Applied Animal Ethology, 4 (1978) 301-313 301

o Elsevier Scientific Publishing Company, Amsterdam - Printed in The Netherlands

AN ANALYSIS OF INGESTIVE BEHAVIOUR AND ACTIVITY OF CATTLE

UNDER FIELD CONDITIONS

Y. RUCKEBUSCH and L. BUENO

Laboratoire de Physiologie et Pharmacodynamie, Ecole Nationale VgtGrinaire - 31076
Toulouse Gdex (France)
(Received 17 October 1977)

ABSTRACT

Ruckebusch, Y. and Bueno, L., 1978. An analysis of ingestive behaviour and activity of
cattle under field conditions. Appl. Anim. Ethol., 4: 301-313.

A modified vibracorder was used to record the time spent by cattle in grazing and
ruminating under field conditions for 24-h periods in autumn and winter during 3 con-
secutive years. An unmodified vibracorder device was also used to obtain concurrent in-
formation about walking, loafing and resting time.
Three major influences were examined: the size of the field, the seasonal changes of
grass availability and the weather. The time spent walking was correlated (r = 0.92) to that
of grazing and the time of their onset (and reset) was correlated (r = 0.89) to the correspond-
ing times of sunrise (and sunset). An excess of time spent walking compared with that of
grazing was found in both the 0.5 and 3 ha fields but the circadian pattern of grazing was
disrupted when cattle were restricted to the small area. When the cows were turned into
the large field both walking and loafing were transiently increased due to the novelty of
the environment. Then the time spent walking but not loafing decreased from November
to March in relation with the lower amount of grass available. A further reduction in time
spent walking was observed for windy weather.
It is concluded that the size of the field and weather interact with the circadian patterns
of both walking and grazing while the seasonal changes in ingestive behaviour are due to the
amount and quality of grass available.

INTRODUCTION

The ingestive behaviour and particularly the grazing time of cattle under
field conditions has received much attention (Tribe, 1950; Hughes and Reid,
1951; Corbett, 1953) and Hafez et al. (1969) provide a general review of eating
and related activities. However, most of the observations have been conducted
over short periods of a few days and the time spent in a given behaviour was
usually recorded at 10 (Czako et al., 1969) or 15 min intervals (Gary et al.,
1970) on the assumption that the behaviour of the animals observed remained
the same until the next observation. Castle et al. (1950) and Bubenick (1960)
used infrared equipment, to allow night observations.
More detailed information was obtained by Canaway et al. (1955). Stobbs
302

(1970), and more recently Stricklin et al. (1976), recorded the motor patterns
associated with ingestive behaviour in cattle using a mechanical clock recorder
(vibracorder) operated by a pendulum activated by the movements of the
animal.
Ingestive behaviour itself has been accurately recorded by monitoring jaw
movements in stall-fed ruminants using pneumatic systems (Duckworth and
Shirlaw, 1955; Ohashi and Goto, 1968; Balch, 1971; Ruckebusch and Bueno,
1972) or by electric monitoring (Law and Sudweeks, 1975). Automatic
measurement of jaw movements during grazing and rumination has been ob-
tained from cattle in the field using either telemetric equipment, numerical
recorder assemblies attached to the halter (Stobbs and Cowper, 1972) or a
modified vibracorder operated by a pneumatic system (Ruckebusch and
Bueno, 1973).
In a preliminary field study of ingestive behaviour, it was found that time
spent grazing could be overestimated when measured by weekly charts of
somatic (vibracorder) activity compared to daily charts of jaw movements by
a modified vibracorder. Overestimates occurred particularly when the herbage
was scant even when only the periods of activity indicated by continuous
bands adjoining strokes were considered (Ruckebusch and Bueno, 1973).
This study was carried out to determine the changes in circadian patterns of
ingestion and activity when herbage intake was restricted by (1) providing a
limited grazing area and (2) by normal seasonal changes.
Since the pressure balloon located in the submandibular space and connected
to the modified vibracorder for recording mastication movements could also
indicate the periods during which the head is fully extended on the ground in
an attitude of complete relaxation of the neck musculature, usually occurring
during deep sleep in a recumbent animal (Ruckebusch, 1975), these values will
be reported as additional resting behaviour data.

METHODS AND MATERIALS

Apparatus

The recorders (type TFW 3/8 and TFW 24/8 Kienzle, Villingen, W. Germany)
used consisted of a heavy pendulum connected to a stylus (vertical movement
2 mm) which registered vibrations or oscillations on a waxed paper recording
disc (12.5 mm diameter). The disc driven by a clock mechanism rotated at an
angular speed of 120 degrees every hour for TFW 3/8 or every 8 h for TWF
24/8. By means of a free cam which rotated slowly once in 1 or 8 days the
pendulum and stylus were gradually lowered and thus a continuous recording
of events was obtained as 8 concentric, slightly-diminishing, spiral traces. To
record the tilting and swinging movements during the activity of the animal
throughout the 24 h, the vibracorder TFW 24/8 was attached to the neck.
In order to record jaw movements, the counterweight of the pendulum was
removed near to the attachment of the pull-back spring of a Vibracorder TFW
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3/8. The remaining part of the pendulum was lightened by perforations and a
thin steel plate was soldered on to the shortened pendulum. A glass syringe
was inserted through a hole in the side of the recorder so that the plunger
butted on the thin steel plate. The syringe casing was fixed in place by medical
silastic (Fig. 1). A length of rubber tubing was used to connect the free end of
the syringe to a thick rubber bulb (125 ml) mounted in the submandibular
space. This was attached to the connecting piece between the nose band and
throat strap of a leather head stall. The modified recorder was placed on the
animal’s back by two girth straps (Fig. 2). About 10 g of silica gel was placed
in the bulb to avoid water condensation in the syringe which might block the
pneumatic transmission of the jaw movements at night. At times, a third
recorder was placed on the other side of the animal’s withers and connected
by a rhythm divider circuit to the afore-mentioned device. It was electromag-
netically activated to record 1 stroke per 20 jaw movements registered.

Fig. 1. A diagram showing the modification of the vibracorder. (A), the counter-weight of
the pendulum is removed and a steelplate is soldered (a), the plunger of a reversed syringe
(b) is butted on the shortened pendulum which is lightened by perforations (c). (B), chart
recording obtained under field conditions covering a period of 24 h. Three behavioural
patterns of ingestive behaviour are shown on the chart: grazing (l), ruminating (2) and
resting (3). Below, the magnification of the trace also shows a 5.5 min period during which
the head rests fully on the ground (*). This particular interval occurred at the end of a
period of rumination.
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Record analysis

Ingestive behaviour was of two types; (a) grazing, when the trace showed a
continuous band of coalescent strokes interrupted by pauses of less than
0.25 min and (b) ruminating, when regular well-defined bands of 0.8 to 1.1 min
duration were separated by intervals of about 5 s (see Fig. 1). The periods of
rest were calculated by subtraction and those of deep sleep were obtained from
the uninterrupted period when deflection of the stylus reached its highest
level corresponding to continuous pressure on the bulb when the animal’s head
was resting with its full weight on the ground.
Walking, loafing and resting times were identified from the frequency of
animal movements indicated by the vibracorder attached to the neck. General
“activity” behaviour was characterized as walking when the vertical strokes
were not separated (more than 20 per min for external spiral traces, lo-15
per min for internal spiral traces), as loafing when the strokes were irregular,
but with no coalescence (less than lo-15 per min) and as resting, either
standing or recumbent, when the strokes occurred with a frequency of at
the most one per min (see Fig. 3B).
Each cow was trained to come to a part of the field to receive about 200 g
of grass cubes each morning, and while eating the cubes the chart was removed,

Fig. 2. An Aubrac cow during grazing with a recording unit attached on its right side. The
vibracorder may be connected to a rhythm divider which activates electromagnetically a
second recorder on the left side, after each 20 jaw movements.
 Resting area

OOrn

v V
_~ -__
3m
V

v V V v
V V V 1

3”tTl

150m

Fig. 3. A diagram showing the two field areas of 5000 (A) and 30 000 (B) square meters
allowed for grazing. The corresponding recording chart of activity, from each area (A and
B) covers the period of 6 days on the 3rd and 6th week outdoors. Three patterns of activity
are shown: walking, loafing and lying down. Grazing occurred from midnight until mid-
day on the small area (A) and the cow was at the resting area during the whole afternoon.
On the large area (B), grazing occurred during the day and was seen in 2 major periods
during the night. The mean values of walking are 670 min per day in A (range: 480 and
780) and 790 min in B (range: 660 and 930). The head is resting on the ground for 50 min
(25-80) per day in A and 30 min (20-40) in B.
306

a new recording disc was inserted and the face cam was returned to its starting
position.

Experiment 1

TFW 3/S modified recorders were used on three cows of the Aubrac breed,
each weighing approximately 500 kg, and 5, 7 and 10 years old, termed G, U
and J, respectively. Recordings began in September-October 1974 on
animals which had earlier been fed on hay in a small barn (6 X 4 m), then
turned out for 3 weeks into a 0.5 ha field of rye grass (5-8 cm high) and
finally placed in a 3 ha field for 4 weeks (Fig. 3). Clement weather with sun-
shine was the rule, except for 48 h of continuous rain when they were in the
large field.

Experiment 2

Recordings of both activity and ingestive behaviour were carried out in the
same 3 cows. The animals were turned into the 3 ha field from May to June
where the grass was 10-12 cm in length. They were then stall-fed on hay
and turned out again in the 3 ha field from October 1975 to March 1976. The
height of grass available decreased from 10 to 5 cm, during this period, and
only the daily activity was recorded. Hay was available ad libitum at the
resting area. An analysis of the charts was performed as for Experiment 1,
except that a more accurate indication of the general activity was obtained
by using a TFW 3/S unmodified vibracorder instead of the TFW 8/24. During
the 6 months over the winter period, the mean amount of walking, loafing and
resting per 24 h was grouped according the weather, i.e. wet or windy. In both
cases a factorial analysis of variance was carried out.

RESULTS

Walking versus grazing behaviour

A comparison of the percentage of time spent grazing and walking in

September-October 1974 showed a good correlation (r = 0.92) between
these two components of ingestive behaviour; that of walking exceeding that
of grazing by lo-15%. Two major periods of grazing occurred at sunrise and
sunset for fine weather while cows were on the large area. These periods were
no longer separated for the 48 consecutive hours of rain and periods of
rumination were then distributed throughout the day (Fig. 4). However, the
mean values of both walking and loafing remained similar whatever the weather,
the major feature being longer periods of resting for fine weather.
The total time spent either walking or grazing in May 1975 took, on average,
30% of the day when grazing rye grass of lo-12 cm in length in the spring
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Minutes
Fine weather
60,

O_
OI
; 20_

; 40_
!s
Raining

O_

.E 20-
“0
5 40_
E
?? cntghtlme - .
z

I I I I
24 08 16 24 08 16 24 HOURS

Fig. 4. Hourly distribution patterns of grazing and rumination in cow G in the large area
during 48 h of fine weather (24°C) and subsequent 48 h of wet weather (22°C).

TABLE I

Time spent on ingestive behaviour and activity by the cows (May 1975). Mean 2 SD for
10 days. In brackets, mean percentage per 24 h

General activity

Walking Loafing Resting

488 ? 18.5 359 f 58 593 ? 118

(33.8) (24.9) (41.2)

Ingestive behaviour

Grazing’ Ruminating* Non chewing

activity

447 i 51 284 + 69 708 + 115

(31.1) (19.8) (49.1)

‘Jaw movement recorder shows that cows were not always harvesting food actively during
walking.
2Ruminating occurred when the cows were at rest either standing or recumbent.
308

time (Table I). The relationship between grazing (x) and walking (y) times in
minutes was
y = (0.96 + 0.04)~ + (56.8 f 17.4) (r = 0.93, n = 20) for cow G
y = (1.16 f 0.13)~ - (13.2 + 18.4) (r = 0.87, n = 20) for cow U
y = (0.95 + 0.06)~ + (68.9 + 26.1) (r = 0.96, n = 17) for cow J
Again the time spent walking exceeded that of grazing and rumination was
found to occur for 50% of the time when the animal was resting, i.e. when no
activation of the pendulum of the vibracorder occurred. A one-month com-
parison between ruminating and resting time showed a significant correlation
(r = 0.82) confirming that in the field, rumination occurred mostly when the
animal was lying down. The mean duration of resting the head on the ground
was 35 min per 24 h in 5-7 periods of 5-7 min each.

Small versus large field conditions

The mean walking time was slightly higher when the animals were on the
large area, as was the grazing time determined from jaw movements (Table II).
The time spent walking was always greater than the time spent grazing, although
a good daily correlation (r = 0.83) was found in all 3 cows between grazing
and walking which was similar to that observed on a larger area. A typical
circadian pattern of grazing in 2 major periods during daytime (75%), shown
in the large area, was not present for grazing on the small area. In this latter
case, a period of grazing was regularly found around midnight which continued
until midmorning.

Seasonal changes

From October 1975 to March 1976, the morning period of walking started
14 + 13 min after sunrise. The last period of walking continued for 87 f 17 min
beyond sunset (Fig. 5). For two cows, G and U, a 6-month analysis showed
two linear correlations: firstly between the mean weekly times of the onset
of grazing (y) and of sunrise (x), the mean equation being y = 0.64 x + 2.13
(r = 0.87, IZ= 22); secondly in the evening between the end of grazing (y’) and
sunset (x’), y’ = 0.39 x’ + 13.12 (r = 0.67, n = 22). Walking time varied both
with changes in the weather and with the change of season. During winter
days values of 444 + 67 min are shown compared with 531 + 88 min for sun-
shine or wet days and also the end of the winter period where only a small
amount of grass was available (Table III). Loafing values were the lowest in
November and December, i.e. for the period of shortest day length. Except
for October, when loafing averaged 6-7 h per day, the mean duration of
loafing was 2-3 h and the duration of resting increased considerably during
the whole winter period.
 309

TABLE II

A comparison of the time spent on ingestive behaviour and activity by three cows in the
small and large fields (September-October, 1974). Mean * SD for 12 consecutive days
recorded after the 2nd week outside the barn. In brackets, mean percentage per 24 h

Cow G cow u Cow J

Day Night Day Night Day Night

8-20 h 20-8 h 8-20 h 20-8 h S-20 h 20-8 h

Small area
General activity
Walking’ 305 228 290 260 360 260
f 54 f 60 + 60 f 40 t 80 * 70
(37.0) (38.1) (43.0)
Loafing 301 105 350 89 259 121
t 34 f 31 * 61 + 27 f 41 + 81
(28.1) (30.5) (25.7)

Ingestive behaviour
Grazing’ 265 121 247 140 290 160
f 91 f 21 1 80 t 31 + 60 f 47
(26.8) (26.8) (24.3)
Ruminating2 89 272 101 204 87 240
f 72 f 67 + 61 % 51 f 21 f 37
(25.0) (21.1) (22.7)

Large area3
General activity
Walking 460 188 480 204 540 190
f 87 f 40 f 81 + 36 f 78 + 27
(40.8) (47.5) (50.6)
Loafing 200 180 150 170 201 121
+ 37 f 27 + 30 f 41 f 31 f 21
(26.3) (22.2) (22.3)

Ingestive behaviour
Grazing 325 83 350 70 409 89
+ 59 + 68 + 60 f 37 f 49 t- 51
(27.9) (29.1) (34.5)
Ruminating 64 217 104 307 79 180
+ 41 f 35 + 41 f 17 * 17 * 29
(18.9) (28.5) (17.9)

‘Variance analysis of walking and grazing showed no significant differences (F = 3.04)

between cows, but was significant (F = 208, P < 0.01) for day versus night-time.
‘Variance analysis of rumination for day versus night-time (F = 39, P G 0.05).
3WaIking but not loafing was significantly increased in the larger area (F = 22.9,P Q 0.05).
TABLE III

Monthly variations in time spent walking and loafing by 3 cows. Mean + SD for 30 days per month in three subjects.
Resting is indicated as its mean percentage per 24 h

Weather’
Fine Windy
_
Walking Loafing Resting Walking Loafing Resting Walking Loafing Resting
(min) (min) (%) (min) (min) (%) (min) (min) (%)

October* 128 * 87 460 + 84 17 768 f 52 366 -r 38 22 666 ? 31 514 + 27 19

November 548 ? 76 192 -t 104 49 484 + 49 113 ? 81 59 475 + 61 91* 30 61
December 554 * 81 102 + 51 55 521 * 61 115 i 19 58 512 f 23 110 + 29 58
January 518 it 58 157 i 48 49 475 f 69 146 2 44 57 312 t 66 214 + 95 53
February 469 F 62 194 i 64 54 415 r 59 201 f 37 57 342 ? 46 184 i 26 63
March 471 + 87 213 i 81 53 452 + 41 202 + 63 55 358 * 43 195 + 44 52

m + SD 558 L 88 220 ? 124 520 i 61 190 ?C86 444 i 61 218 i 139

~___ - ___ ~~ _ __ ~~ ~~_~
Analysis of variance showed that the activity was correlated with the weather’ F = 9.86,P G 0.01,and with the month’
F = 19.0,Ps 0.001.
 311

hours

act. 1 nov. 1 dec. 1 Jan. 1 feb. 1 march

Fig. 5. Mean weekly times for sunrise and sunset from October 1975 to March 1976 (top
panel) and the beginning and end of the daily grazing periods (bottom panel). The times
are based on the mean value + SD for the three subjects.

DISCUSSION

The continuous objective recording of animal behaviour under field con-

ditions requires low-cost, light-weight equipment. If an electromagnetically
operated counter can give the total number of certain movements, the dis-
tribution of these activities over a 24-h period requires a recording on moving
paper. The unit described here permits a record of the time spent by an
animal in grazing, ruminating and resting, as the recording disc is driven by
a clock mechanism at a speed which permits discrimination between these
different states. The modifications that the Kienzle-Recorder TFW 3/8
requires to record jaw movements are simple and do not interfere with the
rubber insulation between the casing and the lid necessary for protection
against humidity. The difficulties normally encountered in recording jaw
movements by various types of contact switches in the submandibular space
are avoided.
The behaviour recorded for cattle in the field confirmed previously gathered
data and the analysis of variance showed no major differences between the
3 cows under the experimental situations. In fact, the cows used were particular
312

ly quiet animals which were chosen for the absence of individual differences
when they were stall-fed. A striking feature is that the jaw movement recorder
showed that cows were not always actively harvesting food when walking. An
excess of time spent walking compared with that spent grazing was found in
September-October 1974 and to a lesser extent in May 1975. The larger
amount of grass available in the spring, as well as a lower rate of defoliation
(Chacon and Stobbs, 1976), may account for the difference.
The patterns of grazing activity were disrupted when cows were restricted
to a small area. In this case, the nocturnal phase of grazing already observed
by Stobbs (1970) was prolonged until the morning (see Fig. 3), and the per-
centage of grazing which occurred during the night rose beyond the 17%
reported by Gary et al. (1970). As expected, the time spent walking per 24 h
was increased when animals were in the larger area from October 1975 to
March 1976, especially for the first month. This phenomenon was also
recorded for loafing so that the time spent at rest did not exceed 25% per
day in October and was nearly doubled for the subsequent period. This may
be explained as an effect of the novelty of the environment since the time
required is about the same as that observed in sleep studies for stability of
the hypnogram (Ruckebusch, 1975).
Another point of interest is that the major periods of grazing occurring
at sunrise and sunset were not clearly delineated during wet weather (Fig. 4).
In this case, the animals seemed likely to avoid long periods of inactivity
during the daytime and even during night-time. Under natural conditions,
activity rhythms usually become synchronized with the earth’s rotational
movement which give rise to certain periodic environmental changes, illumina-
tion being one of the most important. Many animals customarily begin or
cease activity during twilight; simulated twilight can influence the activity of
numerous captive mammals (Kavanau and Peters, 1976). Although it was
difficult to determine the precise influence of twilight in field studies because
of other confounding influences, such as the degree of cloud cover, temperature,
and food availability, the effect of daylight and darkness was readily apparent.
At dawn, or soon after, the cows began eating. They continued throughout
the day beyond darkness until dusk. Decrease in food availability and reduced
protein digestibility may account for the earlier onset of activity in autumn.
When leaf yield was low in winter, i.e. from November to March, the effect
of rain, and to a greater extent the effect of wind, became more evident.
Finally, the periodicity of grazing of cows is highly predictable within a
season. The unmodified TFW 3/B and the modified Vibracorder prove to be
useful instruments both in long term investigation of eating and in related ac-
tivities. These interact with factors such as weather, available forage, novelty
of the environment and size of the available pasture area.

ACKNOWLEDGEMENTS

We are grateful to Dr. R. Kilgour for his assistance in preparation and

criticism of the manuscript.
 313

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