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Food intake and grazing behaviour of sheep

varying in body condition

G. W. Arnold and H. A. Birrell

Animal Production / Volume 24 / Issue 03 / June 1977, pp 343 - 353

DOI: 10.1017/S0003356100011855, Published online: 02 September 2010

Link to this article: http://journals.cambridge.org/abstract_S0003356100011855

How to cite this article:

G. W. Arnold and H. A. Birrell (1977). Food intake and grazing behaviour of sheep
varying in body condition. Animal Production, 24, pp 343-353 doi:10.1017/
S0003356100011855

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Anim. Prod. 1977, 24: 343-353

FOOD INTAKE AND GRAZING BEHAVIOUR OF SHEEP

VARYING IN BODY CONDITION

G. W. ARNOLD
Division of Land Resources Management, CSIRO, Private Bag,
PO Wembley, Western Australia 6014
AND
H . A. BlRRELL
Department of Agriculture, Pastoral Research Station, Box 180,
Hamilton, Victoria, Australia 3300

SUMMARY

Herbage intake and grazing times were measured in four experiments

in which adult sheep grazed short and long pastures. The digestible
organic matter (DOM) intake of adult wooled Merino and Corriedale
wethers was higher, both absolutely and per kg live weight, when
they were lower in live weight due to prior under-nutrition. The
relative differences were similar on both abundant and scant pasture
and were proportional to the relative differences in live weight. The
higher intakes were achieved by differences in the time spent grazing
and in the rate of intake per hour of grazing. DOM intake of shorn
sheep was increased by up to 44 % per kg live weight within 5 weeks
of shearing in two experiments when the mean temperature was
8°C but no increase in DOM intake was observed in four other
experiments when the mean temperature was 11°C. Shorn sheep
achieved higher intakes on both abundant and scant pastures by
increasing intake per hour of grazing.

INTRODUCTION

THE intake of herbage by grazing sheep is influenced by the age, size, weight
and physiological state of the animal, climatic conditions and the availability
and quality of herbage on offer (Arnold, 1970). Little is known, however,
about the interactions between body condition, cold stress and pasture
availability on the intake of grazing sheep.
Donnelly, Davidson and Freer (1974) found that sheep which had been
fed for 6 weeks so that they differed by 9 kg in live weight, had similar food
intakes but ate less on sparse than on abundant pastures. Several studies
with housed sheep have indicated variable increases in food intake following
shearing (Wodzicka-Tomaszewska, 1963a and b, 1964, 1966; Donnelly,
Lynch and Webster, 1974) but reasons for these differences have not been
identified. Under grazing conditions, at Armidale, New South Wales,
Wheeler, Reardon and Lambourne (1963) reported that food intake increased
by 43 % in the 2 weeks following shearing in spring or autumn, and Hutchinson
and McRae (1969) reported increases of 30% in the first 6 days and 59% in
the second 6 days post shearing in winter. In neither study were changes
343
 344 ARNOLD AND BIRRELL

after 2 weeks examined. Hutchinson and McRae found that the relative
increases in intake following shearing were similar in sheep differing by
10 kg in live weight and that similar relative increases occurred at low and very
low levels of pasture availability. They also found that the previous rate of
loss of live weight affected food intake.
The effects of fleece length, temperature, wind velocity and level of
nutrition on heat production and critical temperatures have been studied
in housed sheep (Armstrong, Blaxter, Graham and Wainman, 1959; Graham,
1964; Joyce and Blaxter, 1964). Physiological resistance to a sudden extreme
cold stress is influenced more by previous rate of loss of live weight than by
current level of nutrition (Slee and Sykes, 1967; Sykes and Slee, 1968).
Donnelly, Lynch and Webster (1974) found that acclimatization to a moderate
cold stress occurs within 14 days of shearing in summer so that critical
temperature decreased by 10°C. The behavioural response to cold stress
will be influenced by the above factors and will be expressed in the time
sheep spend grazing, the rate of eating and the use of available shelter.
Hutchinson and McRae (1969) found that the higher food intake of shorn
sheep was achieved mainly because shorn sheep ate faster than wooled sheep.
Shorn sheep, however, stood for longer and this would possibly increase their
heat losses. In none of the studies of response to cold stress has the energy
balance of the sheep been measured, even in terms of change in live weight.
In a preliminary study (Arnold, 1976) the behavioural responses of thin
and fat shorn adult Merino wethers to cold stress were shown to differ
markedly even though rate of loss of live weight before shearing was similar.
This result was at variance with that of Hutchinson and McRae (1969).
This paper reports the results of a series of experiments which were
conducted at three locations in southern Australia to examine the effects
of shearing, body condition and pasture availability on food intake, grazing
behaviour and live-weight change in a range of climatic conditions. The aim
was to try to relate responses to shearing to climatic conditions.

MATERIAL AND METHODS

Seven experiments were conducted in different years in different locations
using castrated adult male sheep (wethers). The procedures were similar in
all experiments and details of the sheep and the pastures are given in Table 1.
Meteorological data are given in Table 2.

TABLE 1
Location, type of sheep, body condition and pasture levels in the experiments
Body condition Pasture level
No. and type of (kg live weight) (kg/ha)
sheep and no. per
Experiment Location treatment Type of pasture Fat Thin Abundant Scani
1 CSIRO, Dickson Expt. 6 3-yr-old Merino Phalaris, sub. clover, 62 38 2200 580
Stn, Canberra, ACT. wethers annual grasses
2 CSIRO Yalanbee Expt. 6 3-yr-old Merino Sub. clover, annual grasses, 55 44 3760 1680
Stn, Bakers Hill, WA. wethers capeweed
3 Victoria Dept. of Agric, 8 3-yr-old Corrie- Perennial ryegrass, sub. 51 38 2500 700
4 Pastoral Research Stn, dale wethers clover, annual grasses 43 33 2250 450
Hamilton, Vic. 8 2-yr-old Corrie-
dale wethers
5 CSIRO, Yalanbee Expt. 12 \ 3-yr-old Sub. clover, annual grasses, Not varied
6 Stn, Bakers Hill, WA. 10 Merino capeweed
7 10 J wethers
 FOOD INTAKE AND GRAZING BEHAVIOUR OF SHEEP 345

TABLE 2
Mean minimum and maximum temperatures (°C), number of 'wef'f days and
windspeed at 09.00 hr (mjsec) in Experiments 2-7
Measurement Experiment 2 Experiment 3 Experiment 4 Experiment 5 Experiment 6 Experiment 7
WCCK ditcr
shearing Min. Max. Min. Max. ^Min. Max. Min. Max. Min. Max. Min. Max.
1 6-2 131 3-7 13-9 4-8 12-9 5-2 14-3 80 14-5 6-2 15-3
2 5-8 17-7 5-4 12-1 2-9 10-8
3 6-5 181 3-4 150 3-9 11-6
4 6-8 19-9 2-6 100 3-9 13-5

Days Experiment 2 Experiment 3 Experiment 4 Experiment 5 Experiment 6 Experiment 7

Wet da;ys Wind Wet days Wind Wet days Wind Wet days Wind Wet days Wind Wet days Wind
0-10 0 2-4 3 5-2 1 2-3 2 40 3 3-2 5 3-3
11-20 0 1-9 4 4-7 6 4-9
21-30 0 2-3 4 6-7 9 4-7
t Days with rainfall in excess of 1 mm.

Experiment 1
This was a factorial experiment with two levels of pasture (high and low)
and two levels of body condition (fat and thin). The herbage intake and
grazing times of all sheep were measured over two 72-hr periods. The first
period began 5 days after sheep were put on to the pastures, and the second
period commenced 5 weeks later.
Experiment 2
This was a factorial experiment with two levels of pasture (high and low),
two levels of body condition (fat and thin) and two levels of wool (6 mo
fleece v. just shorn). Herbage intake by all sheep and grazing times for three
sheep per group were measured. Sheep were shorn on 25 August 1967 and
intake measurements began 3 days later and continued for 4 weeks. Rectal
temperatures were measured at 10.00 hr on days 12 and 14 after shearing.
Sheep were weighed weekly.

Experiment 3
The design and measurements were the same as for Experiment 2. Six
of the eight sheep were chosen randomly for measuring herbage intake and
grazing time. The sheep were shorn on 28 August 1968. Rectal temperatures
were measured at 06.00 hr on days 2, 3, 5, 9 and 41 after shearing.

Experiment 4
This was a repeat of Experiment 3 in the following year. Sheep were
shorn on 20 August 1969. Rectal temperatures were measured at 06.00 hr
on days 8, 9, 16, 19 and 26 after shearing.

Experiments 5, 6 and 7
All three experiments were located on the same Experiment Station. In
each experiment, comparisons of herbage intake were made on flocks of
Merino wethers grazing abundant pasture; half the sheep were shorn the
week after measurements began. In Experiment 5 groups of 12 sheep were
used, and in Experiments 6 and 7 groups of 10 sheep. Fleece lengths after
shearing were measured in Experiments 5 and 7 by means of Vernier calipers
at 10 sites on the body. Rectal temperatures were taken daily at 08.00 hr.
346 ARNOLD AND BIRREIX

General procedures
Experimental sheep were selected to represent the middle range of live
weight in flocks of about 200 sheep and were allocated to treatment groups
by random stratification on live weights. Differences in body condition
were achieved by varying the stocking rate for a period of at least 2 mo.
Since live weight and body composition are closely correlated (Gharaybeh,
Arnold, McManus, and Dudzinski, 1966) the live weights were considered
to reflect differences in body composition.
Where shearing was imposed as a treatment, sheep were shorn 2 days
before they were put on to the experimental pastures in Experiments 2 to 4
and 1 week after going on to the experimental pastures in Experiments 5 to 7.
Experimental pastures with high and low feed availability were obtained
by imposing differential grazing pressures for several months prior to the
experiment.
In Experiments 2 to 4, all groups of sheep were separate but, within each
pasture level, groups were moved between paddocks daily to avoid con-
founding any paddock differences with sheep treatments. In Experiments 5,
6 and 7, shorn and unshorn sheep grazed together.
Measurements began within 5 days of shearing in Experiments 2 to 4,
but sheep were trained to wear faecal harnesses and grazing clocks for 2
weeks prior to this. In Experiments 5 to 7, intakes were measured for 1
week before shearing as well as after shearing. Grazing times were measured
using vibrarecorders (Allden, 1962). Herbage intake was estimated from
the total faecal nitrogen in oven-dried faeces using a regression of digestible
organic matter (DOM) intake on total faecal nitrogen (TFN) (D. B. Purser,
personal communication). The equation used was
DOM intake = 83-3 xTFN+74 g±35.
In Experiments 2 to 4, the grazing times of sheep were recorded on days
1 to 4 and faeces were collected on days 2 to 5 of each week, both inclusive.
In Experiment 5, 6 and 7 measurements were made daily for 10 days after
shearing.
Faeces were bulked for each sheep in each period, dried at 80°C in a
forced-draught oven, and then sub-sampled. The sub-sample was ground
to pass through a 1 mm sieve, and samples were analysed for nitrogen by
an automatic colorimetric method which involved distillation and the use
of indophenol blue, and for ash content by ignition in a muffle furnace for
3 hr at 55O°C.
Rectal temperatures were taken 15 to 30 min after the sheep had been
moved quietly from the pastures to yards. A 30 sec maximum, small bulb
clinical thermometer was used.
Estimates of pasture yield were made each week when intake was
measured, using either an electronic capacitance meter or a visual estimation
technique (Campbell and Arnold, 1973).
Climatic data were obtained either on the experimental sites, or at
standard meteorological stations about 1 km from the sites. The record of
wind was the daily reading of the speed at 09.00 hr except for Experiment 3
for which 24-hr wind runs were measured; the 09.00 hr wind speed and the
24-hr wind run were linearly related (r = 0-9) with a slope of 10 and an
intercept of zero.
 FOOD INTAKE AND GRAZING BEHAVIOUR OF SHEEP 347

RESULTS

Weather conditions during Experiments 2 to 7

Mean minimum and maximum temperatures at Hamilton (Experiments 3
and 4) were about 3°C lower than at Bakers Hill (Experiments 2, 5, 6 and 7)
(see Table 2). Wind speeds were higher at Hamilton on average. The
number of days on which more than 1 mm of rain fell in the first 10 days
after shearing varied from 0 to 5 for Experiments 2 to 7 (Table 2).

Differences in live weight and pasture conditions

In Experiment 1, the groups of thin and fat sheep differed by 24 kg in
live weight but in Experiments 2 to 4 the difference ranged from 10 to 13 kg.
High and low pasture levels differed by about 2000 kg/ha in all experiments
(Table 1).

Experiment 1
There were no treatment interactions between body condition and
pasture level. Thin sheep ate 3 1 % more DOM (P<0-01) and 117% rflore
per kg live weight (P< 0-001) (Table 3). The higher intake was achieved
by the sheep eating about 25 % faster (P < 0-05).

TABLE 3
Results for Experiment 1 at Canberra, ACT (pooled over two periods)
Treatment
r————* , OM DOM DOM intake/ Grazing OM intake
Body intake intake kg live weight time /hr of
condition Pasture (g/day) (g/day) (g/day) (hr/day) grazing (g)
Fat Abundant 600 471 7-7 5-7 109
Thin Abundant 711 558 15-0 5-6 132
Fat Scant 332 270 4-4 7-2 46
Thin Scant 510 415 11-1 8-3 61
SE of treatment means 119 92 3-3 1-1 20!

Food intake was less on scant pasture (P < 0-001) because, despite 39 %
more time spent grazing (P< 0-001), intake per hour of grazing was only
44% of that on abundant pasture (P< 0-001).

Experiment 2
Although there were some interactions with time in the intake para-
meters these were small and will not be considered. The intake of organic
matter, however, was significantly (P< 0-01) higher in weeks 3 and 4 than
in weeks 1 and 2. There were no interactions between the main treatments.
Thin sheep ate 33% more DOM (P< 0-001) and 55% more per kg live
weight than fat sheep (Table 4). Neither amount of fleece nor pasture level
had an effect on DOM intake.
Thin sheep grazed for longer than fat sheep on abundant pasture (P < 0001)
but not on scant pasture. The trend for rate of eating was for thin sheep
to eat faster on scant pasture but not on abundant pasture, but this was not
significant.
 348 ARNOLD AND BIRRELL

Over the 4 weeks, fat sheep gained a similar amount of weight whether
wooled or shorn, but thin sheep gained less weight when shorn (Figure 1).
Rectal temperatures were similar for all groups.

TABLE 4
Results for Experiment 2 at Bakers Hill, WA
Treatment
DOM intake/kg
Body OM intake DOM intake live weight Grazing OM intake/hr
condition Fleece Pasture (g/day) (g/day) (g/day) time (hr/day) of grazing (g)
Fat Wool Abundant 745 576 11-0 5-2 136
Thin Wool Abundant 1157 901 19-5 7-9 153
Fat Shorn Abundant 896 681 130 4-8 160
Thin Shorn Abundant 1118 845 18-3 6-9 169
Fat Wool Scant 734 585 111 7-7 86
Thin Wool Scant 1001 790 17-6 7-7 134
Fat Shorn Scant 819 642 12-2 6-9 106
Thin Shorn Scant 976 771 17-6 7-9 118
SE of mean 230 177 3-4 1-5 52

60 f EXPERIMENT 2 EXPERIMENT 3

. -o
45

40

35

L
30 -
20

DAYS FROM SHEARING

FIG. 1. Fleece free live weights of shorn and unshorn sheep in Experiments 2 and 3.
(Live weights of sheep with wool have been adjusted to fleece-free live weights by
subtracting the fleece weights obtained from the shorn sheep in the same condition
classes.) • • fat+wool, • # thin+wool, O——O fat—wool, O O
thin—wool.

Experiment 3
Unfortunately, faecal samples were lost so could not be analysed for
nitrogen and intakes thereby estimated. Under similar pasture conditions
faecal dry matter outputs (DMO) generally reflect relative differences in
intake, and this was the case here because the 'scant' pasture was not par-
ticularly short of feed and the digestibility of pasture eaten under both
 FOOD INTAKE AND GRAZING BEHAVIOUR OF SHEEP 349
pasture conditions was probably similar. DMO was 25 % higher (P < 0-001)
for thin sheep than for fat sheep, 10% higher (P<0-05) for shorn than
unshorn sheep and 9 % higher (P < 0-05) on the abundant than on the scant
pasture (Table 5). The same pattern held for DMO per kg live weight,
being 45 % higher (P < 0-001) for thin sheep, 11 % higher (P < 0-001) for shorn
sheep and 8% higher (P<0-01) on abundant pasture.

TABLE 5
Results for Experiment 3 at Hamilton, Victoria
Treatment
P I T"\\>r
JraeCal UJVL
Body Faecal DM output/kg Grazing output/hr of
condition Fleece Pasture output (g) live weight (g) time (hr) grazing (g)
Fat Wool Abundant 302 56 7-4 42
Thin Wool Abundant 389 80 7-6 51
Fat Shorn Abundant 327 60 6-4 51
Thin Shorn Abundant 404 87 6-1 66
Fat Wool Scant 312 57 6-8 46
Thin Wool Scant 409 86 7-5 54
Fat Shorn Scant 381 70 61 62
Thin Shorn Scant 442 96 6-6 67
SE of treatment means 30 16 1-3 12

Grazing times were similar for both pasture levels and for fat and thin
sheep, but shorn sheep spent 15% less time grazing (P<0-05). Thin sheep,
however, excreted 20 % more dry matter per hr of grazing (P < 0-01) than
fat sheep and shorn sheep 29% more per hr (P<0-01) than wooled sheep.

TABLE 6
Rectal temperatures CO in Experiments 3 and 4 at Hamilton, Victoria
Days after shearing
Treatment Experiment 3 Experiment 4
»
Body
condition Fleece Pasture 2 3 5 9 41 8 9 16 19 46
Fat Wool Abundant 40-2 40-7 40-4 400 39-9 40-4 40-2 40-6 39-9 40-3
Thin Wool Abundant 40-4 40-2 400 40-1 39-8 40-4 40-4 40-5 39-9 40-3
Fat Shorn Abundant 39-7 39-5 400 39-8 400 39-9 39-6 39-8 39-8 400
Thin Shorn Abundant 391 39-0 39-9 39-8 39-9 ' 39-5 39-5 39-5 39-7 400
Fat Wool Scant 40-4 40-2 400 40-1 400 40-3 40-3 39-9 39-9 401
Thin Wool Scant 400 40-1 40-3 39-9 40-2 39-9 39-9 40-2 39-8 40-3
Fat Shorn Scant 39-8 39-6 39-6 39-6 39-8 39-3 39-8 40-5 39-8 39-8
Thin Shorn Scant 38-8 38-6 39-4 39-5 39-8 39-4 39-4 39-4 39-5 39-8
SE of treatment means 0-24 0-30 0-21 0-23 0-25 0-20 0-34 0-30 0-25 0-2

Thin sheep gained more weight than fat sheep (10-0 v. 4-2 kg, P<0-01)
and shorn sheep gained less weight than wooled sheep (5-0 v. 9-2 kg, P<0-01)
(Figure 1). Rectal temperatures were 0-3 to 1 -4°C lower in shorn sheep up
to day 9 after shearing; thin, shorn sheep had lower temperatures than fat,
shorn sheep until day 5 (Table 6). Wind speed varied from 1 -7 to 6-3 m sec - i .

Experiment 4
Intake increased over time for thin sheep but not for fat sheep, and for
sheep on scant pasture but not for sheep on abundant pasture. These inter-
actions were significant (P< 0-001).
 350 ARNOLD AND BIRRELL

The three measurements of intake (organic matter, DOM and DOM/kg

wool free live weight) showed the same effects (Table 7). Thin sheep ate
more than fat sheep (P< 0-001), and shorn sheep ate more than wooled
sheep (P<0001). The proportionate increase for thin sheep, however, was
greater on scant pasture, and that for shorn sheep was proportionately
greater on abundant pasture. Shorn sheep had a proportionately higher
intake than wooled sheep when they were thin than when fat.
Thin sheep had a longer grazing time than fat sheep on abundant pasture,
but not on scant pasture (P<005); likewise, thin shorn sheep grazed for
longer than wooled sheep only on abundant pasture (P < 0-01).
Organic matter intake per hr of grazing was similar for thin and fat
sheep on abundant pasture, but thin sheep ate much faster than fat sheep
on scant pasture. Shorn sheep consumed organic matter faster than wooled
sheep.

TABLE 7
Results for Experiment 4 at Hamilton, Victt
Treatment
DOM intake/kg
Body OM intake DOM intake live weight Grazing time OM intake/hr
condition Fleece Pasture (g/day) (g/day) (g/day) (hr/day) (g)
Fat Wool Abundant 973 755 16-6 7-7 128
Thin Wool Abundant 1109 876 20-9 91 123
Fat Shorn Abundant 1366 1094 22-8 7-5 178
Thin Shorn Abundant 1350 1086 26-8 7-7 174
Fat Wool Scant 573 428 9-9 9-7 59
Thin Wool Scant 709 529 14-3 101 67
Fat Shorn Scant 597 436 10-9 9-0 64
Thin Shorn Scant 1008 777 22-1 8-2 119
SE of treatment means 244 193 41 0-9 28

As in 1968, thin sheep gained more weight than fat sheep (7-0 v. 5-4 kg,
P<0-05) and shorn sheep gained less weight than wooled sheep (5-6 v. 6-7
kg, P<0-05). Rectal temperatures were lower in shorn sheep up to day 26
from shearing and were lowest in thin shorn sheep for most of the period of
measurement (Table 6).

Experiments 5, 6 and 7
Fleece lengths of shorn sheep were 10 mm in Experiment 5 and 3 mm
in Experiment 7 compared with 41 mm and 89 mm respectively for wooled
sheep. Rain fell every day for the first week after shearing in Experiments 6
and 7 but not at all in this period in Experiment 5. In none of the experi-
ments did intake differ between shorn and unshorn sheep. Rectal tempera-
tures never differed by more than 0-1CC between shorn and unshorn sheep.

DISCUSSION
Body condition
Adult sheep that were in poor body condition increased their food intake
both absolutely and per kg live weight, and did this on both short and
on abundant pasture. The higher food intakes were achieved by thin sheep
eating faster in most situations. In two of four experiments, however, thin
sheep on abundant pasture grazed for longer than fat sheep but did not eat
faster than them.
 FOOD INTAKE AND GRAZING BEHAVIOUR OF SHEEP 351

These results differ from those of Hutchinson and McRae (1969) and
Donnelly, Davidson and Freer (1974) who found that thin sheep ate more
per unit live weight but not absolutely more than fat sheep; the differences
in live weight between thin and fat sheep were similar in all three studies.
The present food intake results agree with those of Allden and Young (1964)
and Langlands (1968) who subjected two groups of Merino wethers of from
3 mo to 7 yr of age to 3 months' differential nutrition. With one exception,
however, the differences in live weight were confounded with a different
rate of live-weight change prior to the studies. Hutchinson and McRae
(1969) have shown that feed intake is influenced by previous rate of live-
weight loss.
The differences in grazing time and rate of intake fit a pattern found in
earlier work. Arnold (1975) showed that pregnant and lactating sheep
achieved higher intakes than dry sheep primarily by increasing rate of intake,
and only on abundant pasture did they graze for longer than dry sheep.

Cold stress
Shorn sheep had significantly higher feed intakes than wooled sheep only
at Hamilton (Experiments 3 and 4) and they achieved higher intakes by
eating faster. The differences in intake were greatest in Experiment 4 and
similar to those obtained by Hutchinson and McRae (1969).
There are several factors other than climatic ones that can affect response
to cold stress but the influence of these factors cannot be assessed in the
present experiments. Since the experiments were all conducted in a similar
way and at a similar time of year, however, it is thought that they may have
acted in a similar way in each experiment. If this is so, the major climatic
differences between Bakers Hill and Hamilton were in temperature and in
wind speed. The number of wet days in the week after shearing was similar
in Experiments 3 to 7.
At Hamilton the shorn sheep had lower rectal temperatures than wooled
sheep and ate more; the mean ambient temperature was 8-3°C compared
with 10-8°C at Bakers Hill and the wind speeds 4-7 and 2-9 m/sec respectively.
The small difference in temperature could have been sufficient to cause cold
stress at Hamilton. The relationship between ambient temperature and heat
production in shorn sheep is exponential (Armstrong et ai, 1959) and the
difference in heat production would be about 10% due to temperature with
an increase due to wind.
A feature of the Hamilton results was the lower rectal temperatures in
thin shorn sheep than in fat shorn sheep in the first week after shearing. A
similar result was obtained under much colder conditions at Canberra
(Arnold, 1976). Both results indicate that tissue insulation was important.
There are no other such reports although Joyce and Blaxter (1964) suggested
that this factor contributed to differences in response to cold stress by
different breeds of sheep. No increases in intake (as indicated by DMO)
due to cold stress occurred in the first 2 weeks after shearing in Experiment 3,
which agrees with other studies reported (e.g. Wheeler et ah, 1963). In
Experiment 4, however, DOM intake increased by 17% in the first week
and by 35 % in week 2 where it remained throughout the measurement period
of 5 weeks. These increases compare with values of 30 % and 59 % reported
by Hutchinson and McRae (1969).
24/3—E
352 ARNOLD AND BIRRELL

If shorn sheep lose more weight or gain less weight than unshorn sheep
this indicates that they cannot fully adjust energy intake to meet the extra
energy demand to keep warm. Wodzicka-Tomaszewska (1964) reported
that weight changes were similar in shorn and unshorn sheep, but in our
studies at lower mean ambient temperatures, shorn sheep failed to make up
the energy deficit due to cold at Hamilton. Theoretically, the deficit would
be greater at lower temperatures, and would be influenced by both wind and
rain but the basic physiological work so far is inadequate to describe precisely
when sheep will be under cold stress.

ACKNOWLEDGEMENTS
It is a pleasure to acknowledge the skilled technical assistance of Messrs I. G. Bush,
L. Osborne and S. R. Wallace. Miss E. Rickman and Mrs G. Knight, Division of Mathe-
matical Statistics, CSIRO, did most of the statistical analyses.

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{Received7 July 1976)