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S T U D I E S O N T H E F E E D I N G

A N D S O C I A L B E H A V I O U R

O F D O M E S T I C H O R S E S

Volmne II

Katherine Francis -Smith

Ph. D University of Edinburgh 1979 CONTENTS

Volume II Page

11 Figures 1

12 Plates 35

13 Bibliography u 7

14 Appendix 77

15 Appended Papers 106 11 FIGURES FIGURE 1 A plan of the fields and animal accommodation Field 0 the Veterinary Station Î NORTH

1 -7 Fields 8 Fenced track 9 Stalls 10 Stableyard 11 Indoor riding school 12 Veterinary Field Station 13 Animal Hospital 3

FIGURE 2 Ground plan of the stalls

9 m 4

FIGURE 3 The mean nursing time (+ sem) of foals, per daylight hour, recorded during Weeks 1 to 8

700

I 600

500

400 I

300

200

100

0 1 2 3 4 5 6 7 8

Weeks after birth FIGURE 4 The mean time (-I- sem) spent by foals, expressed as a percentage of the dams' eating time during Weeks 1 to 7

6o

50

40

30

20 I

I 10

Z

0 1 2 3 4 5 6 7

Weeks after birth FIGURE 5 The number of incidents of coprophagia expressed as a percentage of the number of mares' defaecations that were observed during Weeks 1 to 8

4o

20

10

5 6 7

Weeks after birth FIGURE 6 The mean percentage of the observation time that the mares and foals spent feeding, resting and

in all the other' activities in Weeks 1 to 24 100

á 80

o ó 60 d 4o N Ñ a,o 20 MARES 1111 11[1I F R '0' F R '0' F R '0' F R '0' F R '0' F R '0' F R '0' F R '0' F R '0' F R '0' F R '0' 0 o 20 24 1 2 2 3 4 6 8 12 16 (stable) (field) Weeks after birth 100

8o 4,O m + 6

a) 4o

2o

FOALS R '0' F R '0' F R '0' F R '0' F R '0' F R '0' F R '0' F R '0' F R '0' F R '0' F R '0' F 16 20 24 2 2 3 4 6 8 12 1 Weeks after birth (stable) (field) of time F = feeding Nursing time expressed as a percentage feeding R = resting '0' = other activities FIGURE 7 The mean frequency of changes in activity per hour recorded from five mare -foal pairs during the first seven days after birth

100

90

80 .r.,

0 70

1 t t t i. 1 60 / t / t % 1 ( 1 i i. 50 .. V i ¡ / i / Y I 40 I R `J 7') 0 30 a

o 20

10

1 2 3 4 5 6 7 Days after birth

Mares

Foals FIGURE 8 The mean frequency per hour with which mares and foals changed their activities in Weeks 1 to 24

>, 100 .,4-) 7 90 + U (6 808 r{ mo2 70 ,0 6o w tH o 50 U C'. o 4o 4) t-i 30

20 1 2 3 4 6 8 12 20 24 MARES Weeks after birth

100

+3 ., U (tl 80 . mo; 70 0 .0

ó ai 60 a o 50 U i". 40

G) FI 1 30

20 1 234 6 8 12 16 20 24 FOALS Weeks after birth

- ---- Stable

Field -'10 -

FIGURE 9 The negative exponential curve of intervals

in nursing activity recorded from five foals during Week 1

6.1 6.0

5.9

5.8

5.7 5.6

5.5

5.4

Y 5.3

5.2

5.1

5.0 4.9

4.8

4.7 4.6 1 21 41 61 81 101 121 141 161 181 201 221 .241 261

Interval length, i, secs x

y = loge fi FIGURE 10 The number of nursing bouts of different durations recorded from five foals

during A) Week 1 B) Weeks 6 to 24 inclusive 6o 6o

50

4o

30 q w o

F1 20 1 /------

10 .: . "//%%%% 10

.

A: laile 4444 404 4,4g..i./% ii B: 22 2 .2 :2 102 122 1 2 1.2 22 42 62 82 102 122 142 162 182 202 222 242 262 282 302

Nursing bout duration, secs Nursing bout duration, secs

Successful nursing bouts Successful and unsuccessful nursing bouts Unsuccessful nursing bouts - 12 -

Frequency of FIGURE 11 The mean (+ sem) A) Nursing duration per hour, B)

nursing bouts per hour and C) Nursing bout duration recorded during 7 hr

of observation on each of 5 foals during Weeks 1 to 24 800

700 I

600

r I r 100 I A:

1 2 3 4 6 : 12 20 24

Weeks after birth

I

I I r r r

B: 0 1 2 3 4 6 8 12 16 20 24

Weeks after birth m110 e

r0100 I

m 90 I ó 80

g 70 I I I

z 60

C: 50 1 2 3 4 6 8 12 16 20 24

Weeks after birth - 13

FIGURE 12 The mean time spent in 'interval' behaviour (-1- sem) expressed as a percentage of total =sing duration recorded from five foals during Weeks 1 to 24

IIl I

III I I I

1234 6 8 12 16 20 24 Weeks after birth - 14 -

FIGURE 13 The mean frequency of A) 'Resistances' and B)

'Pushes' per 100 sec of nursing behaviour recorded from five mare -foal pairs during Weeks 1 to 24 n o o 2.5 m 2.0

o 1.5

w A: 0 1 2 3 4 6 8 12 16 20 24 weeks after birth

2.5

2.0

1.5

1.0 0.5

B: 0 1 2 3 : 12 1. 20 24 Weeks after birth - 15

FIGURE 14 The time that Jemma spent in sucking, nosing

and 'interval' behaviour, expressed as a percentage of total

successful nursing time recorded during 7 hr observation in

Weeks 1 to 4

80

70

0 60 E r-I 50 0 ri 4o . ------

o 1 2 3 4 Weeks after birth

Sucking behaviour

Nosing behaviour

'Interval behaviour - 16 -

FIGURE 15 The mean percentage of nursing bouts ended by the dam

1234 i6 l'li Weeks after birth -17-

FIGURE 16 The mean time that mares spent standing still and

flexing a hind leg, expressed as a percentage of the mean nursing

duration in weeks 1 to 24

1 2 3 12 16 20 24 Weeks after birth

Standing still

- - - - - Flexing a hind leg - 18 -

FIGURE 17 The mean time, min, of mares and foals recorded during 7hr observation on each of five mare -foal pairs

(four pairs during Weeks 3,_ 4 and 24) in Weeks 3 to 24 after birth

350

300

250 200 ., E - 150 i a) E -P 100 ., 50 ,

o 3 4 6 8 12 16 20 24

Weeks after birth

Mares

Foals - 19 -

FIGURE 18 The mean number, + sem, of A) mare -to -foal and B) foal -to -mare

interactions recorded during 7hr observation on each of 5 mare -foal

pairs in Weeks 1 to 24

A:

Weeks after birth

260

240

220

200

180

160

140

120

loo

8o

6

20

B: 1 2 3 12

Weeks after birth

recordings made in the field

recordings made in the stable - 20 -

FIGURE 19 The mean frequency of mare -to -foal interactions per hour recorded on days 1 to 7 after birth

4o

30

20

10

0

1 2 3 4 5 6

Days after birth - 21 -

FIGURE 20 The mean time per hour that each mare and foal spent interacting with other mares and foals during Weeks 2 to 24

160

140

120

100 . . i . . 8o i

I %,`. 6o --

40

20

o 2 3 4 6 8 12 16 20 21+

Weeks after birth

Interactions between a mare and other mares and foals

------Interactions between a foal and other mares and foals FIGURE 21 The dominance hierarchy of the mares

POLLY > EMMA > GAY 2> FLATS PIN 2 KIRSTY

(Polly > Emma indicates that Polly was dominant to Emma; the dotted line indicates a temporary unsettled relationship with the arrow towards the submissive horse) - 23 -

FIGURE 22 The number of incidents of coprophagia recorded in Weeks 1 to 24, including adventitious recordings

1 2 3 4 6 8 12 16 20 24

Weeks after birth FIGURE 23 The mean percentage of 4hr observation, between 10.45 and 14.45, that each foal spent in activities shown in A) Week 24, B) 10:11:77, C) 11 :11 :77 D) 12 :11:77

50 50 50

40 4o 4o

30 30 30

20 20 20

10 10 10

A: 0 B: 0 D: 0 1 2 3 4 5 6 1 2 3 4 5 6

1. Standing alert 2. Pawing, bucking and running in circles 3. General activities 4. Resting 5. Grazing or eating hay 6. Nursing Eating hay FIGURE 24 The diurnal distribution of

Grazing behaviour patterns recorded from 5 thoroughbred

mares one 24hr in Resting standing during period February 1975

Resting lying

'Other' behaviour patterns

Emma %EIIIII=EMENIEff1-liM=%//._1111IhI IREIMI%/.IM . ;: Y:t-::: Mi 1%/ l'.1 Willow 1%=_ 1 , %=lif=1-1R5511 :.:.:..:'ì::r.;t: Kirsty NINIMIEN1E= MIME/ K® Polly i1 AI =1 =7011 i111141 Melody /. :':%ICIrINI

Ó7 68 10 41 11 1j 14 '0 1 t, 1l] 18 2b 21 2 3 cb di 02 o4 d5 66 0 Greenwich mean time

The times when the out to /r mares were turned grass and'brought in one of the and behaviour of a group of horses during FIGURE 25 The diurnal distribution grazing resting and 1976 24 hr period in each of the months June, July August

..OPPI

Gay VI. Flatspin II. Kirsty ONO INN Passion . . i Albert T

Emma ...... n. Gay .. Flatspin "POPO. "s...... ,..,.,a . Kirsty ...... N Dux "_POPO. .,. Passion ....,..4 /. 5¡ "POPO. i Albert T T

Emma ... .,...... _a .,. .... MI.AI,.. Gay irisa Flatspin .,,.. .i. ".5... ./I. /././. 0 Kirsty i '.5.4 Albert ,...... VOA T T

, , 1 I , 06 08 18 20 21 22 23 00 01 02 03 04 05 07 08 09 10 11 12 13 14 15 16 17 19 British summer time

Periods of grazing behaviour

Periods of resting behaviour

The times of dawn and dusk - 27 -

FIGURE 26 The frequency distribution of intervals in

grazing behaviour recorded from a group of horses during

one 24 hr period in each of the months June, July and

August 1976

260

240

220

200

180

16o

14o

120

100

80

6o

4o

20

o 10 20 30 40 50 60 70 80 90 100 110 120 130 Interval length, min - 28 -

FIGURE 27 The negative exponential curve of intervals in grazing behaviour recorded from a group of horses during one

24 hr period in each of the months June, July and August 1976

5

4

3

2

1

0 10 20 30 40 50 60 70 80 90 100 110 120 Interval length, min

y = loge fi n 29

FIGURE 28 A map of Field 1 showing the areas of lawns, roughs and paths in May 1976

71m

160m

180m

0 Lawns Li Roughs

Paths

v

66m - 30 -

FIGURE 29 The total dm weight (g) and the moisture content

1 ( %) of herbage samples collected from Field on the dates

shown

80

70

60

50

4o

30

20

10

o 4-, 4, CO m d) G) Hp, H w 9 0 h h h 4 4 ON Kl L` O -3' L N N N

4-, 4-3 !D [D 4) Z Hs óOD I'D h I'D 4 4 ON rn N- O N N N - 31

FIGURE 30 The mercury tilt switch at angles of 52° and 26° to the vertical

This figure represents the angle of the mercury switch when the horse's head B is in the positions shown in Plate 8 .

The mercury switch in the 'off' position. See Plate 8i.

Glycerine

61

Mercury

Brass electrode The mercury switch in the 'on' position. See Plate 8 ii. - 32 -

FIGURE 31 A mercury tilt switch shown in position in

a recorder box (actual size)

Integrated circuits

Aluminium box

Plastic tubing

Plastic T -piece

Brass electrode

6v battery FIGURE 32 The diurnal distribution of the grazing behaviour of two horses recorded

using the Grunch during one 24 hr period in August 1978

Adam

Cupid LL 11E11 1 11111111111111 T T

1 1 I Z T -1 -I T 1 I I 11+ 15 16 17 18 19 20 21 22 00 01 23 02 03 04 05 06 07 08 09 10 11 12 13 14

Periods of grazing behaviour T The time of dawn and dusk FIGURE 33 The diurnal distribution of grazing behaviour recorded from one horse, Kirsty

using the Grunch during seven 24 hr periods in December 1978

5. 12.78 G.12.78

7. 12.78 9.12.78

10.12.78 AIM 11.12.78 12.12.78 T

. . . , .. . ' , . . . . 14 15 16 1? 1 19 0 21 2 23 00 01 02 d3 04 05 06 07 d8 09 10 11 12 13 14 - Periods of grazing behaviour Greenwich mean time

'Ì -The times of dawn and dusk 12 PLATES - 36 -

PLATE I Chance eating faeces recently deposited by Melody -37-

PLATE 2 Flatspin and Bashful in typical nursing posture: Bashful is in the reverse parallel position and

Flatspin has her right hind leg flexed -38-

PLATE 3 Emma showing flehmen while standing in the oestrous posture -39-

PLATE 4- Field 1: The hay area and surrounding pasture heavily soiled with faeces -40 -

PLATE 5 Field 5: Emma, while grazing a lawn, pushes faeces aside to graze underneath - 41 -

PLATE 6 (i) Field 1, June 1976; in the foreground is a rough - 42 -

PLATE 6 (ii) Field 1, July 1976: the roughs are less abundant than in June -43 -

PLATE 6 (iii) Field 1, August 1976: the roughs have been grazed and in some areas the herbage is less than 5 cm tall

me' PLATE 7 A horse wearing a headcollar with a pouch containing a recorder box -45

PLATE 8 The angle of the longitudinal axis of the recorder box to the vertical i) when the horse is standing = 52° -46-

PLATE 8 ii) When the horse is grazing = 26° 13 BIBLIOGRAPHY -48 -

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14 APPENDIX -78-

14:1 Analytical procedure for feed /faeces samples

Note: All containers, flasks, etc. used to contain reagents or samples must be clearly marked as follows:

1. Name of analyst

2. Nature of contents

3. Date

The analysis of a feeding stuff usually involves what is known as a "proximate analysis ". Such an analysis determines the amount of water, ash, crude protein

(C.P.), ether extract (E.E.), modified acid detergent

fibre (M.A.D.F.) in the feed. The total of these sub-

tracted from 1000 gives the amount of nitrogen -free

extract (N.F.E.), which is included in the proximate

analysis.

Sampling and Preparation for Analysis

Before undertaking an analysis the results of which are

to be used to represent the composition of a crop or

a feeding stuff, you should assure yourself that the

sample originally taken was sufficient in amount and

that it was selected properly from the bulk so as to be

fairly representative of it.

The sample supplied must, if necessary, be dried

sufficiently to enable it to be finely ground and should

always be well mixed before withdrawing any portion -79- for analysis.

1. Dry Matter (D.M.) determination

When a sample is received in a wet state it must be weighed into a tray (large for roughage, small for concentrates) and placed in a 100° oven for 48 hours to dry. When dry it is reweighed and the loss in weight gives the moisture content.

Fresh weight - Dry weight x 1000 Fresh weight

g Moisture /kg Fresh sample

The sample is then milled to a fine powder to facilitate the taking of representative samples of material. Un- fortunately, finely divided materials tend to be hygro- scopic and it is impractical to keep milled samples out of contact with the atmosphere. We must take account of this inevitable uptake of water from the atmosphere and this is conveniently carried out routinely in the ash determination.

2. Ash, Mineral and Silica determinations

About 2 g of milled sample are weighed accurately into a weighed silica basin and placed in a 100° oven for 48 hours. Then the basin is .placed in a dessicator, allowed to cool, reweighed and the loss of weight gives the moisture which has been absorbed after milling. - 80 -

Weight of dried milled sample x 1000 Weight of milled sample

= g DM/Kg Milled sample

This figure can be used to correct the results obtained

for E.E., M.A.D.F., C.P. and T.P. which are calculated

on an "as analysed" basis which ignores the presence of

water which has been absorbed by the milled sample.

i.e. Ash determination reveals a dry matter content in

the milled sample of 980 /Kg milled sample.

Ether extract obtained was 20g /Kg milled sample.

Thus to express the E.E. on a D.M. basis we perform the

following calculation:

1000 20 x = g E.E./Kg D.M. 980

The basin is now placed in a muffle furnace at 550° until

ashed (overnight). When ashed it is placed in a dessicator

to cool, then weighed to obtain the weight of ASH.

a. Weight of silica basin + Ash

Weight of silica basin Weight of ash b. Weight of ash x 1000 Weight of dried milled sample

g Ash /Kg dried milled sample

About 10 ml of concentrated hydrochloric acid is added to

the ash (to convert minerals to the chloride form) which

is evaporated to dryness on a water bath to dehydrate and - 81 - render the silica insoluble. About 10 ml of dilute hydrochloric acid is added to take up the soluble chlorides.

The contents of the basin are filtered through a Whatmans

No. 1 filter paper into a dl volumetric flask. The flask is made up to the mark with deionised water when cool.

This solution is now ready for mineral analysis.

Analysis of Ash solution a) Phosphorus

The stock standard is diluted 1/10 and 5 ml is taken for tube A. 5 ml deionised water is placed in tube B. An appropriate dilution of the ash solution is made and 1 ml taken and placed in tube C. 4 ml deionised water is added to tube C.

1 ml molybdate solution is added to tubes A, B and C and then 1 ml metol is added to each tube. All tubes are made up to a volume of 10 ml with deionised water and set aside for 30 minutes in the dark to allow maximum colour development and which is finally read at 680 m U.

Calculate as follows: mg /Kg = Strength of standard x Reading of test Reading of standard

x 1000 x Dilution factors Weight of sample

b) Calcium and Magnesium

Appropriate dilutions of the ash solution are made for - 82 - estimation using ATOMIC ABSORPTION.

The filter paper is then placed into the silica basin used for the ash determination and placed in a muffle furnace at 550° to ash. Then place in a dessicator to cool, and weigh to obtain weight of SILICA. a) Weight of silica basin + silica

Weight of silica basin Weight of silica b) Weight of silica x 1000 Weight of dry milled sample

g Silica /Kg dried milled sample.

3. Ether Extract (E.E.) determination

About 2 g of milled sample are weighed accurately into a

Soxhlet extraction thimble which is then plugged with cotton wool. A clean dry soxhlet flask is weighed.

The thimble is placed in the barrel of the soxhlet extractor and the flask filled 4full with petroleum ether (B. pt.

400-600). The sample is solvent extracted for 4 -6 hours at mantle setting 4.

The ether is distilled off and the flask dried in a forced draught oven at 60 °. Place in a dessicator to cool and weigh to obtain weight of E.E. a) Weight of flask + E.E.

Weight of flask Weight of E.E. -83- b) Weight of E.E. x 1000 Weight of milled sample

g E.E. /Kg of milled sample.

4. Modified Acid Detergent Fibre ( M.A.D.F.) determination

About 1 g of milled sample is weighed accurately into a flask. A dl of acid - cetyl trimethylammonium bromide

(C.T.A.B. solution) is added and the mixture is boiled gently under reflux for 2 hours. Filter hot through a weighed sintered glass crucible (porosity 1) under vacuum.

Wash residue with hot water and then acetone. Dry crucible and contents in a 100° oven overnight. Cool in a dessicator and weigh. This gives the M.A.D.F. a) Weight of crucible + M.A.D.F.

Weight of crucible Weight of M.A.D.F. b) Weight of M.A.D.F. x 1000 Weight of milled sample

= g M.A.D.F./Kg of milled sample.

5. Crude Protein (C.P.) determination

About 2 g of milled sample are weighed accurately into a

Kjeldahl flask and about 30 ml of concentrated sulphuric

acid and two crude protein catalyst tablets are added.

The flask is than placed on the rack and heated,

gradually at first and then at full heat. Digestion is

continued until the sample is clear and then for a further

20 minutes. - 84 -

The flask is allowed to cool and then the neck washed down carefully, gently mixing the water with the flask contents.

Leave to cool:

Transfer contents of digestion flask to a dl volumetric flask and make up to mark with deionised water when cold.

This solution is now ready for the estimation of its nitrogen content.

The particular form of apparatus used is the Markham

Distillation Apparatus which allows volatile substances

(in the present case - ammonia) to be steam distilled with complete collection of the distillate. Provision is made for the introduction of strong alkali (40 g /dl sodium hydroxide) to bring about the release of ammonia from the acid solution.

Steam out the apparatus for about 10 minutes before use.

Pipette 5 ml of sample solution into the body of the apparatus via the small funnel aperture. Place the tip of the condenser into 10 ml of boric acid plus indicator solution in a conical flask. When ready fill the funnel with sodium hydroxide and by lifting the funnel plug, run in the sodium hydroxide very carefully. Ensure excess sodium hydroxide is added. An indication that sufficient alkali has been added is a change in colour of the solution within the apparatus. To avoid "suck-back" remove the receiving flask from the condenser tip, but - 85 - quickly return it to its normal position as soon as possible. Continue the distillation for 10 mins, at the end of which time hold the flask clear of the condenser tip and allow for drainage from the condenser.

Titrate the contents of the receiving flask using 0.01 M hydrochloric acid. The boric acid, while preventing loss

of ammonia by the formation of ammonium borate, is a very weak acid and does not affect the indicator in any way. The titration may then be thought of as being essentially between ammonium hydroxide and hydrochloric

acid.

1 G.M.W. H Cl = 1 G.M.W. NH3

1 1 of 1.01M H Cl = 17 g of NH3 = 14g N

1 1 of 0.01M H Cl = 0.17 g of NH3 = 0.14g N

la of 0.01M H Cl = 0.00014 g N

= 0.14mgN

The g /Kg N is multiplied by 6.25 and the result is

expressed as C.P.

Therefore: Titration x 0.14 x 6.25 x 100 5

x 1000 x 1 Wt of milled sample 1000

= g C.P. /Kg of milled sample.

6. True Protein (T.P.) determination

The nitrogenous compounds of feedingstuffs, apart from

proteins include amides, amino- acids, and perhaps ammonium -86-

salts and nitrates. In root crops and green crops a particularly high proportion of nitrogen is present as non -protein nitrogen (N.P.N.).

To determine the T.P. advantage is taken of the fact that

the N.P.N. substances are soluble in water and that soluble

proteins are precipitated by certain metallic salts, e.g.

copper salts in alkaline solution.

About 2 g of milled sample are weighed accurately into a

2.5 dl beaker provided with a rubber ended stirring rod.

Add 75 ml of distilled water, heat to 60° and maintain

at that temperature for 10 - 15 mins.

Add 25 ml of 0.24M copper sulphate solution, stirring all

the time and heat again to 60 °. Add 25 ml of 0.3'M sodium

hydroxide solution with stirring, heat to 60° to

precipitate all the copper and the T.P. Filter hot

through a Whatmans No. 4 filter paper. Wash with hot

distilled water and after leaving to dry in the funnel for

several hours or overnight, the filter paper and contents

are transferred to a Kjeldahl flask. After the addition

of T.P. catalyst tablets, the procedure is the same as

that described for 0.P. - 87 -

14:2 To calculate the area of Paddock 1 covered by roughs, lawns and paths

Area of paddock = 12 730 m2

Wt. of 10 maps = 22.105 g

Wt of 1 map = 2.2105 g.

2.2105 g paper represents 12 730 m2

m2 Therefore 1 g of paper represents 12 730 2.2105

= 5759 m2

Wt of Wt of Are % of Total 10 (g) 1 (g) ( m ) Area

Roughs 13.838 1.3838 7969 63%

Lawns 7.361 0.7361 4239 33% Paths 0.860 0.0860 495 4% 1 :3

TABLE 1 The observed and the expected number of runs of 1) Grazing 2) Resting and 3) 'Other' behaviour patterns, shown by a group of horses during one period of

24 hr in each of the months June, July and August 1976

1) Grazing behaviour Emma Gay Flatspin Kirsty Dux Passion Albert Mean

w Obsd - 37 40 34 - 39 33 37 Expt - 63 64 63 - 58 61 62 S.D. 3.66 3.73 3.66 3.40 3.60 3.61

H Obsd 10 19 24 17 21 15 20 18 Expt 68 71 71 63 67 63 65 67 S.D. 3.98 4.15 4.17 3.71 3.90 3.66 3.81 3.91

20 24 - - m Obsd 19 23 27 23 0 Expt 54 62 65 58 - - 64 61 S.D. 3.17 3.62 3.79 3.40 - - 3.75 3.54 TABLE 1 (cont.)

2) Resting behaviour Emma Gay Flatspin Kirsty Dux Passion Albert Mean NObsd - 11 13 13 - 9 8 11 - Expd - 42 45 /I /I 31 35 39 S.D. - 2.46 2.61 2.57 - 1.79 2.01 2.29 H Obsd 7 10 13 9 9 6 7 9 61 Expd 64 66 65 59 61 56 57 S.D. 3.73 3.88 3.84 3.47 3.55 3.29 3.32 3.58 N mObsd 8 8 9 8 - - 8 8 - - 0Expd 45 51 53 40 /I /I 47 'S.D. 2.61 2.95 3.08 2.34 - - 2.57 2.71

3) 'Other' behaviour patterns Mean Emma Gay Flatspin Kirsty Dux Passion Albert

34 33 30 37 34 34 3 32 37 35 Expd 34 32.30 2.04 GbsdS.D. 1.9 7 1.9 3 1.84 .18 14 HObsd 8 14 21 9 15 12 17 16 12 18 23 9 15 13 19 Expd 1.07 0.89 S.D. 0.71 1.02 1.32 0.49 0.87 0.76 - - Obsd 16 20 21 27 30 23 - - 25 0Expd 15 21 25 28 35 - - 2.00 1 .42 S.D. 0.87 1.22 1.42 1.61 - 90 -

14 :4 To calculate the significance of differences in the time that mares or juveniles spent grazing lawns and roughs, and the proportion of paddock covered by lawns and roughs

The horses grazed the paddock at random spending a proportion of time on lawns and roughs similar to the proportion of paddock covered by these areas.

Under Ho, 1 - r = (grazing time on 1 in daylight x 100 %)

(total grazing time in daylight )

- (grazing time on r in daylight x 100 %)

(total grazing time in daylight )

= area of lawns x 100% total area of paddock

- area of roughs x 100% total area of paddock

= -30% TABLE 1 The mean 1 - r values for mares and two -year -olds and their t values for June, July and August

Mares Juveniles Residual t value n mean l -r n mean 1 -r mean square d.f. mares juveniles June 3 12.8 2 48.8 20.0/1115 3 16.5693 24.8847 July 4 5.4 3 62.2 495.8039 5 3.1801 7.1730 August 4 -28.3 1 80.0 37.11/114 3 0.5475 17.9755

= - t mean (1 -r) expected (1 -r) mean square iresidual - 92 -

14 :5 To calculate the difference between mares and juveniles in the time they grazed the lawns, 1, and roughs, r

Ho. There is no difference between mares and juveniles in the time they spent grazing the lawns and roughs.

Sample mean for mares = xl

for juveniles = X Sample mean 2

Under Rol x = x2 - Xi and will have a normal distribution with mean zero and variance ñ62. Analysis of Variance

TABLE 1 The values of 1 - r for mares and juveniles in June, July and August

Emma Gay Flatspin Kirsty Dux Passion Albert x x1 x2 June - - 15.6 15.2 7.7 12.8 'I JI .2 53.4 48.8 27.2 July 45.2 1.6 -7.6 -17.6 5.4 69.8 51.3 65.5 62.2 29.7 -32.7 -21.0 - - August -25.6 -34.0 -28.3 80.0 80.0 -6.7

1 - r = (grazing time on 1 in x 100 - daylight %) (grazing time on r in daylight x 100 %) (total time in grazing daylight ) (total grazing time in daylight ) -94-

TABLE 2 The F ratio for June

Sums of Squares d.f. Mean Square

Mares v juveniles 1550.8830 1 1550.8830 Group 1632.0164 4

Residual 81.1334 3 20.0'1'15

If Ho is true, the F ratio

Mares v juveniles mean square = 1 Residual mean square

The F ratio = 77.3720 d.f. = 1,3 * * -95-

TABLE 3 The F ratio for July

Sums of Squares d.f. Mean Square v Mares juveniles 5532.3196 1 5532.3196

Group 8011.3389 6

Residual 2479.0193 5 495.8039

The F ratio = 11.1583 d.f. = 1,5 * * -96-

TABLE 4 The F ratio for August

Sums of Squares d.f. Mean Square Mares v juveniles 9385.7114 1 9385.7114 Group 9498.0355 4

Residual 112.3241 3 37.111114

The F ratio = 250.6774 d.f. = 1,3 * * - 97 -

14:6 The technique used to estimate the minimum distance travelled by Gay during one period of 24 hr in

June 1976

1. Map plottings had been made at 5 min intervals during daylight hours.

2. On the map 1 mm represented 2/3 m.

3. The sum of the distances between successive map plottings for Gay in June = x mm = 8842 mm.

4. x represents 8842 x 2 = 5895 m. 3 5. During daylight hours Gay spent 1110 min active.

6. The horses were stationary when resting.

7. The rate of travelling = 5895 = 5.31 m /min activity. 1110 8. During 24 hr Gay spent 1185 min active.

9. The estimated minimum distance travelled by Gay in

24 hr = 5.31 x 1185 = 6300 m (to the nearest 100 m). -98-

14:7 The grazing recorder

The grazing recorder is a self -contained sampled data recorder using integrated electronic circuits powered by nickel -cadmium rechargeable cells. A mercury tilt switch

senses the attitude of the recorder, and provides a

signal which is registered in the memory at pre- determined

intervals and stored in there until the replay box is used to sequentially recall it for analysis.

14:7:1 Sample mode operation

Refer to Figures 14:1, 14 :2, 14 :3.

Before use, the memory (IC1) is cleared by disconnecting

the battery for a short time and the sample counter is

reset to zero using the replay box. With the sample/

hold switch in the 'hold' position, the internal clock

is inhibited and the memory is in its read mode. When

the switch is moved to the 'sample' position, the memory

enters its write mode and the clock (IC4) begins to

oscillate. The clock output is divided (Ic3 cd part

IC2) to provide 1 pulse per 2 min to the sample counter

(part IC2). At the end of each pulse, the signal from

the switch, which is applied to the memory input via

IC5c is recorded and the memory address is incremented.

When a count of 1024 is reached, further recording is

inhibited by IC6c and the signals are held for replay.

Recording may be stopped at any time by moving the switch

to the 'hold' position. - 99 -

14:7:2 Replay mode operation

Refer to Figure 14:3

The replay box is connected to the grazing recorder to

retrieve the recorded data. With the sample /hold switch

in the 'hold' position the replay box provides an external

count input to the sample counter which may be incremented

at will to display the switch signal as recorded for any

sample. The replay box also had a sampling indicator

which allowed the sampling time interval to be checked when

the box was connected to the grazing recorder while the

recorder was in the sample mode.

A mains power unit and a charger circuit for the nickel

cadmium cells used in the recorder were contained in the

replay box.

I:7:3 The switch monitor

Refer to Figure 14:4

The switch monitor allowed the signal from the mercury

tilt switch to be monitored while the recorder was in use.

A buffer circuit and lamp driver was used to avoid upsetting

the recorder's electronics which might be damaged by a

simple battery and lamp circuit. The monitor was powered

by its own rechargeable battery.

14:7:4 Problems with the present design and possible

improvements

The clock circuit in the present design consumed most of - 100 - the power (approximately 10 mA) and hence relatively large nickel cadmium cells were required. In addition, the sampling interval was difficult to set precisely and maintain over the normal operating temperature range. Since the

Grunch was designed an improved version of the clock IC has become available which would considerably reduce the current drawn to less than 0.5 mA and hence the battery

size and weight. A better solution however would be to use one of the more recently available wrist -watch type if integrated circuits and quartz crystal to give a very low power, frequency stable clock source. If this latter

option were used, the size of the recorder box might be reduced to approximately one third of its present volume.

A further improvement could be made by increasing the number of samples held in the memory. Integrated circuits

are currently available which would give sixteen times the memory capacity which could be used to either increase

the frequency of sampling or record several channels of

data simultaneously. In the event of the number of samples being increased improvements to the replay box would also be necessary. The latter could be adapted to connect to

a printing device to provide a hard copy of the results or, better still, a direct connection could be made to a suitable

computer which could then process the results as they are

retrieved. FIGURE 14+:1 Block diagram for the Recorder Box

SRMPkE M o 6 E_

Merloey 5*,vrcm Rh6RE .% NJPar ---+~ Se 1PLE < G.oct M?H pcY COWM R di I cE,c 1019 air

O __ ,N W1GTE

1

RE-NA y M oDE

.x--,-.

QH Ho-) r(O(

t I IOK 3 lo io Q11 D1 9 Tcb L IG S lc I. ICI

L

13 Z LA-SP. 11 J 10 i OutR+T e3 u _L8 4.i

`° " \ I i r- Cy Qcsuc 1 Cowl Q 1C6o- - - loK - - VL4 W,1t b01,1t t 6v Dl1CWp1L i

SnrIPtiL HOLD MONO FIGURE 14:3 Circuit diagram for the data replay box

i) Replay circuit +Sv -1r

I

ED. bek-rA ---; or.kleAT u t 4 12. a 1 II 44"\/ Ts_

oo -1ygo 7iD -moo I3 { 7 + 12 :1 ça i6,s r 4411 u1

't1PJ4 ii) Circuit for the power supply and battery charger -ig( -ro oP4 DiSPAY )1loct ÁtsCqY

J E FIGURE 14+ :4 Circuit diagram for the switch monitor

CD.4oI I 1 rJ RAT FQoM <

4 12.1.4.ickt <----- 15 APPENDED PAPERS FRANCIS- SMITH, K. (1977). Behaviour patterns of horses grazing in

paddocks. In: Proceedings of the Society of Veterinary Ethology

London, 8 December 1976. Appl. Anim. Ethol., 3, 293 -293.

Royal Dick Veterinary College, University of Edinburgh, Edinburgh

(Great Britain)

ABSTRACT

In June, July and August, 1976, a group of seven thoroughbred

horses were observed spending on average 16.68, 15.28 and 16.87

h/24h grazing, 3.17, 1.37 and 2.28 h/24h in other activities and

4.15, 7.35 and 4.85 h /24h resting. The increase in time spent

resting in July was attributed to higher average temperatures.

The average number and length of resting bouts per 24h was 11

bouts of 22 min in June, nine bouts of 48 min in July and eight

bouts each lasting 35 min in August. In July, each horse lay down,

on average, for 169 min in 24h compared with 55 min in June and

46 min in August. Grazing bouts lasted from 5 min to 10h 45 min,

with average values of 108, 125 and 140 min in June, July and

August, respectively.

The animals showed a definite choice of the areas called lawns for

grazing and the areas called roughs for excretion. They grazed the

roughs more in August than in June or July and this was believed

to be a result of a drop in available herbage. Three 2 -year olds

in the group were found to maintain a high grazing time on the

lawns throughout the summer compared with the adult mares. This

was attributed to either lower nutritional requirements or a greater aversion to grazing roughs (two of the 2 -year olds were geldings).

Social interactions showed that there was a linear hierarchy in the group. Animals stayed closest to the animals near them in the hierarchy; the 2 -year olds were the lowest ranking animals and they kept the largest distance between themselves and all the adult mares. FRANCIS -SMITH, K. (1978) The nursing behaviour of foals.

(Comportement d'allaitement chez les poulains) In: Proceedings of the First World Congress on Ethology as Applied to Zootechnics,

Madrid, 23 - 27 October 1978, 2, p. 97.

Dept. of Animal Health, Royal (Dick) School of Veterinary Medicine,

University of Edinburgh, Veterinary Field Station, Easter Bush,

Roslin, Midlothian, SCOTLAND, U.K.

SUMMARY

Foals are the 'following' type of ungulate offspring. They stay close by their mother throughout the day and are able to nurse at frequent intervals. The nursing behaviour of 5 thoroughbred foals was observed. The average time spent nursing decreased with age, the biggest change was during the first 4 weeks post- partum, after which the drop in nursing time with age was more gradual. A foal did not always suck when nursing. The foals were observed to suck from the teats in a series of bursts, usually swopping teats between bursts. Initially the mare helped her foal to nurse by standing still, stepping forward to bring

the udder to the foal's head and by flexing the opposite hind

leg. While nursing the foal pushed its nose into the udder before

and during sucking, this caused the mare to actively resist the

foal's attempts at nursing. The mare's 'resistances' and the

frequency of foal's 'pushes' reached a maximum in Week 11. The

foal reacted to frustration by pawing, cantering and kicking the

mare. RFSUME

Les poulains sont des progénitures ungulées du genre suivant. Ils restent près de leur mere pendant la journée et ils peuvent s'allaiter par intervalles fréquents. On a observé la conduite d'allaitement de cinq poulains de pur sang. Le temps moyen passé en allaitant s'abaissait selon l'aage du poulain; le changement le plus remarquable fut pendant les quatre premieres semaines après naissance; après cette période l'abaissement du temps passé en allaitant selon l'age fut plus graduel. Un poulain ne suçait pas toujours pendant l'allaitement. On remarque que les poulains suçaient les tetons'd'une poissée d'activité, en principe changeant les tetons entre ces périodes. D'abord, la jument aidait au poulain à allaiter en restant immobile, et elle s'avancait pour donner le pis au poulain tout en fléchissant la jambe arrière en face. Pendant l'allaitement le poulain poussait son nez dans le pis avant sucer, et mame pendant l'acte de sucer; cela causait la résistance active de la jument contre les tentatives d'allaiter.

La 'resistance' de la mère et la fréquence des 'poussées' du poulain furent au maximum pendant la deuxième semaine. Le poulain réagissait contre la frustration en grattant du pied, en allant au petit galop et en donnant des coups de patte á la mère. Equine vet. J. (1977), 9 (3), 155 -157 I

Coprophagia as seen in Thoroughbred Foals

KATHERINE FRANCIS -SMITH AND D. G. M. WOOD -GUSH* Department of Animal Health, Royal (Dick) School of Veterinary Studies, Easter Bush, Roslin, Midlothian

SUMMARY Four Thoroughbred foals were seen to quickly eat part of the faeces deposited by their own dams on some 40 per cent of the mare -defaecating occasions observed between the second and fifth week after birth. They did not do it before or after this period. This behaviour was thought to be a feeding pattern which formed a normal part of the foal's development.

INTRODUCTION Recordings were made on a portable cassette recorder COPROPHAGIA is a behaviour pattern recognised as The weeks post partum on which observations were occurring in the foal (Tyler, 1972; Taylor, 1954; Hafez, made are shown in Table I. The mare /foal pairs were Williams and Wierzbowski, 1969). It is seldom seen observed for a total of 234 hours. A total of 18 in the adult equine (Feist, 1971). The incidence and observation periods a week were made on each mare normality or abnormality of coprophagia is uncertain and foal distributed to cover the time between 08.00 h to and observation of young foals was considered necessary 17.00 h. Observations began on the day the foal was to investigate its nature. born (for foal 3 they began on Day 8 post partum). The observer sat at the side of the stable and aroused little MATERIALS AND METHOD curiosity from the mare after the first day. The foal treated the with The subjects were 4 Thoroughbred filly foals born observer the same curiosity that it gave the rest of the environment. between April and July 1976 (Table I). Each of the mares was subjected to the same husbandry, they all gave birth in a loose box bedded with wheat RESULTS straw, and all received a diet of 4.5 kg oats, 1 kg bran and A total of 25 incidents of coprophagia were recorded 7 kg hay a day, which the foals were able to share. In (Table II). In 17 cases the foal approached the faeces fine weather each mare and foal were turned out to immediately the mother had passed them and in 3 cases grass together for a few hours. within 30 seconds. The longest time a foal took to Observations were made during half hour periods react was 345 seconds. In 18 cases the foal ate the to record the behaviour of the mare and foal at 15 second faeces immediately and in 7 cases the foal sniffed and intervals and all details of incidents of coprophagia. pawed the pile for up to 75 seconds before eating (Table III). TABLE I TABLE II DETAILS OF THE FOALS OBSERVED LENGTH AND NUMBER OF INCIDENTS OF COPROPHAGIA SHOWN BY EACH FOAL Weeks Age Age Post First First Partem Age when No. of Average Range in Ate Ate when Foal Coprophagia was incidents length length Foal Date Sex Wt at Hay Oats obser- first and last seen (seconds) (seconds) of Birth (days) (days) vations Birth (kg) made 1 11 -32 days 4 169 105 -240 2 3 -20 days 12 170 60-285 3 10-31 days 5 125 75 -210 1 3.4.76 F 45.36 7 13 1 -7 4 6 days 4 161 75 -240 2 7.4.76 F 48.98 8 19 1 -6 -20 3 26.6.76 F 47.17 8 15 2 -8 4 9.7.76 F 32.65 3 17 1 -6 Each foal ate faeces in a characteristic manner. With its knees bent and mouth open the foal pressed its muzzle hard into the pile for up to 10 seconds before straightening * Present address: Agricultural Research Council's Poultry Research Centre, West Mains Road, Edinburgh, EH9 3JS. its forelegs and chewing a mouthful of faeces. The foal 2 TABLE III 4 cases the foal stopped eating hay or grass, in 8 cases RESULTS OF THE TIME ELAPSING AFTER THE MARE the foal stopped exploring and in 5 cases the foal was DEFAECATED, BEFORE THE FOAL APPROACHED THE standing still before eating faeces. Similarly, after PILE (A) AND THE LENGTH OF TIME THE FOAL EX- coprophagia, the foal engaged in one of these behaviour PLORED THE FAECES BEFORE EATING (B) patterns. In 3 instances the foal walked to its mother and sucked, in 6 cases it began to eat hay, oats or grass, in 11 cases the foal explored the stable and in 5 cases it A B walked to its mother and stood still. In one instance the foal was lying down when its Time No. of Time No. of mother defaecated, the foal stood up immediately and (seconds) cases (seconds) cases sucked then ate the faeces. It was impossible to estimate the amount of faeces eaten each instance. 0 -15 17 0 -15 18 15 -30 3 15 -30 5 DISCUSSION 30-45 1 30-45 I Taylor (1954) believed that the presence of strongyloides 75 -90 1 60-75 I eggs in the foals' faeces meant that foals ate faeces from 180 -195 1 adult horses. It has since been shown that Strongyloides 300 -315 1 westeri larvae are passed to the foal in the mare's milk 345 -360 1 (Lyons, Drudge and Tolliver, 1973). However, Taylor also suggested that foals ate adult horse faeces to acquire flora for their intestine, a theory sup- chewed slowly with pauses of up to 12 seconds between the proper bacterial et al. (1969). Tyler (1972) saw New chews dropping lumps of faeces that were stuck round its ported by Hafez, one day to 14 weeks eating mouth as it chewed. When the incident ended the foal Forest pony foals aged from most common at 3 -4 weeks moved away from the faeces to engage in one of a number faeces and noted that it was the faeces eaten nearly always other activities. In 6 cases the foal continued chewing old. Tyler stated that of twice a foal was seen after it had walked away from the pile. This time was belonged to the foal's mother but included in the incident length. eating its own faeces. not to eat faeces during In all cases except one the foal ate from a pile of Zebra foals have been reported 1972). Adult horses faeces once only, when the incident had finished the their first few days of life (Klingel, eat faeces through hunger (Feist, 1971) foal showed no more interest in the same pile. Foal 3 occasionally captivity (Altmann, 1969). made the exception. After eating faeces for 75 seconds or boredom as a result of as an abnormality in the foal was startled by a dog and walked to mare 3, Coprophagia has been reported piglets. There is no record of this behaviour in 195 seconds later foal 3 returned to the same pile and young ate for a further 30 seconds. ungulates. foal did not react to all the faeces passed by its The 4 foals observed showed coprophagia between Each ate mother. Table IV shows the number of defaecations 1 and 5 weeks post partum, during this time they freshly passed faeces. between and including the first and last one eaten by from 41 per cent of "available" its mother only ate each foal, and the foals' reactions to the piles. Foals Each foal ate fresh faeces from and In most cases (20) copro- ate from 25 (41 per cent) of 61 piles of freshly passed from a pile of faeces once. the mother defaecated. faeces. Of these 25 incidents, 3 occurred in the field. phagia was stimulated when These did not appear to be any different to incidents in Each incident appeared to be purposeful and the foal the stable. Foals did not show any interest in their ate the faeces in a distinctive manner, unlike feeding own faeces, or those of other mares when together in behaviour when eating hay, oats or grass, or exploratory the field other than sniffing the piles if they grazed nearby. behaviour. When exploring, the foal nosed, sniffed, Similarly the mares occasionally sniffed faeces from licked and chewed all objects in its environment, it was their foals or from other mares with no other associated easily distracted and would return to the same object behaviour patterns. many times. Incidents of exploratory behaviour lasted To eat faeces each foal interrupted another behaviour from a few seconds to 5 minutes and were interrupted by and standing behaviour pattern, in 8 cases the foal ate faeces after sucking, in cantering, sucking, eating patterns. TABLE IV The term coprophagia implies that the foals swallowed NUMBER OF DEFAECATIONS FROM EACH MARE the faeces. The only evidence that the foals did swallow BETWEEN AND INCLUDING. THE FIRST AND LAST faeces was the fact that they chewed them, then, in some ONE EATEN BY EACH FOAL cases moved on to another ingestive behaviour; the foals were never seen to drop mouthfuls of chewed faeces in the way they sometimes dropped chewed hay. Foal's Reaction Tyler (1972) said that foals ate parts of 1 or 2 faecal pellets. Mare No. of No. No. No. Coprophagia may be a feeding -exploratory activity Defae- ignored explored eaten and this idea cannot be completely dismissed until cations only more is known about the behaviour. The indications that it is not exploratory are: I 19 11 4 4 (i) The foal ate its mother's faeces only. 2 19 6 1 12 (ii) The foal ate from a pile once only. 3 14 8 1 5 (iii) Foal 2 which showed 12 incidents of coprophagia did 4 9 2 3 4 not score any higher on total exploratory time than the other foals. 3 Although only 4 mare /foal pairs were observed, Klingel, H. (1972). Das Verhalten der Pferde (Equidae) coprophagia had appeared in each foal before the end Handbuch der Zoologie. 10, 1 -68. of the 2nd week post partum and was not seen after the Lyons, E. T., Drudge, J. H. and Tolliver, S. C. (1973). On the end fifth week, life -cycle of Strongyloides westeri in the equine. J. Parasit. of the because of this and the fact that 59,780 -787. the foals ate from 41 per cent of fresh faeces it is possible Odberg, F. O. and Francis- Smith, K. (1976). A study on that coprophagia is a normal part of a foal's development, Eliminative and Grazing Behaviour -The Use of the Field which, if an ingestive behaviour, may introduce bacterial by Captive Horses. Equine vet. J. 8, 147 -149. flora into the foal's gut and supply certain which Odberg, F. O. and Francis -Smith, K. (1977). Studies on the the foal is unable to synthesize. formation of ungrazed eliminative areas in fields used by horses. An aversion to eating faeces or eating near faeces is Appl. Anim. Ethol. 3, 27-34. present in adult horses (Odberg and Francis-Smith, 1976; Taylor, E. L. (1954). Grazing behaviour and Helminthic Disease. 1977) and coprophagia shown by adult horses is generally Brit. J. Aninz. Behay. 2, 61 -62. considered to be an abnormal behaviour. Tyler, S. J. (1972). The Behaviour and Social Organization of the New Forest Ponies. Anim. Behay. Monographs 5, 85 -196. ACKNOWLEDGEMENT The authors wish to thank Dr. P. Imlah for his critical reading RÉSUMÉ of the manuscript. On pense qu'il s'agit plus d'une attitude nutritionnelle REFERENCES que d'un comportement exploratoire du monde extérieur. Altmann, D. (1969). Cited by Habenburg, L. (1971). Verhalten bei Einhufern. Neue Brehm-Bucherei No. 427. Wittenberg - Lutherstadt: Ziemsen. ZUSAMMENFASSUNG Feist, J. D. (1971). Behaviour of Feral Horses in the Pryor Vier Vollblutfohlen wurden bei der Aufnahme von Mountain Wild Horse Range. Master's thesis. University frischem Kot ihrer Mütter einer of Michigan. während beschränkten Zeitdauer beobachtet. Dieses Verhalten wurde als Hafez, E. S. E., Williams, M. and Wierzbowski, S. (1969). Fressverhalten interpretiert als The Behaviour of Horses. In: The Behaviour of Domestic und nicht "Entdeckungs- Animals, ed. E. S. E. Hafez. 2nd edition. Bailliere, Tindall verhalten" und als normalen Bestandteil der Fohlenent- and Cassell, London. wicklung betrachtet. Accepted for publication 29.4.77