Influence of breed on reactivity of sheep to humans

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Influence of breed on reactivity of sheep to humans

P Le Neindre, P Poindron, G Trillat, P Orgeur

To cite this version:

Original

article

Influence

of breed

on

reactivity

of

_sheep

to

humans

P Le

Neindre

1

P

Poindron

G Trillat

P

_Orgeur

1

INRA,

Laboratoire de

_{l’Adaptation}

des Herbivores aux _Milieux,

Theix,

63122

_{Saint-Genes-Champanelle;}

2

INRA/CNRS

(URA

1291),

Laboratoire de

Comportement Animal,

Station de

_Physiologie

de la

_{Reproduction,}

37380

_Nouzilly,

France

(Received

10 June 1991;

accepted

28

May

1993)

Summary - Ewe lambs from 2 breeds

_(M6rinos

d’Arles and

_Romanov)

and their

cross-breds,

from M6rinos ewes sired

by

Romanov rams, were observed

during

5 individual tests.

During

the first 3 tests there were no other

sheep

in

sight

and animals were

_alone,

with

concentrate or with a human.

_During

the last 2 tests, some _penmates were in

_sight

and the

experimental

animal was alone or with a human. Romanov animals were much more

reac-tive than the M6rinos.

They

eliminated more, ate less and avoided the human more. For

most of the criteria, crossbreds were closer to the Romanov than to the M6rinos

_purebreds.

This seemed to be due to

_genetic

differences and not to direct maternal influence.

sheep / breed

_/

_reactivity to man

_/

Mérinos

_/

Romanov

Résumé - Influence de la race sur le comportement d’ovins vis-à-vis de la présence

humaine. Des

_agnelles

de deux races

(Mérinos

d’Arles et

Romanov)

et des croisées issues

de mères Mérinos saillies _par des mâles Romanov ont été obervées dans _cinq

_épreuves

individuelles. Pendant les trois

_{premières épreuves}

il

_n’y

avait _pas de

congénères

en vue

et les animaux étaient

seuls,

avec de l’aliment concentré ou avec un homme. Pendant les deux dernières

_épreuves,

des

_congénères

étaient

_visibles,

dans un _{parc contigu,} et les

animaux étaient seuls ou avec un homme. Les

agnelles

de race Romanov ont été

beaucoup

plus

réactives _que celles de race Mérinos d’Arles. Elles

déféquaient

et urinaient

plus

fréquemment,

mangeaient moins et évitaient

_plus

l’homme. Pour la

_plupart

des critères les croisées avaient des

_{performances plus proches}

de celles des Romanov que de celles des

INTRODUCTION

For domestic animals human

_beings

are a common feature of the

_environment,

and

their

_adaptation

to human _{presence may} be

_important

for animal

production

and welfare. The

_adaptation

of animals to

_production

_systems

with

_{differing degrees}

of human intervention

_{depends partly}

on their

_reactivity

towards man. In this

_respect,

some breeds _may be better

adapted

than others because

_they

are

_basically

less

disturbed

_by

the _presence of man. There are indeed indications that the

_reactivity

of animals to humans varies between breeds and can be influenced

_by

_genotype.

This has been shown in

_poultry

_(Faure

and

Folmer, 1975;

Murphy

and

Duncan,

1977),

and also in domestic

_ungulates

such as cattle

(Dickson

et

_{al, 1970;}

_Murphey

et

al,

1980, 1981;

Boivin,

1991),

pigs

(Hemsworth

et

_al,

₁₉₉₀₎

and

_goats

_(Lyons

et

at, 1988).

’

In

_sheep,

several studies have been carried out

_concerning

the behavioural

reactivity

of animals to a novel environment

_(ie

_open

_field),

but data

_concerning

reactivity

to man and

possible

breed differences in this

_respect

are

_lacking.

For

example,

Zito et at

₍₁₉₇₇₎

and

_Moberg

and Wood

₍₁₉₈₂₎

_compared

the behaviour of lambs reared in

_isolation,

_{in groups} or with their mothers.

Similarly,

Lachaux

et at

₍₁₉₈₃₎

and Winfield et at

₍₁₉₈₁₎

_analysed

the reaction of

_sheep

_depending

on their

_familiarity

with each other. These studies showed differences in behaviour between the various

_types

of animals in various social or non-social situations.

Putu et at

₍₁₉₈₈₎

found differences in

_open-field

behaviour of ewe lambs that

were correlated with later maternal behaviour at first

_lambing.

_They

also

_reported

intraracial differences in the

_reactivity

of animals to humans. On the other

_hand,

breed differences are known to exist in several

behaviours,

such as selection of

lambing

sites

_(Alexander

et

_o!,

_1990),

_{mother-young relationships}

_(Alexander

et

al, 1983;

Shillito Walser et

_{al, 1983;}

Poindron et

al,

1984),

or

_open-field

behaviour

and reaction to the _presence of a

_dog

_{(Torres-Hernandez}

and

Hohenboken,

1979).

There are also indications that fear

_reactions,

_including

fear of

_humans,

_vary with the breed

_(Romeyer

and

Bouissou,

personal

communication).

It is

_possible

that the breed differences found in _open field reflect

_genotypic

variations in

_reactivity

of animals to various stressful

_situations,

_including

the presence of man. This would

appear to be the case from the results of

_Romeyer

and Bouissou

_concerning

fear

reactions.

_However,

other data on cattle fail to confirm this

_hypothesis;

Boivin

(1991)

found no correlation between

_open-field

behaviour and reaction of animals to their

_{handling by}

man.

Similarly,

_Boissy

and Bouissou

₍₁₉₈₈₎

did not find a

clear-cut relation between

_open-field

behaviour and reactions to man in heifers.

Obviously,

more

_{investigations}

are needed to

_fully

assess the

_possibility

of

_genotypic

influence in the reaction of

sheep

to man.

To further

_investigate

the

_possibility

that breeds of

_sheep

_may differ in their

re-activity

towards

_humans,

we

_compared

the behaviour of 2 breeds of

_sheep

_(M6rinos

d’Arles and

_Romanov)

that are

_subjectively

_reported

as

_differing

_widely

in their

reactivity.

We also studied one of their crosses in

_preliminary

_attempt

to

distin-guish

between direct and maternal effects. To this

_end,

we studied the behaviour

of females in the presence of man, in a standard situation. This necessitated

tak-ing

the animals to a closed area that was

_previously

unknown to

_them,

as well as

and also studied

_reactivity

to a novel environment. This allowed clarification of

the extent to which the behavioural reactions observed were

_specific

for

_reactivity

to man, and their

possible

relations with other

aspects

of

_general

_reactivity

that

are

_usually

taken into consideration when

_studying

_temperament

(Price

and

Thos,

1980;

Boissy

and

Bouissou,

1988;

see also

Archer,

1973 and

Jones,

_1993,

for

general

reviews).

MATERIALS AND METHODS Animals

The

_study

was carried out in 1988 at the ENSA--INRA

_experimental

station of Le Merle in south-east France. This station is run

_according

to the traditional

system

in this

_part

of

_France,

_{involving daily}

contact with man for most of the

year. Animals are

kept

indoors all

_winter,

and let out to

_pasture

_daily

for the rest of the _year,

_except

_during

summer when

_they

are driven to

_pasture

in the

_Alps

under the constant

_supervision

of a

shepherd.

Fifty-five

female lambs _{1 yr} of _age were used in the

experiment.

They

had been

reared

by

their mothers in the same _group from

birth,

weaned at 3 months of age and

kept

thereafter in one

single

all-female _group.

They

were of 2 breeds: M6rinos d’Arles

(N

=

_19,

_a French

population

from south-east

_France)

and Romanov

_(N

=

_20,

_a

Russian breed used for its

_high

_prolificacy)

and their cross

_(N

=

_16,

M6rinos ewes

sired

_by

Romanov

_rams)

that will be referred to as &dquo;crossbred&dquo;.

Testing

procedure

Five individual tests of 4 min each were conducted over a 5-d

_period.

Two _persons drove each animal from the barn to the

_testing

room, which consisted of a 4 x 6 m

enclosure with

plain

walls. Six squares were marked on the floor with

plaster powder

(fig 1).

The test were as follows:

- Test 1:

subject

alone. This

corresponded

to the classical

open-field

test described in the literature

_{(Archer, 1973);}

- Test

2:

_subject

alone. Concentrate

_{(100 g)}

was

_placed

in the middle of the enclosure

_{(square 2).}

The

_latency

before

_eating

and the duration of

_feeding

_provided

objective

measurements of the

_degree

of distress caused

_by

social isolation and

novelty;

- Test 3:

a human stood

_stationary

in square

_{5 ;}

- Test 4: five

non-experimental

animals from the same flock were set behind a

wire-mesh door in front of square 5. The

subject

was alone in the

enclosure;

-

Test 5: as in test

_4,

but with a human

standing

in square 5.

Each animal was tested

only

once

daily.

On the first 3

d,

approximately

one-third

of the

_subjects

from each breed were allocated to one of the first 3 tests

_(table

_I).

On the last 2

_d,

half of the animals from each breed were allocated to 1 of the last 2 tests. On the first d of

_study,

animals were selected at

random,

_except for

a

_crossbred;

the order

_changed

for each _group of 3 animals. The 3 females were

tested alone.

_Then,

the next 3

_(again

one of each

_genotype)

were tested with

_food,

and the next 3 with man. This sequence was

repeated

until all animals had been

tested,

and each animal was marked on the

head,

the back or the rump,

according

to the

_type

of test carried out. The

_{following day,}

the same

_{testing procedure}

was

used. A similar

_procedure

was used for tests 4 and 5. In this _way,

_type

of test and breed were

kept

balanced

throughout

the

study.

The

_personnel

remained the same

_during

the whole

study,

and all

_operators

wore similar blue

clothing

(overalls

and a

jacket).

_During

the _tests, animals could see the observer who sat on a

platform

2 m above the

_ground.

A data recorder

with an internal clock

(DATAMYTE

1006)

was used to record the activities. All the

personnel,

including

the man

standing

in the

_testing

_pen, were unknown to the animals before the start of the

_study.

Behaviour recorded

The

_following

behaviour was recorded:

- while the

ewe was

_being

driven to the

_testing

room, the number of times it turned back and tried to force a _passage

through

the

personnel

and back to the

barn;

-

during

all 5 tests: the number of _squares

_crossed,

the number of times the animals sniffed the

_ground,

door and

_walls,

the number of low and

_high-pitched

bleats,

defaecations and

urinations,

looks in the direction of the

_{observer, rearings}

against

the

wall ;

-

- -

during

tests 3 and

_5,

the

_latency

before

_sniffing

and number of sniffs at the

observer;

-

during

tests 4 and

_5,

the time

_spent

_{in square} 5. Statistical

_analyses

The overall results for the breeds over the 5 d were

_{compared using}

_Mann-Whitney

tests. Parameters that were measured in the 5 tests were summed. The numbers of

times animals came back when driven to the

_testing

_pen when there was no other

sheep

in

_sight

_(first

3

_tests)

were summed.

More detailed

_analyses

were then made of 4 variates

(numbers

of _squares

_crossed,

of

_high

and low

_pitched

bleats and of

_{eliminations).}

_Square

roots of these variates

were used to obtain

_homogeneity

of the variances. Influences of

_type

of test, order of

day

and breed were tested with variance

analyses according

to the

_following

basic model:

where:

Y

i1kl

is the

_performance

of the lth individual with the

_jth

genotype

in the kth

type

of test on the ith

_day;

C is a _constant;

G

j

is the effect of the

_jth

_genotype;

T!

is the effect of the kth

_type

of test;

GT!!

is the effect of the interaction between the

_jth

_genotype

and the !cth

_type

of

_test;

e2!!! is a random effect.

These

_analyses

were made

_separately

for test without _congeners

_(first

3

_d)

and for tests with _congeners

_(last

2

_d).

The

_{intragenotype}

correlations of the sums of the results from tests 1 and 3

(respectively

alone or with a

_human,

first

_period)

with those from tests 4 and 5

(with

peers in

sight

when alone or with a

human,

second

period)

were calculated.

Correlations between variables within the same

_periods

were also calculated.

Effect of the _presence of congeners was

tested,

on these 4 variates and on the

number of times the animals came back when driven to the

_testing

pen

by studying

the differences between the results obtained in

_periods

1 and 2. As

_period

and treatment are

_confounded,

their effects cannot be isolated in this

_analysis.

RESULTS

Influence of breed

_(overall

_comparisons

of table

_II)

When the results of the 5 tests were considered

_together,

M6rinos animals differed

significantly

from Romanov animals in most behaviours studied. Merinos sniffed the

_ground

and door more

frequently,

and emitted more

high-pitched

but fewer

low-pitched

bleats than Romanov ewes.

_They

eliminated

(defaecated-urinated)

and

looked in the direction of the observer less often.

_They

spent more time in square 2

eating

concentrate. In the _presence of the human

_{observer, they}

sniffed him sooner

and more often. The 2 breeds

_spent

a similar amount of _{time in square} 5 near the

congeners in the absence of an _operator

(test 4).

_By

contrast, in the latter’s _presence the time spent near the _congeners was

_{significantly}

lower in the Romanov

_{(test 5)}

while in the M6rinos there was little difference whether or not a human was _present

during

the test. When driven to the enclosure the Merinos turned back less often than the Romanov.

_However,

_despite

all these differences the number of squares

crossed,

the usual variate for

_{indicating general}

_activity,

did not differ

_{significantly}

between the 2 breeds.

Crossbred animals had

_performances

_midway

between those of their

_parental

genotypes

for 4 parameters and in 3 cases

_(sniffing

the

_door,

_feeding

_time,

time in

square 5 with a

human)

the results did not differ from those of the other _{2 groups.} The differences

_regarding

both Mérinos and Romanov were

significant only

for the

number of times the animals turned back when driven to the

_testing

pen. For 5 of the

_remaining

_parameters

_performances

of crossbreds were similar to those of the Romanov.

_Only

in 2 cases did

_crossing

result in

performances

similar to those of

the Merinos

_genotype

_(number

of

_low-pitched

bleats and time in square

2).

Only

one

mobility

_parameter

(numbers

of _squares

crossed)

was lower for the crossbred

Analyses

of variance

The influences of

_day

and the interaction between test and breed were never

significant

in the variance

_analyses

for the 4 variates

_analysed.

On the other

_hand,

the 2 other factors

_(the

_type

of test and

_breed)

had

_significant

influences.

Type

of test

_{(table III)}

The presence of food was associated with a decrease in the numbers of _squares crossed and of

_high-pitched

bleats. A similar

_tendency

was observed for the influence

of human _presence on the number of crossed _squares. In _contrast, eliminative

behaviour and

low-pitched

bleats did not

_change.

The _presence of _congeners

Effects of

day

within the 2

_periods

were not

_significant.

It is then

_possible

to consider that the effect of

_periods

is also not

_significant

and that differences can be attributed

mainly

to the absence or _presence of _congeners. The _presence of _congeners was

associated with a clear-cut reduction in locomotor

_activity

and a reduction in

eliminative behaviour

_{(-2.06 !}

1.82,

p <

0.01:

-0.87 ±

0.97,

p <

_0.01).

The difference in

_high-pitched

bleats was not

_significant

_{(0.57 ! 2.11, NS).}

_Low-pitched

bleats increased

_{significantly}

_only

for the Romanov

_{(+1.51 !}

_1.65,

_p <

_0.01).

The number of times the animals came back when driven to the

_testing

pen

decreased

_sharply

when there were _congeners

_penned

next to the

_testing

enclosure

(-1.92 !

1.86,

p <

_0.01)

which could be seen

_by

the

_subject

while it was

_being

driven to be tested.

Relationships

between

variates

Correlations of measures

_according

to the _presence or not of _congeners The correlations between results of the same variate from the sums of tests 1 and 3

with those of tests 4 and 5 were

highly significant

for all the variables _except for the

number of

low-pitched

bleats

_(squares

_crossed,

!° =

_0.402,

_p <

_0.01;

_high-pitched

bleats,

r =

0.64,

p <

0.01;

low-pitched,

bleats,

r

= 0.16,

NS :

eliminations,

r =

0.38,

p <

0.01).

Intraperiod

relationships

between variates

Within a

_period

_(with

or without the _presence of a

_human)

the numbers of crossed

squares and the number of

high-pitched

bleats were

significantly

related

(first

period,

r =

0.31,

p <

_{0.05; second period,}

r =

_0.28,

p <

_0.05).

The number of

low-pitched

bleats and number of crossed _squares was related

only during

the first

period

(r

=

0.52,

p <

_0.05).

The number of eliminations and number of crossed

DISCUSSION

A number of

_points

emerge from the

study.

The differences observed between the 3

types

of animals indicate that

_they

differ _very

_clearly

in their behavioural reactions

to man, as well as in their reactions to a novel situation. The

_good

correlation of measurements in the _presence or in the absence of _congeners, the lack of effects of the

_day

of

_testing

and the order used for the various tests further demonstrate the

_validity

of these behavioural tests and their

_ability

to discriminate between

sheep

on the basis of

_temperament.

Frequency

of

eliminations,

usually

considered an

indicator of fear or behavioural

_discomfort,

was

_higher

in Romanov than in M6rinos.

Similarly, feeding

time and time _spent in the square

containing

food were lower in

Romanov, again

indicating

a

_higher

degree

of

_reactivity

in this breed. This was

also evident in the reactions towards man, where human presence was

_perceived

as

unpleasant,

because he was avoided even when mates were

_present.

In this context, Romanov females

_emerged

as

_being

more reactive and disturbed

_by

man in an

open field than M6rinos. The Romanov also took

longer

to

_approach

the human and were driven to the

_testing

pen with more

difficulty.

Because

they

also looked more often at the

observer,

it cannot be excluded that differences between breeds

in the absence of a human were

_partly

due to the _presence of the

_observer,

and not

only

to isolation and a new environment.

Consequently,

even

though

our results

appear to _agree with the

_hypothesis

that there is a relation between behavioural

reactivity

in an _open field and

_reactivity

to man, as

_suggested

_by

the results of

Romeyer

and Bouissou

_{(personal communication)}

on

fear,

this may still need to be confirmed.

The 2 breeds can also be

_{distinguished}

_by

a number of other factors. For

_example,

overall

_{olfactory investigations}

and vocal

_activity

were lower in Romanov than in

Merinos. This

_might

be related to the

_higher

_reactivity

of the

Romanov,

whose behaviour

_might

be more affected

_by

the

_anxiogenic

characteristics of the tests. In

support

of this

_suggestion

it can be noted that the

_disturbing

_presence of humans

was also associated with a reduction in

vocalizations,

mainly

due to a reduction in

high-pitched

bleats. This is in

_agreement

with the results of Price and Thos

₍₁₉₈₀₎

and Zito et al

_(1977),

who found that the presence of a human reduced

mobility

and

_{bleating. However,}

in contrast with the conclusion of Price and Thos

_(1980),

our results indicate that the role

of

a person as a substitute for penmates is not

appropriate

in our

_situation,

since females avoided the human even in the _presence of mates.

It is

_possible

that the breed differences found here do not reflect

_only

differences

in behavioural

_reactivity.

_{They might}

also _express some basic differences in the

relative

_importance

of the sensory cues characteristic of each

breed,

independent

of

_behaviour,

as

_already

shown for

_example

in mutual

_mother-young

_{relationships}

(Shillito

et

al,

1982).

It cannot be excluded for

example

that M6rilios are

simply

more vocal than

_Romanov,

without this

niarking

a

higher

_reactivity.

On the

other

_hand,

breed differences in vocalizations have also been found

_by

others

(Torres-Hernandes

and

Hohenboken,

1979),

with variations

_probably

associated with

_differing

levels of behavioural distress due to _age and

habituation.

This

The

_study

on crossbred animals sheds some

light

on the differences found in

the _pure breeds. Because crossbred females were born to M6rinos

_mothers,

the differences cannot be

_explained

_{solely by}

the influence of the mother. Otherwise the crossbreds would not have differed from the _pure

_M6rinos,

whereas in fact

they

were in many variates closer to their Romanov father than to their M6rinos mother. Thus _{it appears} that the differences

_reflect,

at least in

_part,

some

_genetic

variability

of

_general

_reactivity,

_including

behavioural

_reactivity

to the _presence of

man. Our results therefore

_suggest

that it

_might

be

_possible

to select animals on

the basis of their reaction to man in an

_attempt

to facilitate the

_management

of

sheep.

A better

_knowledge

of the

_weight

of the various

_parameters

_influencing

the measured

behaviours,

reaction to

_novelty,

social

_tendency,

and fear of man,

however,

would

_certainly

facilitate the

_design

of such a selection _programme.

_Similarly,

the

genetic

components

of the various behaviours

_reflecting

_reactivity

to man

(direct

and maternal

_effects,

_heterosis)

have still to be clarified.

ACKNOWLEDGMENTS

Thanks are due to the staff of the

_experimental

ENSA-INRA farm of Le Merle who _gave their support to the

_{study by providing}

the animals and technical assistance.

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