APPLIED ANIMAL BEHAVIOUR SCIENCE

ELSEVIER

Applied Animal Behaviour Science 49 (1996) 321-333

Behavioural comparison of layer and broiler fowl: measuring fear responses

S. Keer-Keer, B.O. Hughes *, P.M. Hocking, R.B. Jones
Roslin Institute (Edinburgh), Roslin EH25 9PS, UK

Accepted 27 February 1996

Abstract Broilers and White Leghorns have different temperaments; White Leghorns are regarded as flighty and broilers as docile. This difference in temperament is sometimes interpreted as a difference between the strains in underlying fearfulness. Five experiments were carried out to compare fear responses of the two strains. All experimental birds were fed a medium-energy (breeder) replacement diet to reduce strain differences in body weight and locomotor activity which accompany provision of standard broiler grower diets. The nature of the test stimuli, the test situation and the age of the bird were varied between tests. In three of the experiments, differences in fear response between strains were minimal or absent. These included: exposure of isolated IO-day-old chicks to a person situated at one end of a novel arena (Experiment l), exposure of individually caged 1l-week-old birds to a person approaching from 1 m away (Experiment 31, and exposure of the same birds to a novel object placed at the front of the home cage (Experiment 4). In two experiments, strain differences were apparent. These were: exposure of 3-week-old chicks, placed in pairs in a novel cage, to a person approaching from 4 m away (Experiment 2) and exposure of 16-week-old birds, in groups of seven, to a large novel object (road hazard cone) placed in the test pen (Experiment 5). In both cases the Leghorns showed more marked avoidance responses than broilers. Leghorns physical response is often more vigorous than that of broilers, and it was difficult to separate putative strain differences in the type of behavioural strategy adopted upon exposure to threat from the effects of underlying fearfulness. Results indicate either that there is no difference in fear between the two strains, or that Leghorns are more fearful than

* Corresponding author. Tel.: (0131) 527-4200; fax: (0131) 440 0434. 016%1591/96/$15.00 Copyright 0 1996 Elsevier Science B.V. All rights reserved. PII SOl68-1591(96)01055-6

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broilers, but strain differences only become apparent in a social situation or when the levels of fear evoked by the test are moderate.
Keywords:

Chicken; Fearful behaviour; Genetics

1. Introduction The UK Farm Animal Welfare Council have identified five freedoms as being important for safeguarding animal welfare (Webster and Nicol, 1988). One of these is freedom from fear . Fearfulness can be described as a propensity to be easily frightened by a variety of potentially alarming situations (Jones and Waddington, 1992; Jones, 1996). Domestication has produced animals which seem to show lower fearfulness than their progenitors but there is still pronounced variation within and between strains. There has long been a consensus (Murphy, 1977; Murphy and Wood-Gush, 1978; Duncan, 1985; Jones, 1987a) that breeds, stocks and strains show consistent differences in temperament (distinct nature and character). One of the most obvious traits is flightiness , which manifests as rapid movement or flight away from a recognised disturbing stimulus. Stocks selected for meat production are regarded as less flighty than those selected for egg production (Siegel, 1984; Appleby et al., 1992). Strain differences in flightiness have been interpreted in terms of differences in underlying fearfulness or emotionality (Murphy and Wood-Gush, 1978; Appleby et al., 1992). There have been numerous comparisons of fear between light- and medium-hybrid laying fowl (Gallup et al., 1976; Jones, 1977; Murphy, 1977; Jones and Mills, 19831, but few in which White Leghorns and broiler chickens were directly compared. This paucity of information may reflect the difficulty in developing a reliable and appropriate methodology. Fear is recognised by behavioural changes which include diverse forms of defensive behaviour such as passive avoidance, freezing, tonic immobility, withdrawal and vigorous escape (Kruijt, 1964; Duncan, 1985; Jones, 1987b). However, there are large differences between stocks in their inherent level of locomotor activity. Some, like Leghorns, are active, while others, like broilers, are lethargic (although the locomotor activity of broilers is greatly increased when they are restricted fed; Hocking et al., 1993). Pronounced differences in locomotion make it difficult to use activity-based measures as indices of fear when comparing stocks. White Leghorns and broilers in the present study were fed a medium-energy (breeder) replacement diet; this would have reduced, though not eliminated, differences in body weight and locomotor activity which would otherwise have accompanied the provision of standard broiler grower diets. Duncan (1981) suggested that although fear is a hypothetical intervening variable, it is still possible to define it operationally. Fear ideally functions to protect the animal from danger (Toates, 1980; Jones, 1987b) and because it takes precedence over and inhibits all other behaviour systems (Jones, 1987b; Jones, 19961, it is argued that the extent to which a test stimulus is avoided reflects the birds fear of it. Several factors are known to induce or influence fearfulness in domestic fowl; separation from conspecifics, exposure to a novel environment or to an unfamiliar object, approach of or proximity to a human being. The objective of the series of

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experiments described in this paper was to examine systematically the withdrawal/avoidance response of a White Leghorn and a broiler line in a number of tests in which these factors differed considerably.

2. General methods The comparisons were between both sexes of two parent-stock pure lines: a commercial male-line broiler (Ross Breeder Ltd., Newbridge, UK) and a White Leghorn line maintained as unselected stock for six generations at the Roslin Institute. The birds were housed at 1 day of age in single-sex, single-strain groups of 25, in eight pens (2.44 m X 1.52 ml containing wood shavings. The pens were arranged in a systematic order to control for possible environmental effects. At 7 weeks of age the birds in each pen were split into two groups, one group was removed to a new pen, creating a total of 16 pens. All birds were fed a standard replacement (breeder) medium-energy diet ad libitum. Water was provided ad libitum in suspended bell-shaped drinkers, one per pen.

3. Experiment

1: Box plus experimenter

test

This test (Jones and Waddington, 1992) measures the approach/avoidance responses of chicks to a human visible behind a wire wall, at the end of an unfamiliar arena. Because withdrawal is thought to be a functional fear behaviour (Toates, 1980), marked avoidance in this test situation is considered to reflect increased fear of human beings. Chicks will view a stimulus either with the left or right eye according to its nature or novelty value (Dharmaretnam and Andrew, 1994; Vallortigara and Andrew, 1994) and they may use this strategy to self regulate their state (R.J. Andrew, personal communication, 1995). Therefore, lateralisation of viewing the experimenter, was also recorded. 3.1. Method The chicks were tested on successive days when 10 or 11 days old; approximately half of each strain were tested on each day. Twenty-eight Leghorns (15 females, 13 males) and 28 broilers (14 females, 14 males) were caught individually at random from successive pens (all pens represented) and carried to a nearby empty pen. Four individual Leghorns were tested, followed by four individual broilers. Each chick was placed in the centre of a rectangular box of 62 cm X 40 cm X 30 cm (length X width X height), successive chicks alternately facing the left or right side of the box. The box had three wooden walls and one of wire mesh (mesh size 1.5 cm>, the floor was covered in wood shavings. The observer (S. K-K) sat directly in front of and facing the wire mesh wall, with head and upper torso visible to the chick and approximately 30 cm away from the box. The box was divided widthways into four imaginary areas 15 cm X 40 cm (length X width), and the chick s position recorded every 15 s during the 5 min test period. At each scan the chick scored 1 if it was in the area closest to the experimenter, through 2 and 3, to 4 at the far end. The total scored (avoidance) by each chick was the

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Table I Experiment

I: means (f SE) scores of Leghorn Leghorn

and broiler chicks in box plus experimenter Broiler 3.00*0.091 - 1.00+ 1.14 3.29 + 0.56 Significance NS NS 0.001

test (N = 28)

(P 1

Avoidance Lateralisation No. of jumps

2.84 + 0.077 0.39 + 1.05 10.36f 1.51

mean of 20 scans; high scores reflected high avoidance. While the position of each chick was recorded, the eye with which it appeared to be observing the experimenter was simultaneously noted. A lateralisation of viewing score was derived by calculating the total number of times the chick was recorded viewing with the right eye minus the number of times it was recorded viewing with the left. The number of jumps made by each chick during the 5 min test period was also recorded. 3.1.1. Statistical analysis The effects of sex, strain, initial position and test day, and their interactions, were analysed using an ANOVA. Numbers of jumps by strains were compared using the Mann-Whitney U-test.
3.2. Results

The results are shown in Table 1. Neither the avoidance nor the lateralisation of viewing scores differed between strains, but Leghorns jumped significantly more often than broilers. An almost significant interaction of sex and strain (P = 0.054) suggested that male Leghorns showed greater avoidance than females, whereas male and female broilers had a similar avoidance score to each other. Gender exerted no detectable influence on lateralisation of viewing or jumping.

4. Experiment 2: Avoidance response to an approaching

human (corridor test)

This experiment measures the behavioural response of a bird to a human approaching along a corridor and is based on methods described by Jones et al. (19811, Hemsworth and Bamett (1989) and Bamett et al. (1992). Chicks were tested in pairs in this experiment because pilot trials suggested that increased fear levels associated with isolation could mask responses to the approaching human. 4.1. Materials and methods 4.1 .I. Birds Birds were tested in random order when 3 weeks old as same-sex, same-strain pairs; they were different individuals to those used in Experiment 1. Eight pairs (four male,

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four female) of each strain were tested, each pair once only. They were caught manually in their home pens and transported (a 5 min journey) as test pairs in a closed box. 4.1.2. Testing environment This was a separate room, lit by moderate-intensity (120 lux) fluorescent lights and maintained at a temperature of 24C. Four cages of 40 cm X 60 cm X 68 cm (width X length X height) were used; they had plywood sides, 2-cm wire-mesh fronts, tops and backs, and flat, littered floors. A cage was placed on a lOO-cm-high table 30 cm from the end of each of four separate parallel corridors (5.3 m X 1.0 m; length X width), the walls and one end (opposite end to the cage) of which were formed of pale beige, cotton material suspended on wires, like a curtain. A video camera (Panasonic S-VHS 625, wide-angle lens), placed behind each cage, recorded the bird s behaviour; the approaching human was also visible in the field of view. 4.1.3. Procedure Each pair was left to acclimatise in the test cage for l-2.5 h, mean acclimatisation time was the same for both lines. Thirty minutes before testing, the video camera was placed behind the cage. The camera was switched on remotely 5 s prior to testing. At the start of the test the experimenter (S. K-K) emerged from behind the curtain at the end of the corridor at a distance of 4.3 m from the cage front. She advanced slowly, pausing for a 10 s interval at 3.5, 2.5 and 1.5 m, and for 5 s at 1.0, 0.5 and 0 m, before finally placing a hand over the top of the cage. The behaviour of the birds was recorded on videotape and analysed. Their flight and avoidance reactions were scored on a three-point scale (see Table 2). High scores reflect increased agitation. In almost all cases both birds were visible and both were engaged in the same activity. Where birds were engaged in different activities both were scored and a mean value was calculated. Where only one bird could be seen clearly only its score was taken. 4.1.4. Statistical analysis Avoidance scores were compared between strains using the Mann-Whitney U-test. The distribution of number of birds showing their first category 3 (flight) response by approach stage, was analysed using an extended Fisher exact test. The effect of pre-test acclimatisation time and different test corridor were analysed using ANOVA. 4.2. Results The mean avoidance Fig. 1. At all positions behaviour scores of both strains at each stage are summarised in from 2.5 m onwards the Leghorns scored more highly (greater

Table 2 Experiment

2: behavioural Description

categories

and their associated

scores

score
0

1
2

Sit, alert (head movements), stand still Major body movement away from stimulus Flight response: agitated movements at rear of cage, sudden crouch, jump, panic, frantic mMing

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4.3

3.5

2.5

1.5

1.0

0.5

0.0

0.0

a
in response to an

Approach

stage (m)

Fig. 1. Experiment 2: mean ( + SE) avoidance behaviour scores of broilers and Leghorns approaching human. At Oa m the experimenter s hand was placed on top of the cage.

the broilers, and the mean scores overall were significantly higher (1.80) than for broilers (1.27). The Leghorns showed a panic or flight response at an earlier stage in the approach (mean distance 1.2 m> than the broilers (mean distance 0 m), and the distribution of first flight response by approach stage (Table 3) was significantly different between lines (P < 0.004). Corridor used and pre-test acclimatisation time were not found to have an effect on mean behavioural score. No sex by strain interactions were detected.
(P = 0.004) for Leghorns

agitation)

than

5. Experiment 3: Home cage approach In this test, like the previous one, we measured the responses of birds to human approach. However, here the birds were tested individually in a home cage at 11 weeks

Table 3 Experiment 2: numbers of bird pairs showing category 2 (flight response) (N = 8). At Oa m, the experimenter s hand was placed on top of the cage Distance of approaching 4.3 Leghorns Broilers 0 0 3.5 0 0 2.5 person from birds (m) 1.5 2 0 1.0 2 0 0.5 2 0 0 0 3

at successive

stages of approach No response

Oa 1 3 0 2

1
0

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old, and the human approached from a distance one described by Bamett et al. (1992). 5.1. Method

of 1 m in 20 s. The test was based on

When the birds were 10.5 weeks old, 12 males of each strain were removed from their home pens (three from each pen) and transferred to individual cages in the top and middle tier of a battery, measuring 40 cm X 48 cm X 58 cm (width X length X height) and 30 cm X 48 cm X 51 cm, respectively. The cages were allocated according to a blocking design to control for possible environmental effects such as tier or cage size. There were three blocks (two in the top tier and one in the middle tier) with eight birds (four of each strain) per block. Broilers and Leghorns were housed in alternate cages because of a requirement of a subsequent experiment. The photoperiod ran from 08:30 to 23:OO h. The birds had 1 week of acclimatisation in the cages before testing. Birds were selected in random order for testing, with the proviso that the bird was not in a cage adjacent to that used in the previous test. All birds in the experiment were tested on the same day. The experimenter (S. K-K) assumed a position directly opposite and 1 m away from the cage of the bird being tested, with her face at cage level, then approached in three stages: 1. stood motionless at 1 m distance for 10 s, keeping her hands by her side; 2. moved forward to 0.5 m from the cage, and held the position for 10 s; 3. moved forward to 0 m from the cage, placed a gloved hand and face (protected with goggles) on the bars of the cage front. This procedure was achieved for top tier birds by stepping on stools. The orientation of the bird was recorded on a five-point scale at 5 s intervals, that is, twice at each of the three distances. The scoring system was a modified version of the novel object scoring system (Hughes and Black, 1974a; Sefton, 1976). The scoring was as follows: 0, head out of the front of the cage; 1, head within the cage with bird oriented towards front; 2, bird facing the sides of the cage; 3, bird oriented towards the rear of the cage; 4, bird exhibiting escape behaviour, usually at the back of the cage. The total orientation score was the mean of the scores at the three stages: higher scores represent an increasing degree of fear/avoidance. Mean avoidance scores were compared between strains using a one-way ANOVA. 5.2. Results The mean ( f SE) avoidance scores were 2.17 f 0.08 for Leghorns for broilers; they were not significantly different. and 2.05 f 0.20

6. Experiment 4: Response to a novel object This test, based on one first described by Hughes and Black (1974a), measures the avoidance response of birds to a brightly coloured novel object placed in the food trough of their home cage.

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6. I. Method The birds tested here were the same as those used in Experiment 3 and were tested the day after Experiment 3. Birds were selected in random order for testing and all tests were completed on the same day. The novel object was a wooden rod 34 cm X 2.5 cm (height X width) rising vertically from a 10 cm X 15 cm square base. The rod was covered with five different colours of 2-cm-wide plastic tape forming a multicoloured series of bands. It was placed vertically in the centre of the food trough, at the front of the cage. The observer stood 3 m to the side of the cage. The bird s response to the object was recorded at 10-s intervals over a 3-min period (a total of 18 observations). The responses were scored as described in Experiment 3. The total scored by each bird was the mean of 18 scans. Mean avoidance scores were compared between strains using a one-way ANOVA. 6.2. Results The mean ( f SE) avoidance scores were 2.21 + 0.09 for Leghorns for broilers; they were not significantly different. and 2.3 1 f 0.08

7. Experiment 5: Pen and cone test In this test the responses of Leghorns and broilers to a measured. However, here the birds were tested in groups avoidance of the novel object was measured by calculating from the object. The test was based on one described by Jones the birds were recorded using video cameras. 7.1. Method The four test pens (two pairs) were in the same hut as the birds were housed. Each pen (2.44 m X 1.53 m) contained a food hopper and bell water drinker; they were similar to the home pens except that the litter was fresh. A black and white camera (Panasonic CCTV) was suspended above the centre of the pen, and the birds behaviour was videotaped. Two tests were filmed simultaneously, tests took place between 14:00 and 16:30 h, and four tests were filmed on each of four test days. Twenty-four hours before testing (at 16 weeks of age), four (same-strain, same-sex) groups of seven birds were removed from pre-selected pens and transferred to the four test pens. Immediately prior to testing, the food and water were removed from the first two pens and the camera suspended above the pen. The experimenter then left the room. For the next 30 min ( pre-cone session) the birds moved freely around the test pens. The experimenter returned after 30 min, placed a red road hazard cone (0.45 m high, 0.26 m diameter base) in the centre of each pen, and left the birds for a further 30 min ( with-cone session). The experimenter then returned and repeated the procedure for the other two test pens. The video tapes were analysed at a later date. novel object were again (of seven) in pens and the distance of each bird (1987c). The responses of

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A
0 A 0

Pre-cone Leghorn Pre-cone broiler

Cone IlO-

Cone Leghorn Cone broiler

loo-

= E S 8 5 D ._ n

90-

80-

Pre-cone

session

Cone session

Time period (mins) Fig. 2. Experiment 5: the mean distance of broilers and White Leghorns from the centre of the pen during pre-cone and cone sessions of the pen and cone test. Points represent means of previous five scans. Lines fitted: broiler, pre-cone y = - 0.091 x + 7 1.692, with-cone y = -0.459.x + 85.477; Leghorn, pre-cone y = 0.594~ + 63.724, with-cone y = - 0.272 x + 97.35 1. Maximum possible distance from centre, 120 cm; mean distance predicted by chance, 75 cm.

havioural inhibition may have reduced jumping in the broilers. Neither of these explanations is convincing, given the lack of difference in avoidance scores. Thirdly, chicks of both strains may have been equally frightened but social reinstatement motivation may have been greater in the Leghorns. Lastly, the difference may have reflected early line divergence in locomotor ability or motivation. In the home cage approach and novel object tests (Experiments 3 and 4) there was no detectable strain difference in avoidance. Because individual Leghorns and broilers in these tests were housed alternately, the influence of neighbouring birds, through a mechanism such as cultural transmission, on their fear behaviour should be considered. However, previously documented effects of mixing strains on their behaviour or performance were found to be slight (Hughes and Black, 1974b; Savory, 1975). These two experiments used the same individual birds and this may have produced a greater similarity in result between them than between other experiments which used different individuals.

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The two experiments (2 and 5) where a strain difference in fear was detected; corridor and pen and cone , were those where the subjects had one or more conspecifics present. Strain differences in fear may only become apparent in social test situations and it is conceivable that Leghorns are more sensitive to the transmission of alarm from conspecifics. In the pen and cone tests fear of the cone may have motivated avoidance of it or attraction to the walls. There is evidence in the literature suggesting that the positions of birds in a pen is influenced by the pen walls (Newberry and Hall, 1990; Pamment et al., 1982). This wall seeking behaviour is thought to be shelter or escape seeking behaviour, motivated by fear. Sex by strain interactions apparent in Experiments 1 and 5 suggested that male Leghorns were more fearful than female Leghorns, whereas male and female broilers behaved similarly to each other. The effect of gender on fear behaviour of fowl is reviewed in Jones (1987b) and where sex differences are reported results suggested that males were more fearful than females. In these experiments we cannot discount the effects of age on fear: Brake et al. (1994) found an increase in the duration of tonic immobility with age in broiler breeder pullets and Candland et al. (1963) found that freezing increased in White Leghorns up to the age of 20-45 days and then declined. It could be argued that the shorter flight distance and the greater avoidance of an approaching human and of the cone shown by Leghorns reflected a strain difference in the type of behavioural strategy adopted upon alarm. In other words, Leghorns may not be more fearful than broilers but they may react to the same stimuli in a different (flighty) way. For example, Duncan and Filshie (1979) exposed White Leghorns and broilers to frightening stimuli whilst simultaneously measuring the heart rate of the bird using a subcutaneous radio telemetry device. They found the behavioural reactions of the two strains followed a predictable trend, with Leghorns showing extreme panic behaviour and broilers a less violent alarm reaction. They also found that the heart rate of the Leghorns reached a higher maximum than the broilers, but the broilers heart rate took longer to return to resting levels. Both trends are associated with increased fear. They concluded that some reappraisal of the flighty or docile classification is required, since docile birds may be as frightened as flighty birds in physiological terms. Conversely, the results of Experiments 2 and 5 might represent a real strain difference in fearfulness, rather than just a difference in behavioural strategy. Thus, the findings present us with a contradiction; in two experiments a clear difference in fear response was demonstrable between the two strains, whereas in the other three no difference could be shown. It is possible that the experiments where no strain difference was detected induced particularly high (1) or low (3 and 4) levels of fear and those where a difference was detected (2 and 5), may have induced moderate levels of fear. This proposition could be tested by presenting birds with different stimulus intensities within a given experiment. Murphy and Wood-Gush (1978) suggested that certain procedures produce too high a level of fear to reveal strain differences and Jones and Faure (1982) found that increased novelty of the test environment abolished sex differences in chicks open field behaviour.

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The problems inherent in measuring fear have been addressed by Jones (1987b) and Jones (1987~) and although methodologies have been developed which detect fear differences in birds of the same strain, they may be less useful, or may require adaptation, when comparing fear between very different strains of birds.

Acknowledgements This work was supported by Ministry of Agriculture, Fisheries and Food. We thank Dave Waddington for statistical advice. Broiler chicks were kindly donated by Ross Breeders Ltd., Newbridge, UK.

References
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