Download as pdf or txt
Download as pdf or txt
You are on page 1of 8

APPLIED ANIMAL

BEHAVIOUR
SCIENCE
EJSEVIER Applied Animal Behaviour Science 54 (1997) 89-96

Activity bout criteria for grazing dairy cows


A.J. Rook *, C.A. Huckle
lrrstirurr qf Grassland arzd Em~ironmmr Research. North Woke. Okhampton. Demn EX20 ZSB. UK

4ccepted 20 March I997

Abstract

Data on the eating, ruminating and idling activity of grazing dairy cows recorded to a
resolution of 1 min were used to study the choice of activity bout criteria and the implications
of
this choice. An inter-bout length of 5 min was found to suitable for all three activity categories
across a range of treatments, at different times of the day and for all animals. After smoothing
using this criterion bouts contained proportionally up to 0.069 of other activities for eating, 0.076
for ruminating and 0.054 for idling. The proportion of intra-meal intervals decreased slightly
during the main evening meal and increased during meals taken at night. It is concluded that the
intensity of bouts should be reported as well as their number and duration and that total activity
times should be calculated from unsmoothed data. 0 1997 Elsevier Science B.V.

Keywords: Cattle-feeding and nutrition; Grazing behaviour: Bout criteria

1. Introduction

It is well recognised that animals perform activities in discrete bouts rather than at
random (e.g. Rook and Penning, 1991). However, the definition of a bout is not trivial.
because some breaks in the activity pattern are so short as to be regarded as intra-bout
intervals, while others are of sufficient length to be regarded as true inter-bout intervals.
In most situations there is not a priori basis for assigning intervals to one or other of
these categories. It is thus usual practice to derive the bout criterion empirically from the
data.
A simple and widely used method of determining the bout criterion (Forbes et al.,
1986) is to regress the frequency of occurrence of intervals of different duration on the
duration, using a piecewise linear or ‘broken stick’ model. This model is fitted with a

* Corresponding author. Tel.: + 44-1837-82558; fax: + 44-1837-82139: e-mail: aj.rook@bbsrc.ac.uk

016%1591/97/$17.00 Q 1997 Elsevier Science B.V. All rights reserved.


/‘If SOl68-1591(97)00065-8
90 A.J. Rook, C.A. Huckle/Applied Animal Behauiour Science 54 (1997) 89-96

number of different breakpoints and the breakpoint of the best fitting model is chosen as
the bout criterion. Sibly et al. (1990) suggested a more complex method, based on the
concept that there are two independent Poisson processes at work, a fast process
generating within-bout intervals and a slow process generating between meal intervals.
Berdoy (1993) subsequently generalised this concept to a three process model, in which
bouts are themselves clustered.
Discussion of bout criteria has generally been in the context of definition of meals
and has considered only two states, eating and not eating. However, most animal
behaviour studies make use of more complex time budgets. This introduces the
additional problem of how best to smooth the data to remove small intra-bout intervals
when the activity occurring in these intervals may itself be of interest. Further, once
such smoothing has taken place, the consequent properties of the ‘bouts’ so created must
be studied in order to ensure that the term is not used in a misleading way. In this paper
we consider the issues outlined above using the grazing behaviour of dairy cows as an
example.

2. Materials and methods

This study used data from an experiment (Wilkins et al., 1994) that was designed to
investigate the effect of concentrate supplementation and sward clover content on milk
production and grazing behaviour by spring-calving, Holstein-Friesian, dairy cows
continuously stocked on perennial ryegrass (L&urn perenne L.1 - white clover
(Trifdium repens L.) swards that were maintained at a sward height of 60 mm, as
measured by a rising plate meter (Holmes, 1974).
Grazing behaviour measurements were made at only one sward clover content (0.20
of total dry matter (DM) as clover). Three levels of concentrate supplementation (0, 2
and 4 kg/day) were offered in equal feeds at each of two milkings. The concentrate
contained 180 g/kg DM crude protein and 12.8 MJ/kg DM metabolizable energy.
Animals from all concentrate treatments grazed together in the same paddock. Competi-
tion and social facilitation may, therefore, have affected the results obtained (Rook and
Huckle, 1996). Concentrate level had no significant effect on milk or milk constituent
yields.
Two groups of multiparous cows were used for behavioural measurements. There
were eight cows in group 1 and nine cows in group 2. Measurements were made on
cows in group 1 on seven occasions between 8 May and 8 July and on cows in group 2
on nine occasions between 10 May and 10 July. The number of cows measured on each
occasion varied between three and eight, depending on the availability of equipment.
Grazing behaviour was recorded automatically over 24-h periods commencing at
morning milking (approximately 07.00 BST), using recorders that stored analogue
electrical signals of the jaw movements on miniature tape recorders carried on the
animal (Huckle et al., 1989). Subsequent computer analysis of the recorded waveforms
allowed the time spent grazing, ruminating and idling to be obtained. Data were
summarised on a minute by minute basis, an animal being deemed to be engaged in one
of the three activities if it spent at least 30 s in any 1 min engaged in that activity. Some
A.J. Rook I CA. Huckle/Applied Animal Behauinur Science 54 (19971 E-96 ‘)I

other aspects of the grazing behaviour of these animals are discussed by Rook and
Huckle ( 1996).
Meal criteria, defined as the minimum duration of an inter-meal interval, were
established using data for frequency of occurrence of non-eating periods of different
durations. Three methods of defining the criterion were used. The method of Forbes et
al. (1986) relates the frequency of occurrence of periods of non-activity of different
length to that length using piecewise linear regressions. The break point of the best
fitting regression is taken as the criterion. The methods of Sibly et al. (1990) and Berdoy
(1993) involve the explicit assumption that two or three Poisson processes generate the
periods of non-activity and the fitting of these processes using non-linear methods.
Criteria for determining ruminating and idling bouts were obtained in a similar manner.
In addition the frequency distributions of non eating periods consisting of ruminating
alone, idling alone or a mixture of the two activities were also considered. This was in
order to check whether the nature of the interval was affecting the overall distribution of
intervals observed. Data for each concentrate feeding level were analysed separately. In
addition, the two periods at pasture between morning and evening (a.m.) and evening
and morning (p.m.) milkings were analysed separately within each concentrate level. as
were data for individual animals.
Once a common bout criterion for all activities had been established the data were
smoothed. This was achieved by examining a window of 2n - 1 min, where n is the
bout criterion, and assigning to the mid-point minute in the window the predominant
activity, on a minute by minute basis, within the window. Two to three iterations of this
process were usually sufficient to ensure that all bouts smaller than the bout criterion
were removed.
The proportion of each recorded activity reassigned by the smoothing process was
examined. The distribution of the intra-bout activities within the smoothed bouts was
also examined.

3. Results

Meal criteria could not be established using the methods of Sibly et al. (1990) or
Berdoy (1993) as these models proved highly unstable and generally failed to converge
to a solution. The method of Forbes et al. (1986) was, therefore, adopted. The meal
criteria for eating, ruminating and idling were all 5 min regardless of the treatment
imposed. Thus a criterion of 5 min was used in the smoothing process, resulting in a
window of 9 min being used.
There were no significant differences between a.m. and p.m. periods, between
individual animals or (with the exception of ruminating bout duration) between concen-
trate levels for any of the variables examined. The mean results for the three concentrate
levels are, therefore, presented in Table 1 by way of example. The mean proportion of
time spent in an activity that was reallocated as a result of smoothing was 0.058 for
eating, 0.072 for ruminating and 0.081 for idling. The maximum for a 24-h recording on
an individual animal was 0.329 for eating, 0.301 for ruminating and 0.191 for idling.
These reallocations resulted in bouts that contained at most 0.069 of other activities for
92 A.J. Rook, C.A. Huckle/Applied Animal Behauiour Science 54 (1997) 89-96

Table 1
Effect of concentrate level on grazing behaviour
Concentrate level (kg/day) 0 2 4 S.E.D. F. prob

Total daily grazing time (unsmoothed) (mitt/day) 617 581 591 20.7 0.191
Total daily ruminating time (unsmoothed) (mm/day) 321 311 326 19.2 0.676
Total daily idling time (unsmoothed) (min/day) 315 425 405 24.4 0.125
Total daily grazing time (smoothed) (min/dayl 622 590 598 21.3 0.276
Total daily ruminating time (smoothed) (min/day) 331 313 331 21.5 0.625
Total daily idling time (smoothed) (min/day) 366 415 394 25.1 0.147
No of meals/day 8.87 10.12 10.22 0.802 0.163
No of ruminating bouts/day 9.21 10.42 9.58 0.756 0.269
No of idling bouts/day 12.18 13.58 13.11 0.844 0.243
Meal duration (min) 79.1 68.3 66.8 7.55 0.195
Ruminating bout duration (mini 41.8 31.6 38.2 4.05 0.043
Idling bout duration (mitt) 31.0 31.5 31.2 2.00 0.967
Proportion of eating time reallocated 0.054 0.066 0.056 0.010 0.475
Proportion of ruminating time reallocated 0.061 0.083 0.075 0.013 0.258
Proportion of idling time reallocated 0.074 0.082 0.086 0.009 0.383
Intra-meals intervals as proportion of total meal time 0.063 0.077 0.068 0.009 0.26 1
Intra-ruminating intervals as proportion of tota 0.064 0.077 0.08 1 0.010 0.555
ruminating bout time
Intra-idling intervals as proportion of total idling 0.052 0.056 0.055 0.007 0.208
bout time
Ratio smoothed: unsmoothed eating time 1.010 1.101 1.013 0.011 0.960
Ratio smoothed: unsmoothed ruminating time 1.007 0.997 1.009 0.014 0.622
Ratio smoothed: unsmoothed idling time 0.978 0.973 0.967 0.520

eating, 0.076 for ruminating and 0.054 for idling. The ratio of total time in an activity
after smoothing to that before smoothing was 1.012 for eating, 1.005 for ruminating and
0.973 for idling.
The pattern of time regarded as spent in each activity after smoothing for each hour
of the day, together with the proportion of intra-bout intervals is shown in Fig. 1 a, b and
c for eating, ruminating and idling, respectively. The proportion of meals attributable to
intra-meal intervals was rather lower during the evening (1700- 1900 h). Thus not only
do animals take longer meals at this time (Rook et al., 1994) but these meals are also
more intense. Meals that occurred during the night had a higher proportion of intra-meal
intervals. This may reflect an increased need for vigilance when feeding during darkness
or an increase in the search component of foraging, due to increased difficulty of finding
food in the dark. By contrast, bouts of ruminating and idling were more sustained during
the night. These patterns generally reflect the overall time recorded in each activity on
either a smoothed or unsmoothed basis.

4. Discussion

The failure of the Sibly and Berdoy methods to converge suggests that the postulated
underlying Poisson processes may not in fact occur. Rook and Penning (1991) have
A.J. Rook, C.A. Huckle/Applied Animal Behmiour Science 54 (1997) 89-96

.
.
. .
.

. . . . . . .
. . . . .
. .
.

. .
. .
. . ?? ? ?? ? ? ?? ?
? ? ? ?? ?
? ?
? ?? ?
? ?? ?? ?

10

0
00:oO 1 mu OQ:O3

Fig. 1. The mean pattern of activity bouts over 24 h. including intra-bout intervals for (a) eating, (b)
ruminating. Cc) idling.
94 A.J. Rook 1C.A. Huckle/Applied Animal Behuiour Science 54 (1997) 89-96

15:OU

hour ofday

Fig. 1.(conrinued)

shown that while a Markov process can be used to describe the time course of behaviour
in sheep an age-dependent process may in fact be operating. Metz (1975) also found
evidence of an age-dependent process in housed cattle. In these circumstances the
Poisson assumption of randomness does not hold, even for one of the sub-processes. The
data discussed here have a resolution of only 1 min due to the computer algorithms used
to analyse the recordings. However, new equipment and methods for recording jaw
activity will allow resolution down to l/20 s and may help to further elucidate the
patterns involved (Rutter et al., 1993). The three-process model (Berdoy, 1993) may
have more applicability in this context. In addition, Alados et al. (1996) have recently
employed fractal geometry to analyse feeding behaviour, an approach which may be
useful with the type of data discussed here.
Metz (1975) reported a minimum inter-meal interval of 20 min for cows in indoor
feeding trials using survivorship curves. However, his data were only recorded to a
resolution of 4 min and thus a criterion of 5 min such as that reported here could not
have been determined. He also found that there were relatively few short intervals
between ruminating activity and also relatively few short meals. However, when
intervals less than 4 min are included, as here, there is a clear break in the frequency
distribution at 4 min. Forbes et al. (1986) used a meal criterion of 8 min for their dairy
cows fed indoors. Prache (1984) used a meal criterion of 7 min. The ubiquity of the
A.J. Rook. C.A. Huckle/ Applied Animal Behariour Science 54 f 19971 89-96 9s

5min criterion across all three activities and all the treatments, in this study, is a strong
argument for its adoption, especially when multi-activity smoothing is being carried out.
This study emphasises the need to report the intensity of bouts of behaviour, in a
quantitative manner. as well as their number and duration. For example, the ‘determined
character’ of grazing during the evening was first pointed out by Castle et al. (1950) but,
to the authors’ knowledge, this is the first quantitative demonstration of the effect.
Prache (1984) attempted to define two types of meal based on the intensity of eating
activity within meals. The criterion used was greater than (type 11 or less than (type 2)
50% of the meal actually engaged in eating. This would seem to be an excessively lax
criterion; it is questionable whether type 2 meals should be regarded as meals at all.
Such meals would be counted as non-eating using the smoothing method described here.
There was no evidence in this study that intra-meal intervals became longer as a meal
progressed, or that they were concentrated towards the end of a meal. However, Penning
et al. (1993) reported data for sheep that suggested that intra-meal intervals increased in
length and frequency towards the end of a meal. Smoothing could, therefore, under some
circumstances mask important features of the data. It is also clear that unsmoothed,
rather than smoothed. data should be used when reporting total activity duration or
calculating total intake.

Acknowledgements

This work was funded by a commission from the UK Ministry of Agriculture,


Fisheries and Food. We thank J. Whyte and S. Hammond for the care of the animals and
A.J. Clements. N. Daffem and M. Sampson for skilled technical assistance.

References

Alados. C.L., Escos, J.M., Emlen. J.M., 1996. Fractal structure of sequential behaviour patterns: an indicator
of stress. Animal Behaviour 5 1, 337-443.
Berdoy. M., 1993. Defining bouts of behaviour: a three process model. Animal Behaviour 46, 387-396.
Castle, M.E., Foot, AS., Halley, R.J., 1950. Some observations on the behaviour of dairy cattle with particular
reference to grazing. J. Dairy Research 17, 215-230.
Forbes. J.M., Jackson. D.A.. Johnson, C.L., Stockill, P., Hoyle. B.S.. 1986. A method for the automatic
monitoring of food intake and feeding behaviour of individual cattle kept in a group. Research Develop-
ment in Agriculture 3, 175-180.
Holmes, C.W.. 1974. The Massey Grassmeter. Dairy Farming Annual, pp. 26-30.
Huckle, C.A., Clements, A.J., Penning. P.D. A technique to record eating and ruminating behaviour in dairy
cows. In: Phillips, C.J.C., Ed., New Techniques in Cattle Production. Buttetworths, London, pp. 236-237.
Metz, J.H.M., 1975. Time patterns of feeding and rumination in domestic cattle. Meded. Landbouwhogesch.
Wageningen 75, 12.
Penning, P.D.. Orr, R.J., Parsons, A.J., Rutter. S.M. 1993. Factors controlling meal duration and herbage
intake by grazing sheep. 27th International Congress of the International Society for Applied Ethology,
Berlin. 26-30 July, 1993, pp. 498-500.
Prache, S., 1984. Electronic device to determine influence of number of lambs suckled and feed supplementa-
tion on grazing behavjour. Proceedings of the Herbage Intake Workshop at Edinburgh.
Rook, A.J., Huckle, C.A., 1996. Sources of variation in the grazing behaviour of dairy cows. J. Agric. Sci.
(Cambridge) 126. 227-233.
96 A.J. Rook, C.A. Huckle /Applied Animal Behaciour Science 54 (1997189-96

Rook, A.J., Huckle, CA., Penning, P.D., 1994. Effects of sward height and concentrate supplementation on
the ingestive behaviour of spring calving dairy cows grazing grass-clover swards. Applied Animal
Behaviour Science 40, 101-l 12.
Rook, A.J., Penning, P.D., 1991. Stochastic models of grazing behaviour in sheep. Applied Animal Behaviour
Science 32, 167-177.
Rutter, S.M., Penning, P.D.. Champion, R.A., Roberts, G., 1993. Recent developments in the automatic
recording of grazing behaviour in free ranging ruminants. 27th International Congress of the International
Society for Applied Ethology, Berlin. 26-30 July, 1993, pp. 594-596.
Sibly, R.M., Nott, H.M.R., Fletcher, D.J., 1990. Splitting behaviour into bouts. Animal Behaviour 39, 63-69.
Wilkins, R.J.. Gibb. M.J., Huckle, C.A., Clements, A.J., 1994. Effect of supplementation on production by
spring-calving dairy cows grazing swards of differing clover content. Grass and Forage Science 49,
465-475.

You might also like