H. ,  M. C.H. cone . , the 
br eds were essentially equal. If the standards laid down by other 
201 
work rs can be accepted then neither of the flocks was in any way 
anaemic;  in fact their haemoglobin levels were good. The distinct 
breed difference in the total leucocyte counts { Cheviots ( 10 , 000 c .l1llll , 
Romneys ) 10 , 000 c .mmo blood ) is felt to be worthy of further investi­
gation for the high counts for the Romneys agreed perfectly with those 
of Karpov (1941 ) .  
One of the difficulties in this project was at all times the 
tendency for physiologists , biochemists , mathematicians and even 
agriculturalists to regard sheep as being identical but for their 
appearance . Much sheep work is reported without the breed being 
specified , which rather restricted the field of standards for 
comparison {this applied equally to all experimental divisions of 
the project ) and complicated the interpretation of some of the results . 
An interpretation of the breed differential in leucocyte counts is 
difficult, for the total counts were not split down into the composite 
types . However, so far as the high Romney leucocyte counts are 
concerned, it is  felt that they may indicate that th animals were 
livin under constitutional stress or they may be a breed charaot ristic 
possibly conn cted with the breed ' s traditional r sistance to foot 
rot and other marsitiand tA:·oubles . 
The whole blood and red blood cell pot ssium estimations revee� d 
that the Cheviots had thr e fifths , while the Romneys had only one 
quarter high values . There was agreement between the breeds on the 
actual values within a classification . 
202 
The distributions of whole blood and red blood cell potassium 
values pl<>ce the Cheviot and Romney in the categories of hill and 
lowland sheep respectively (Evans and Mounib 1957) even though the 
Romneys were obt ined fran New Zealand hill stocks . 
Evans and Mounib (1957) are inclined to connect with ' hardiness ' 
a large percentage of high blood potassium values within a breed. 
They derive their belief from a wide survey of hill and lowland 
breeds which revealed that the high values are found most often in 
hill sheep. The Romney figures presented in this project agree 
almost perfectly with those of Evans and Mounib for the Romney in 
England. However, the Cheviots do not agree nearly so well . This 
latter is possibly because Evans (1954) was dealing ldth the 
'' Caithness" variety which in breeding, background, hardiness and 
thriftiness (Thomas 1945) is different to the Border Cheviots 
from which the Massey Agricultural College Cheviots are descended . 
Selection of the foundation animals for importation into New Zealand 
may al o be implicated in that , by chance, it could have occurred 
that a large proportion of B.K. Cheviots and L.K. Romneys were 
among the original stud animals . 
Blood volume estimations carried out under the conditions required 
for a comparative study indicat d that the Cheviot had , per anima.l , 
a little over a litre mor blood (25 to JO percent) than the Romney 
and that trds difference remained statistically significant at th 
one percent level when it was related to body weight and ccmputed 
203 
surface area. 
The blood volume differences between the breeds are such as to 
bring the more active Cheviot into line with current theories on 
the requisites for athletic living. Dill (1938) ;  Brody (1945b) ;  Houssay 
(1955 ) ,  consider that a high oxygen capacity (blood volume) is neces­
sary for comparatively athletic living rather than any peculiarly 
high levels for haemoglobin or erythrocytes . The Cheviot would 
therefore appear to fulfill the stated requirements and backs them 
up by living athletically, as is evidenced by the rangemeter studies 
mentioned above. It is interesting to note that when xpressed 1n 
ml./Kg . the blood volume for the Romney corresponds with Courtice ' s 
( 1943) mongrel dogs 77 :79 and the Cheviot approaches the figure 
for greyhounds 93 &114. 
It is considered likely that a high blood volume may be related 
to other fUnctions--for so they may be called--such as hardiness . 
So far as the writer is aware, no oomparati ve studies have been 
carried out on blood volume in sheep stocks in the United Kingdom, 
and it i felt that in view of the pr sent findings , such studi6s 
are well warranted. 
The ha matological flock was also used to obtain general 
production data. It appears that the Cheviot was able to maintain 
a live-w ight differential over the Romney under the particular 
conditions and that by and large under those same conditions the 
breed was the more productive as was evidenced by the rate of lamb 
growth and the numbers of twins born. However, the weaning weights 
of the Romney lambs equalled the Cheviots ; but this may have been 
due to a differenc in lambing dates. 
Wool production is th point on which the Romney scored , for 
fifty percent mor wool is an appreciable difference . 
CONCLUSIONS 
It is concluded that the results obtained from the overall 
project indicate : -
(1 )  That the techniques developed for these investigations 
have proved their usefulness for physiological , behaviour and haem­
atological studies with sheep. 
(2 )  That the Border Cheviot sheep in New Zealand is  still 
possessed of physiological attributes which may explain its ability 
to produce better on poor hill country than the New Zealand Romney 
Marsh. 
(J ) The seasonal trends in the energy metabolism of sheep 
which towards autumn probably run counter to weight increases and 
grazing and ovulation activity, differences in br ed reaction to 
starvation and breed differences in activity and haematological 
picture indicate that a critical re-appraisal of the bases for 
reporting intraspecific nergy metabolis tudies is overdue . 
DIVISION "F" 
BIBLIOGRAPHY 
(In sections corresponding 
to the Div1s1ons 11K� "B'� JC� 
''D''and"Eh of thi Project . ) 
DIVISION "A" BACKGROUND TO PROJECT 
Bakewell , R. (17'25-1795 ) Of Dishley in Leicestershire , England 
Bartholomew. J .  (1953 ) Advanced atlas o f  modern geography 
(Bartholemew, Edinburgh ) 
a : .36 
b : .36 
Brody, s . (1945 ) Bioenergetics and growth (Reinhold , N e w  York , 
u.s .A . ) : 918 
Cresswell , E . (1951 ) Unpublished data 
Dill , D .  B . (1938 ) Life1 heat and altitude ( Cambridge , Harvard, 
University Press )  : 6 
206 
Garrard, G .  H . (1954) A survey of the agriculture of Kent (Royal 
Agricultural Society , England ) 
G.  B . Aeteorological Office (1957 ) Personal communication 
H. F.  R . (1951 ) Report of the Scottish hill farm research committee 
a : 127 
b : 129 
Hunter, R .  F. (1954) Brit . J .  An • Behav . II 2 : 75 
Jones ; l-1. G.  (1945 ) Cited from Thomas , J .  F. H . (1945 ) Sheep 
(Faber and Fab�r • London ) 
a : 129 
b : 129 
Keys , A . ;  Brozek , J . ; Henschel , A . ; Mickels en, O . ; Taylor , H .  Le  
{1950) The biology of human starvation (University of Min­
nesota Press )  : 3.38 
Niehols ; J .  E . (1928) J .  Text . Inst . (Manchr . ) 19 : 329 
Peren; G. S . ; Hewitt , w . R . R . ; Ballard , H . V . ; Phillips , T .  0 .  
(1951) Sheepfm.g Annu. , Massey Agric . Coil . ,  New Zealand 
14 : lll-137 
Reeve ; E. B . (1948) Nutr . Abstr . Rev , 17 : 8.31 
Riohards ; R . (1957) Personal communication 
207 
Scott , Sir \·Jalter (1771-1832 ) Of Abbotsford , Tweeds i de , Scotland 
Thomas , J .  F. H. (1945 ) Sheep ( Faber and Faber , London) 
a 31 
b : 125 
von Borstel, F. (1951) M . Agric . Sci . thesis , Massey Agric . Coll . , 
New Zealand 
(1957 ) l.Vannop , A . R .  
a Personal communication 
b : " " 
Wyatt , J .  A . (1957 ) Personal communication 
Youatt , \-{ . ( 1776-1847 ) 
New Edition (1894) 
a 312 
b : 334 
c : 335 
d 285 
e 286 
Sheep : their breeds management and diseases 
(London , Simkin, Harshall & Co. London ) 
DIVISION "B" ENERGY i·'iET.ABOLIS?-1 
STUDIES 
Benedict , F.  G. (1938) Carn. Inst . vlas h  • .  Pub . 503 
a 62 
b 84 
c 15 
d 39 
e 39-179 
f 148 
g I 206 
h 40 
1 51-55 
Benedict , F . G. ; Lee , R . C . (1937 ) Cam. Inst . Wash. , Pub . 489 1 16 
Blaxter, K. L.  (1954)  Cited from Hammond , J .  (1954) Progress in 
the physiology of farm animals ( Butterworths , London) : 18 
Brody, s .  (1930 ) Univ . Mo. Agric . Exp. Sta . Res . Bull. 143 
a : 7 
b : 13 
(1945 ) Bioenergetics and growth (Reinhold , New York , U.S .A . ) 
208 
a • 417 . 
b : 307 
c : 311 
d : 312 
e : 352-403 
r 40.3 
g I 228 
h 213 
I r 462 
(19.56) Personal communi cation 
Cantarrow, A . ;  Trumper, M . (1949) Clinical biochemistry (Saunders , & C o  • •  
Philadelphia & London) 
a 1 296-298 
b : 299 
Cochran, H. G. & Cox, G. M. (1957) Experimental designs (John 
Wi1ey & Sons , New York, U.S.A.  t Chapman & Hill Ltd. , London) 
Colonna, D. (1956) Personal communication 
Coop, I .  E . (1953) J .  Agric . Sci . 43 : 469 
Dougherty, R. W . ; Meredith , C .  D. ; Barrett, R.  B.  (1955) Amer. 
J .  Vet . Res . 16 : 89 
Forbes, E . B . a Fries , J .  A . ; Kriss, M . (1926) J .  Dairy S ci . 9 : 1.5 
Hel1berg, A . (1949) Roy. Swed . Acad. A ric.  Sci . Sec . report 5 
a t 132 
b : 133 
Keys , A . ; Brozek, J . ; Henschel , A . ; Michelsen, 0 . ; T�lor, H. L. 
(1950) The biology of human starvation (University of �·S.nne s o t a  
Press ) : 338 
Ueiber, • (1947} .Physiol. Rev. 27 
a 527 
b : 588 
c : .518 
Lines , E . w. ,  Pierce , A . t-J . (1931) Bull . Coun. Sci . Indus . Res. 
Aust . 55 : 27 
�tarston, H. R. (1948) Aust . J .  Sci.  Res . B I 
a : 110 
b no 
c r 107 
Merillat. L. A. (1921) Veterinary surge17 Ill (.Alexander Bger, 
Chicago. u.s.A.) • 343•350 
MUl"linw J .R . I Burtco. A. C. (l9�S) J .  Butr. 9 t 23)•2.59 
Pilgrim, A. F. (1�) A,ust .  J • Sc 1.. Res. B I : 132 
P.rosser, O .  L . J Bishop , D. W . a Brown .  F .  A. a Johna c:m ,  T. L· · l Wult£ , 
V. J .  (1?5�) Cemp&l"atlve animal pbyat.olou (Saundere a Co. , 
Ph1ladelph1a & Lond-cn) : 235-2)6 
R1tzm.an, E .  G . ' 'Senedict , F. G.  (19)8) Cam. %net. Wash. Pub. 494 
� : �ll) . 
b c 102 
0 t UJ•142 
d t 128 
8 I 1)6 
t t 142 
£ t 1.� 
'· h I );;(\ 
Rltbnet", M. (18)3) Cited fr Benediet, F. G.  (19)8) Garn. last. 
wash . Pub . 503 : 51 
Snedecor, G. w. (1946) Statistical methoda (Iowa State College 
P.re••> • �4 
Wlggers, C. J .  (l�9) Pbyaiolog 1n h ltb and disease (Lea A 
Feblser,Phlladelphia) 
a z  949 
'b t  9.SO 
Wood , 'l. _ . (1927) Animal nutr1 t:lcn (Un1veralt:r f1ltorial l>reaa) 
• 184 
lit�JUA• E .  o. a Ben.ttiat. r .  c • (1938) earn. :tnst. .  wasb. Pub. 494 
, ,.op 
(b) 1'h ttt!£) gp the&r M!&laol.1g mtea "le!4&Dc 99t .p!UIMJ O.Ut 
so as* e2r: tb£!1 tlleaR-adii 5'ilu ia»rt ttvra�'•• 
210 
Armsby, H. P . (1906)  The principles of animal nutrition (John Wiley 
& Sons , New York , U . S • . A . ) : 374 
( c )  The effect of 1 thyroxine on the oxv�en consumption, resuiratory 
rate and body weight o£ one Romney t1arsh ewe . 
Blaxter , K . L .  (1948 ) J .  Agric . Sci . 38 : 1 
Blaxter, K.  L . ; Reineke , E , P . ;  Crcmpton , E . W. ; Petersen , W. E . 
(1949 ) J .  Anim. Sci . 8 r 307 
Brody, s . (1945 ) Bioenergetics and growth (Reinhold, New York, 
u.s .A . ) z 171 
Hart � D , s . (1955 ) Proc . N . z . Soc . � . Prod. 15 57 
Jordan, R . M . (1954) J .  Anim. Sci . 13 : 438 
Kirton , A , H . (1957 ) M . Agric . Sci . thesis , Massey Agric . Col1 . 
New Zealand 
Turner, C. w. i Reineke , E . P . (1946) Mo • .Agric .  Exp . Sta . Res . 
Bull . z 397 
Warwick , E . J . : Childs , E . C . : Flower , A . E . ; Ham , W. E . (1948) 
J .  Anim. Sci . 7 : 198 
(d )  Methane emalation in sheep , 
Brody, s . (1945) Bioenergetics and growth (Reinhold , New York , 
u .s .A . ) : 315 
Blaxter , K .  L. (1954) Cited from Hammond , J .  (1954) Progress in 
the physiology of farm animals (Butterworths , London) : 26 
Ritzman, E . G. ; Benedict , F. G. (1938) Carn . Inst . Wash. Pub. 
494 : 25 ( Citing Klein, W . ,  Bioohem . Zeitsch (1916) 72 : 169 ) 
( e )  Regurgitation in �heep .. 
Bergmar1, H.  D. ; Dukes , He R . (1926 ) J .  Amer .  vet . m d .  Ass �  
22 : 600 ( Citing Toussaint , H. (1875 ) Arch. de Phisiol . 
norme et path. 7 r 141 ; Colin, G, C .  (1871 ) Triote · de 
Phisiol . comp . des Anim . Ed . 2 , 1 : 631 ; and Flourens , E . J .  
(1844) Mem. d'Anat , et Phisiol . comp . (Paris ) : 30 ) 
Dougherty, R. \i , (1956) Personal communication 
Dougherty, R . w . ; Habel , R . � . (19.5.5 ) Cornell Vet . 4.5 : 4.59 
Dougherty, R . . ;  Meredith , c . r. . (19.5.5 ) Amer . J .  vet . Res . 
16 : 96 
2ll 
tukes , H . H . ( 1955 )  The physioloeY of domestic animals (Boilliers , 
Tindall and Cox, London) : 322 
Kryzwanek , .t• . w. ( 1934) Vet. Bull . 4 : 390 
Stigler, R. (1931) Arch , F . Tier und Tier (B ) 4 : 613 
DIVIS I Ol�  " C" RANG !NG REHAVICUR ST Uf iES 
Burton, H . ; Castle , M . E . (19.50 ) J .  Dairy Res . 17 : 229 
Imgland , c . J . (19.54)  Brit . J .  Anim . Behav . II 2 : 56 
Harumond , J .  (195�) Progress in the phys i o l ocy of farm animal s 
(Butterworths , London ) : .586-6o2 
Hancock , J .  (19.50 ) N . z . J .  Sci . Tech . 32 : 22 
Hunter, R. F. (1954) Bnit. J .  Anim . B ehav . II 2 7.5 
Jones , M . G. ( 1928) Welsh J .  Agric . 4 : 194 
Louw, L . J . ;  Haveng a , C. �. ;  Hamersma, J .  (1948 ) Fmg. s . Afr. 
23 : 753 
Shepherd , J .  H . (1921 ) N . Dakota Agric . Exp. Sta. Bull . 154 : 8 
Stapledon, R. G . ; Jones , M . (1926 ) �elsh Plant Breed . Sta. Bull. 
H 5 : 50-.53 
Tribe , D. E. (1949) Emp. J .  exp. Agric. 17 : 110 
Van Rensburg, J .  A . (19.56 ) Fmg. s . Afr. 31 149 
von Borstel , F. (19.51) J .  Agrie . Sei . thesis , Massey Agrie . Coll . 
New Zealand 
Wallaee , D. B. ;  Kennedy , K. (1944) J .  Agrie . Sei . 34 : 192 
DIVISIO nnu HA.EMATOJ.PGI. L STUDIES 
Alleroft, w. M . (1941) J .  Agrie . Sei . 31 : 323-324 
Barcroft ,  J . ; Kennedy , J .  A . ;  r'.ason , H . F . ( 1939) J .  Physiol . 
95 : 162 
Bermetts , ,.,.  'T "  £1 . �--· • ; Chapman, F . 3 . (1937 ) Aust . vet . J .  13 : 140 
Bro�· , s . (1945 ) ioenergetics and gro��h (Reinhold , New York , 
U o �· • r o ) : 403 
Courtice , F . C .  (1943 ) J .  Physiol . 102 : 291 
Dacie , J .  V . ( 1951 ) Practical haematology ( J .  & A . Churchill 
Ltd . ,  Enoland ) 
212 
Dukes , H . H . ( 1947 ) The physiology of domestic animals ( Comstock 
Publishing Co . ,  Ne·.-1 York , U.S .A . ) 
a 36 , 23 , 46 
b : 61 
Svans , J .  v .  (1954 )  Nature (Lond . ) 174 : 931 
2:vans , J .  V • ; ;.1ounib , N . S . (1957 )  J .  A g r i c . Sci . ,48 436 
i<'ilm�r , J .  F . (1933 )  .tmst . vet . J .  9 : 163 
Gotsev , T . (1939)  J .  Physiol . 94 : 543 
Uregerson , V , j . I . ; Schiro , H. ( 1938 ) Amer . J .  Physiol . 121 291 
Greger.:>on, H . I .  (1951 )  Ann . Rev. Physiol . 13 . 397 . 
Ha.11ersma , P . J .  (1934 ) Onderstepoort J .  vet . Sci .  2 : 153 
Hansard , S . L . ; Butler , W . 0 . ; Comar , C . L . ; Hobbs , C.  s . ( 1953 )  
J .  Anim. Sci . 12 : 407-4ll 
Hewitt , �-J. R . R . (1947 ) Sheepfmg Jmnu. , Massey Agric . Coil . ,  New 
Zealand 10 : 48 
Holman, H. 1 . (1944) J .  Comp . Path. 54 30 
( 1945 ) J .  Comp . Path. 55 : 146 . 147 
(1947 ) Proc. roy • .Soc . Med. 40 : 186 
Josland , s .  W . (1933 ) N. Z . J .  Sci .  & Tech .  14 : 304 
Karpov , A . s . (1941 ) u . R . s . s . , ll . s . 31 : 719-721 
lerr, S.  E./J.W7) J.. biol.. Chem. 117 : 233 
Iushner, H. F• a Kltatava. O. N. (1938) �·� ,,. , J. s .  20 ;  47•.52 
Me Ler oy ,  G .  B. ; Br own ,  E .  !. a Smith, W. A· (19.52) J .  Antal. Sci. 
11 • 791 (hoe.)  
Har a t ont l:I .  R. ; Thomas , R . G . : M�e� D. ; Lines. E . i.<l . L . , MoDonald 
I. W . ; Mo3re , H . 0. ; lull• L .  B .  (1�8) Bull . C oun .  Sei� 
lr\dus . R�s. Au st . 113 : SS 
Mebrotr-. P., N. J Mu1liek, D., • a Kebar, N . D . {19$4) J .  Ani.m. 
Set. lJ 1 ,.026 (?roe.)  
Mill • • T. (19:32) Oorn ll Vet. 22 t  320 
Mollinson, P. L .  (19.')1) Blood transfusions in el1n1cal med1c1n 
(Bl.aekweU, Ox:rord) : :n-34 
Proseer• c. L . ' Bishop, D. w. ; Brown, F. A. a John, T. I •• ; Wultf, 
V .  J .  (1950) C at ive animal phy'siolog;y {Saunders 4 eo. • 
Philadelphia & London) 
a s 534 
b : .534 
Re• •·  • B . (1948) Nut r .  Abstr . Rev . 17 s 814 
Red • M. a thout , A. F. (1951) J .  Amer .  ve t .  med . As a .  118 1 254 
So by • P. (19.52) ord . Vet . lied . '4 : 929 ... 961 
Todd, • c . ; \Tlyant, ' • • ; s t one , w . M. ' Elmu, G .. w . (19.S:n 
Amer . J • v t .  Re a . 1) : 74-76 
Tur e•• c .  • I  Her�: . B .  A. (19�1) Mo . Agric. Exp. Sta. Re e  . 
Bull . 159 
tJnd rwood, · • J . # Harvey, R. J . : 13 eh, A. 13 . (19)9) Ans.t .. J. xp . 
B1ol . med. Set. 1? t 202 I· 
• I .  V t .  R s.  14 : 405 
Widdaa, w . F .  (1954) J .  Pbye1ol. 12.5 : lSp 
Wiggere., o .  J .  (1S49) Pbyaiol 
Febtaer. PhUadolphla} 
a : )86 
b f 390 
in h alth and disea ( a A 
,P}VISIQN "E " OVERMJ., PROJECf DIScuSSION 
Barton. R. A .  (1954) Sheepf'mg Annu. ,  Massey Agric . Co11. , New 
Zealand 17 : 151 
Brody, s . (1945 ) Bioenergetics and growth (Reinho1d, New York , 
u. s .A . ) 
a : 213 
b : 918 
Courtice, F . c .  (1943 ) J .  Physiol. 102 : 300 
214 
Dill , D . B .. (1938) Light, h at and altitude (C  ridge , Harvard, 
Univ. Pre s )  : 22 
Evans , J .  V. (1954) Nature (Lond. ) 174 : 931 
Evans , J .  V . ; Mounib, M. s . (1957 ) J .  Agric . ci . 48 : 436 
Firbank, T .  (1940) I bought a mountain ( G .  o. Harrap & Co . , 
London) : 15 
Hs.m.mond, J .  (1955 ) Progress in the physiology of' farm animals 
(Butterworths London) : 586-602 
Hellberg, A. (1949) Roy, Swed. Aead. Agric. Sci .  Sec . report 
5 : 133 
Houssay, B . A. ;  Lewis , J .  T . ;  Orias , G. ; Braun-Menendez , E . :  Hug , 
E. ; Fog11a, V. G. ; Leloir, L. F. (1955) Human physiology 
(McGraw-Hill , New York and London ) : 507 
Karpov, A • . s . (1941Y u . a . s . s . , N . s . 31 : 721 
Per n,  G. s . :  Hewitt , W . R. R. : Ballard, H. V. ; Phillips , T . O.  
(1951 ) Sheep!mg Annu. Massey Agric. Coll . ,  New Zealand 
14 : lll-137 
Thomas , J .  F. H.  (1945 ) Sheep (Faber & Faber, London) : 125 
APPENDIX "A" 
A.PPEJIDIX A T@ll 1 
Bloek I - 19th Sept.aber - 4th Oct.ober 1956 
C!:rgen 29P'!'PUon recordS 1n ;u.tres* oxrgen connaed per f1ye _, mrt.e peri_ocis, 
lloi!lmeD RMr4ota 
1 2 3 4 5 6 1 8 9 10 
1·.90 l·,JO 1 .99 1'.99 1.)) 1 • .52 1 .62 1 .42 1 .71 1 • .52 
1 .91 2,00 2.14 2·.00 1 • .52 1.72 1.81 1-.71 1 .62 1 .52 
2 .09 1 .90 2.20 1.90 1 .4.S 1.80 1.8.5 1·.80 1.,52 1 • .52 
2.1.7 1 .89 1 .89 1 .76 1 .3) 1 .57 1 .61 1 .42 1 .41 1.46 
l 
2. 10 1.99 2.18 1.71 1.23 1 .42 1 .91 1 .44 1 .80 1 .?2 
2.19 1.70 2,08 1 .81 1 .)3 1.70 1.70 1 .5.5 1 .61 1 .72 
2,02 1 .. 99 2 ,10 1 .81 1 .33 1 .71 1 .80 1 .33 1 .43 1 .)) 
1 ." 1 • .52 1 .96 1 .80 1 .23 1 • .54 1.80 1 .,51 1.42 1.42 
1 .8,5 1.,51 1 .88 1 .41 1 .22 1 .6o 1 .41 1 .42 1 .51 1 .-41 
1 .89 1 .51 1 .80 1.69 1 ,JJ 1 ,42 1 .70 1 .)2 1�42 1.2) 
* Corrected to Standard Temperature and Pressure 13 0\ 
APfiiPll A TAJQ 2 
ll.ock n 1th - 11th October l.9S6 
Qana t9P'Pmt.icz recor4• 1R lliru*&UM C9NJ1Md per !iD NPPH Pftloda 
aamota 
1 2 ' 4 .s ' 7 8 9 10 
1 .?4 1.� 1.54 1.54 o.f)7 1.?4 1.46 1 .)4 1 • .62 1.)0 
1 .. 82 1.64 1.� 1.44 0.96 1 .1.) 1.5) 1.,, 1.72 1.07 
1.5) 1.7) 1.44 1.6) 1.1.5 1.82 1.)4 1 .1) 1.7) 1.,, 
1 .)) 1.61 1.71 1.51 1.52 1.�;, 1 . 42  1.52 1.62 1 .)) 
1.62 1.42 1.11 1 .61 1.)) 1.52 1 ., 1.52 1.81 1 .4) 
1.4) 1 • .5) 1.72 1.24 1.62 1 .4) 1 .)4 1.$) 1 .72 1.24 
1.91 1.24 1.62 1.,5) 1.1.5 1.4) 1 • .5) 1.34 1 .&) 1.)4 
1.64 1.16 1 .4) 1 .24 1 . 10 1 .34 1.44 1.06 1 .4) 1.1.5 24 bre .  
na.rnu.-
1 .)-.S 1.1.5 1.28 o.96 0.96 1.2.5 1 .)4 1.2.5 1.26 0.96 48 ...  
.tarnttoa 
* Corrected to Standard Temperature and Pressure 
� -.,J 
APPEIDIX A T.ABLE 3 
Block m 2)rd January - )l"d February 19.57 
Oxygen conammt.ion ncqrds in ll�,..ox.ys:en coneaed per fi.T! ainut.e peri.odiJ 
llopga Ol!yi.oy 
4 5 1.5 . 16 17 7 8 10 20 21 
1 .7) 1 • .).5 1 .7.) 1 .47 1 .71 1.40 1 .54 2.11 1.)0 1.19 
1 . 19 1 .57 1.61 - ·1.18 1 .76 1 .).) 1 .)3 2.10 1 . 14 1.4.) 
1 .42 1 .61 1.37 1.28 2.04 1.,52 1.42 2.09 . 1.18 1.66 
1 .67 1 .3.) 2 .00 2.19 1.8o 1.28 1.24 1 .)8 1 . 19 1.19 
1 .)8 1.32 1 .,Sl! 1 .71 1 .42 1 . 14 1.3) 1 .86 1 .28 1 .37 
'!f-
1 .70 1 .41 . 1 .79 1 .41 1 .41. 1 .27 1.41 2 07 ' . 1 .�1 1.)2 
1 • .51 1·S7 1 .76 1 .61 1 .47 1 . 14 1.2) 1 .47 1.)8 1 .14 
1 .4) 1.26 1 .j2 1.)8 1.,52 1 .19 1.62 1 .66 1 . 14 1.09 
1 .42 1 . 1) 1.42 1.37 1 .42 1 .1) 1 • .)7 2.09 1.00 1 .18 
1.40 1 .49 1.;4 1.§5 1 • .59 1 . 16 t • .so 1 .40 l .U.O 1.26 
1\) I-' • Correct.8d to Standard '!'�ature � hoeasure m 
APPENDIX A TABLE 4 
Block IIIa - 8th - 12th February 1957 
Qxygen consumption records in litres* oxygen consumed per five minute periods 
Romnen 
4 5 15 16 17 
1 .42 1 .23 1 .69 1 .37 1 .66 
1 .42 1 .37 2 .21 1 .35 1 .61 
1 .54 1 • .54 1 .40 1 . 16 1 .59 
1 .68 1 .44 1 .53 1 .44 1 .. 49 
1 . 7.3 1 .6) 1 .. 87 1 .8) 1 .7) 
• Corrected to Standard Temperature and Pressure 
Cheviots 
7 8 10 20 21 
0.95 0 .99 2 . 18 1 .28 1 .)) 
1 .04 1 . 15 1 .4) 1 .47 1 .24 
1 . 15 1 . 25 1 .64 1 .44 1 .05 
1 . 15 1 .44 1 .43 1 .)4 1 .24 
1 .45 1 .54 2.22 1 .45 1 .30 
1\) 1-' "' 
APPENDIX A TABLE 5 
Block IV 29th March - 7th April 1957 
Qx;ygen consumption records in litres• o:x;,ygen consumed per five minute periods 
Romneys Cheviots 
17 X 16 15 5 8 7 20 y 10 
1 .39 1 . 19 1 . 77 1 . 53 1 .48 1 . 15 1 . 15 1 .20 1 . 19 1 .73 
1 . 14 1 . 09 1 .53 1 .28 1 .29 1 . 09 1 .00 1 . 19 1 . 19 1 .24 
1 .)0 1 .45 1 .54 1 . 25 1 .59 0 .87 1 . 01 1 .25 1 . 06 1 .25 
1 .36 1 .36 1 .42 1 . 13 1 .50 0 .98 0.97 1 . 03 1 . 27 1 .36 
1 . 08 1 .36 1 .45 1 .26 1 .57 1 . 08 1 . 08 0 .97 1 .37 1 . 08 
' 
1 .)7 1 .37 1 .52 1 .47 1 .4) 0 .99 1 .04 1 .24 1 • .56 1 .)7 
1 .)8 1 .)8 1 .48 1 . 52 1 .)8 1 . 1-9 1 . 10 1 . 19 1 . 14 1 .29 
1 . 29 1 .63 1 .5) 1 .53 1 .68 1 .29 1 . 15 1 .24 1 .24 1 .58 
1 .)1  1 .26 1 .)6 1 .46 1 . 16 1 . 17 0 .97 1 .46 1 . 12 1 . 26 
1 .29 1 .24 1 .)8 1 .)8 1 . 18 1 . 13 0 . 89 0 .94 1 .22 1 .27 
l\) 1\) 
• Corrected to Standard Temperature and Pres sure o_ 
APPENDIX A TABLE 6 
Block V 6th - 17th April 1957 
Qxygen consumption records in litres*  oxygen consumed per five minute periods 
Romneys Cheviots 
17 X 16 15 5 a 7 20 y 10 
1 .)7 1 .22 1 .)0 1 .)1 1 . 12 0 .98 0 .88 1 . 12 1 . 02 1 . 17 
1 . 16 1 . 18 0 .98 1 .2) 1 . 08 0 .88 0 .97 0 .9) 1 . 2) 1 . 0) 
1 . 18 0.98 1 . 06 1 . 07 1 . 07 0 .82 1 . 02 0 .88 1 .07 0 .97 
1 .00 1 . 15 0 .96 1 -.24 1 . 00 0 .72 0 .72 0 .77 1 .05 . 0 .91 
1 . 01 1 . 01 1 . 10 1 . 10 1 .06 0 .77 0 .72 0 .86 1 .01 0.91 
1 . 10 1 . 06 1 . 10 0 .96 0 .91 0 .72 0 .72 0 .67 1 .01  0 .91 
0 .95 1 .04  0.9.!) � .91 1 e 09 0.72 0 .76 0 .81 o .81 o.et 
0 .96 1 .25 1 . 01 0.82 0 .87 0 .72 0 .91 0 .72 0 .91 0.77 
1 . 05 1 .04  1 . 09 0 .90 0 .90 0 .66 0 .81 0 .67 0 .81 0 .81 
0 .95 1 . 14 0 .81 0 .80 0 .76 0 .66 0 .66 0 .76 0 .90 0 .76 
N 
• Corrected to Standard Temperature and Pressure � 
APPENDIX A TABLE 7 
Block VI 18th - 27th April 1957 
Oxvgen consumption records in litres* oxygen consumed per five minute periods 
Romnays Cheviots 
17 X 16 15 5 8 7 20 y 10 
0 .85 1 . 09 0.99 0.95 0 .85 0 .71 0 .80 0 .90 0 .99 0 .76 
1 *05 1 . 15 0.91 0.96 0 .86 0 .86 0 .81 0 .96 1 .05 0 .86 
1 . 10 1 . 09 0 .96 1 .29 0 .95 0 .77 0.86 0 .86 1 . 05 0 .86 
1 .23 1 . 13 1 .23 1 . 23 0.95 o . 8o 0 .95 1 . 04 1 .09  1 . 04 
1 .33 1 . 09 1 .23 1 .2e 1 . 13 0 .95 1 . 14 1 .o4 1 .37 0 .99 
1 .24 1 . 15 1 .29 1 .34 1 . 10 0 .86 1 . 14 1 . 10 1 . 19 1 .24 
1 .37 1 . 14 1 .33 1 .23 1 .04 0 .99 0 .99 1 . 18 1 .33 1 . 18 
1 .58 1 . 10 1 .24 1 .29 1 . 10 0 .86 o . 86 1 . 10 1 .09 1 . 24 
1 .42 1 . 1) 1 . 61 1 .)7 1 .)2 1 .2) 1 . 18 1 . 18 1 . 61 1 . 61 
1 .)4 1 . 10 1 .6) 1 .43 1 .29 1 . 24 1 . 14 1 .05 1 .4) 1 .43 
N 
• Corrected to Standard Temperature and Pressure N N 
APPENDIX A TABLE 8 
ANALYSES OF VARIAN� 
Qxygen consumption in litres• consumed per 
Sources D.F.  Variance 
�1o� � 
Breeds 1 1.1859 
Error 8 o.4J99 
Block I;t 
Breeds 1 0.0049 
Error 8 0.2007 
Block III 
Breeds 1 0.)969 
Error 8 0.))68 
ilock Ina 
Breeds 1 0.4570 
Error 8 o.l95J 
�lock !! 
Breeds 1 1.1004 
Error 8 0.1048 
Block !. 
Breeds 1 0 .9025 
Error 8 0 .0472 
BJ:gck VI 
Breeds 1 0 .)721 
Error 8 O.lo4J 
5 minute§ 
F. 
2.70 
0 .02 
1.18 
2 .)4 
10.50 
19 .12 
J-.57 
... Corrected to Standard Temperature and Pressure 
per animal 
Nec.F 
5� 
22J 
1� 
5.)2 ll .JO 
• • 
• " 
" • 
" " 
n .. 
11 " 
224 
APPF;ND!X A T�.BJ& 9 
bHALYS[$ QF VARIAN� 
Ox.vgen oQrununpt'=on 1n 1'-rus• PQilSW!S per 5 minutes pgr Kg. body n1§t 
Source D.F. Variance F , Neo , F 
5� l� 
B;lQg}S ;t 
Breeds 1 0,0005 2 ,50 s .Jz ll,)O 
Error 8 0,0002 
��2gh ;u 
Breed8 1 0,00000 o.oo " " 
Error 8 0,01283 
�:l.aPh �!I 
Breeds l 0,00006 0,)2 " n 
Error 8 0,00019 
Bl2gk I;IIa 
Breeds l 0,000048 o.sJ .. " 
Error 8 0,000090 
i.2gls � 
Breeds l o.oooo6 o.os 11 .. 
Error 8 0,00112 
il.2gls I 
Breeds l 0,00012.54 2.42 " • 
Error 8 0,0000.519 
Bl22ls :a 
l3 eds l 0,0000 o.po • • 
Error 8 0,0000 
• Corrected t.o Standard· T pe ture and Pre sure 
22.5 
APPENDIX A TA.BI.E 10 
ANALYSES OF VARIANCE 
Oxygen consumption in litres• consum&d per 5 minutes per sq. metre 
• 
Block 
I 
II 
Ill 
Ilia 
IV 
V 
VI 
Sources 
Breeds 
Error A 
Days 
Error B 
computed surface area 
D.F.  Variance 
1 1 . 094-1 
8 0 . 2948 
9 1 .0881 
81 0 . 0120 
Breeds ( t9 9th day) 1 0 .0083 
Error A 8 0 .2.071 
Days (to 7th day) 6 0 . 0188 
Error B 54 0 . 0259 
Breeds 1 0 .0075 
Error A 8 0.)106 
Days 9 0 .0457 
Error B 81 0 . 0)29 
Breeds 1 0 .1812 
Error A 8 0.1703 
Days 4 0 .1271 
Error B 36 0 .0300 
Breeds 1 0.4186 
Error A 8 0 .0894 
Days 9 0 . 0459 
Position 9 o .oo76 
Error B 72 0 .0131 
Breeds 1 0 .3982 
Error A 8 0 . 0214 
Days 9 0 .0817 
Position 9 0 .004.5 
Error B 72 0 . 0065 
Breeds 1 0 .1076 
Error A 8 0 .0742 
Days 9 0.1966 
Error B 81 0 . 0116 
F. 
3 .71 
7 .34 
0 .04 
0.73 
0 .02 
1 .)9 
1.06 
4 .24 
4.68 
) .50 
0 .58 
18.61 
12 .57 
o ,69 
1.45 
16.95 
Corrected to Standard Temperature and Pressure 
Ne�.F  · 
5� 1� 
5.32 11 .30 
2 .01 2.67 
5 .)2 11 .)0 
2 . 29 ) .18 
5 .32 11 .)0 
2 . 01 2 .67 
5 .)2 11 .)0 
2 .61 ) .8) 
5 .32 11 .)0 
2 . 01 2 .67 
2 . 01 2 .67 
5 ·32 11 .)0 
2 .01 2 . 67 
2 ,01 2 .67 
5 .)2 11 .30 
2 . 01 2 .67 
2 2 6  
APPENDIX "B" 
228 
APPEN:CIX B TABLE 1 
Analyses of variance production records haematology flock 
Sources D.F.  Variance F. Nec .F 
.5% 1� 
Ewe weights Breeds 1 9006.0 12 .19 3 . 98 ? . 01 
6/?/56 Err;)r 68 739 .0 
Ewe weights Breeds 1 631 . 0  5 -95 4.00 ? . 08 
27/9/56 Error 62 106 . 0 
Ewe 1ofeights Breeds 1 938.0 8 . 85 4. 00 ? . 08 
21/2/57 Error 62 106 . 0 
E�1e weights Bre�ds 1 451 . 3 3 • . 5 5  I.J. . OO 7 . 08 
30/?/57 Error 61 12? .2 
Lamb weights Breeds 1 920 .95 3 .41 4 . 03 ? .1? 
12/12/56 Error 51 269 .95 
Lamb production Breeds 1 ?54 .65 5 .)4 4. 03 ? . 17 
2?/9 - 12/12/56 Error 50 141 .34 
\vool production Breeds 1 133 .60 98 .96 4. 00 ? . 08 
22/ll/56 Error 64 1.35 
229 
APPENDIX B TABLE 2 
Analyses of varian9e haemato1ogical data 9/7/56 
Sources D.F. Variance F.  Nec .F 
5"' 1� 
Haemoglobin Breeds 1 16.0950 20 .,54 3 .98 7 .01 
Error 83 0 .7835 
P . c . v .  Breeds 1 189 .00 11 . 80 ) .98 7 .01 
Error 80 16. 01 
Erythrocytes Breeds 1 11 . 5598 6 .55 4.60 8.86 
Error 14 1 .7654 
Leucocytes Breeds 1 342 .149 108. 58 3 .98 7 . 01 
Error 81 ) .151 
M . c . v .  Breeds 1 1LJ. .8� 3 . 38 4. 6o 8.86 
Error ll} L;..38 
M . C. H . Breeds 1 2.32 1.72 4 . 60 8.86 
Error 14 1 .35 
M .C .H . ( conc . ) Br$eds 1 0 .75 0 . 01 3 .98 7 .01 
Error eo 5.4·0 
2)0 
APPENDIX B TABlE J 
Analyses or variance haematological data 3/10/56 
Sources D.  F. Variance F. Nec .F  
5� 1� 
Haemoglobin Breeds l o.oo o .oo 4 . 00 7.08 
Error 61 1 .24 
P .c .v .  Breeds 1 o .6o 0 . 04 4.oo 7.08 
Error 58 16.49 
Erythrocytes Breeds 1 0.465 0.22 4.22 7.72 
Error 26 2 .080 
Leucooyt.es Breeds 1 96. 02 16.99 4 . 00 7.08 
Error 60 5 . 65 
!1 . C .V . Breeds J_ 10 .11 0.)6 4.24 7 .72 
Error 25 27 .99 
1-1 . c . H . Breeds 1 2.57 0 .92 4 .24 7 .72 
Error 25 2 .79 
1-1 . c.H.  ( cone . )  Breeds 1 0 .22 0 .10 4 . 00 7 .08 
Error 56 2 .27 
231 
APPENDIX B TABLE 4 
Analyses of variance - haematological data 19/2/57 
Sources D.F.  Variance F. Nec.F 
5� 1� 
Haemoglobin Breeds 1 2 .2160 o .ao 4 . 00 z . oa 
Error 57 2 .7572 
P . C. V .  Breeds 1 151 .54 4.13 4 .00 7 .08 
Error 57 36.62 
Erythrocytes Breeds 1 3 .0628 1 .50 4 .24 ? .77 
Error 25 2 .0374 
Leucocytes Breeds 1 36.331 6 .80 4 .oo ? .08 
Error 57 5 -375 
l-1 . C.V . Breeds 1 4.99 1.6) 4 .26 ? .82 
Error 24 3 . 06 
M. C.H . Breeds 1 0.21 0 .07 4 .26 ? . 82 
Error 24 3 .14 
M . c.H.  ( cone . ) Breeds 1 2 .32 0 .12 4,00 ? .08 
Error 56 18.49 
Total blood Breeds· 1 14.25 38.5 4 . 03 ? .17 
vol. litres Error 49 0.37 
Erythrocytes Breeds 1 4.0328 22 .40 4 .03 ? .17 
litres Error 49 0 .1800 
Plasma Breeds 1 3 .6651 31 .60 4 .03 ? .17 
litres Error 49 0 .1160 
Weight in Breeds 1 181.20 8.69 4 .03 ? .17 
kilograms Error 50 20.85 
Blood Breeds 1 3035 .96 21.8  4 .03 ? .17 
mJ../Kg. Error 49 139 .03 
Erythrocytes Breeds 1 1060.38 15 .89 4 .03 ? .17 
mJ./Kg. Error 49 66.75 
232 
APPENDIX B TABLE 4 (Cont1) 
Plasma Breeds • 1 499.56 8.44 4.03 7 .17 
ml. /Kg. Error 49 59.23 
Surface area Breeds 1 0 . 0326 9 .05 4.03 7 . 17 
( Sq.  metres Error 49 0 . 0036 
computed) 
Blood volum.e/ Breeds 1 8. 0936 31. 27 4.03 7 .17 
' 
Sq. metres Error 49 0 .2588 
computed area 
Erythrocytes Breeds 1 2.1205 15 .20 4 .03 7 .17 
vol . /  Error 49 0 . 1395 
Sq. metres 
computed area 
Plasma vol ./  Breeds 1 1 .6038 16.90 4 .03 7 . 17 
Sq. metres Error 49 0 . 0949 
computed area 
APPENDIX B TABlE 5 
Analyses of variance - comparison of haematologieal data between 
Haemoglobin 
Breed 
Cheviots 
Romneys 
Leucocytes 
Breed 
Cheviots 
July : October 1956 
October 1956 : February 1957 
July 1956 : February 1957 
Dates Sources D.F.  Variance F. 
July : Oct . Dates 1 64.870 98.14 
Error 7:3 o. 661 
Oct . :Feb. Dates 1 72 .4:3 45 .84 
Error 61 1 .58 
July:Feb. Dates 1 0 .112 0 .09 
Error 70 1 .210 
July:Oct. Dates 1 10 .445 9 .81 
Error 71 1 . 065 
Oct. :Feb. Dates 1 13 . 079 5 .48 
Error 57 2 .)87 
July :Feb. Dates 1 11 .681 5 .94 
Error 70 1 .966 
July: Oct . Dates 1 2.21 0.44 
Error 71 5 . 04 
Oct. : Feb. Dates 1 6.886 1 .43 
Error 60 4.820 
July zFeb. Dates 1 17 .94 3 -93 
Error 69 4 .56 
Nec.F  
5J, 
) .98 
4.00 
) .98 
3 . 98 
4.00 
) .98 
) .98 
4.00 
) .98 
233 
1� 
7·01 
7 .08 
7.01 
7 .01 
7 . 08 
7 .01 
7 . 01 
7 .08 
7 . 01 
APPENDIX B TABLE 2 !Cont. l 
Romneys July:Oct . Dates 1 0 .386 o . o6 3 .98 7 . 01 
Error 70 6 .no 
Oct . r Feb . Dates 1 0 .949 0.15 4.00 7 . 08 
Error 57 6 .247 
Ju1y: Feb . Dates 1 0 .191 0 . 03 3 .98 7 . 01 
Error 69 6.3.59 
P.c.v . • s  
Breed 
Cheviots July: Oct . Dates 1 1027.34 70 .27 3 .98 7 . 01 
Error 70 14.62 
Oct . : Feb. Dates 1 94.82 ) . 76 4. 00 7 . 08 
Error 58 25. 24 
Jul.y :Feb. Dates 1 182 .41 9 .90 ).98 7 . 01 
Error 70 18 .42 
Romneys July: Oct . Dates 1 399 . 10  22.)5 3 .98 7 .01 
Error 68 17. 86 
Oct. :Feb. Dates 1 30.1  0 .96 4.00 7 .08 
Error 57 31 .2  
July :Feb. Dates 1 194.1  6 .)2 3 .98 7.01 
Error 67 30 .7  
s;rohrogyt.es 
Bre� 
Cheviots July : Oct. Dates 1 35.9278 15 .43 4.38 8 .18 
Error 19 2 .)280 
Oot. : Feb. Dates 1 ).4517 1 .95 4. 24 7 . 77 
Error 25 1.7736 
July: Feb. Dates 1 19.9151 11 .60 4.).5 . 8.10 
Error 20 1.?165 
Romneys 
M, c.v. 
Breed 
Cheviots 
Romneys 
M,C,H, 
Breed 
Cheviots 
APPENDIX B TA."3LE 
July: Oct . Dates 1 
Error 21 
Oct . : Feb Dates 1 
Error 26 
July : Feb , Dates 1 
Error 19 
Ju1y:Oct . Dates 1 
Error 18 
Oct.  : Feb. Dates 1 
Error 24 
July: Feb . Dates 1 
Error 20 
Ju1y : Oct . Dates 1 
Error 21 
Oct. :Feb. Dates 1 
Error 25 
Ju1y : Feb , Dates 1 
Error 18 
July : Oct . Dates 1 
Error 18 
Oct. tFeb, Dates 1 
Error 24 
Ju1y : Feb. Dates 1 
Error 20 
23.5 
5 � Cont1} 
2 .8186 1 .71 4 ,J2 8 ,02 
1 .64.54 
O,J277 0 ,14 4 .22 7 .72 
2,JJJJ 
4 .4877 2 . 06 4 .)8 8 .18 
2 .1748 
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MASSEY AGRICULTURAL COLLEGE 
LIBRARY PAUilER3TON l'iORTH, N.Z.