Equine Energetics. II. Energy Expenditure in Horses During Submaximal Exercise J. D. Pagan and H. F.

Hintz J ANIM SCI 1986, 63:822-830.

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EQUINE ENERGETICS. II. ENERGY EXPENDITURE IN HORSES DURING S U B M A X I M A L EXERCISE

J. D. Pagan a and H. F. Hintz 2'3 Cornell University, Ithaca, NY 14853
ABSTRACT

Energy expenditure was measured in four geldings (433 to 520 kg) during submaximal exercise on a racetrack using a mobile open-circuit indirect respiration calorimeter. A total of 304 5-min measurements of O2 consumed and CO2 produced were taken. Measurements were made with and without riders. The amount of energy expended by the horses was exponentially related to speed and was proportional to the body weight of the riderless horse or the combined weight of the horse plus rider and tack. Total energy expended by the four hones walking, trotting, cantering was best described by the equation: Y = e 3.02+-0065X where Y = energy expended (cal.kg'l.min-t) and X = speed (m/min). Digestible energy (DE) required above maintenance was calculated as DE (kcal.kg-t.h -t) -- e3"~ .57 (Key Words: Energy, Horses, Exercise, Oxygen.) -- 13.92 X .06.

Introduction

Materials and Methods

The main productive function in horses is work. This work may vary from high speed racing at speeds over 1,150 m/min to endurance rides, where horses may travel 167 km at a rate near 250 m/min, to draft work where horses pull or carry heavy loads at slow speeds. In all these types of work, energy is the dietary factor most likely influenced by exercise. The National Research Council estimated energy requirements for various activities (NRC, 1978), but these estimates were based on limited data. The following studies were conducted to measure energy expenditure during submaximal exercise (walking, trotting and cantering) in horses. Feeding standards were developed from these data.

1Present address: Manna Pro Corp., 4929 Wilshire Blvd., Los Angeles, CA 90036. 2The authors gratefully acknowledge the assistance of Terry Kinsman, Bill Tutton, Lisa Lee, Pamela Livesay-Wilkens,L. V. Sodetholm and Janice Williams. ~Send reprint requests to H. F. Hintz, Cornell University, Ithaca, NY 14853. Received August 16, 1985. Accepted April 8, 1986.

The energy expenditure of horses during submaximal exercise was measured using a mobile open-circuit indirect respiration calorimeter (figure 1). The calorimetry system was validated using a nitrogen dilution technique similar to the type used by Fedak et al. (1981). The system was found to measure oxygen consumption with an error of less than 2.0%. A total of 304 5-min measurements were made on four geldings (Quarter Horse weighing 433 kg; grade horse, 490 kg; Appaloosa, 506 kg; Thoroughbred, 520 kg). Each horse's oxygen consumption and carbon dioxide production was measured at rest and over a range of speeds varying from 40 m/min to 390 m/min on an 800-m oval track with a stone-dust surface. Step frequency was measured at each speed and was used to calculate stride length. Measurements were made on the horses both with and without rider. The weight of the rider plus tack averaged 59 kg. During a measurement of 02 consumption and CO2 production, the expired gas from the horse was collected using the face-mask shown in figure 2. This mask was held on the horse's head with a leather strap with velero fasteners. An airtight seal was formed between the mask J. Anim. Sci. 1986.63:822-830

822

EQUINE ENERGETICS--EXERCISE

823

s a m p l e for analysis

--I~ FNO~
MASK

H081~

Figure 1. Calorimeter used to measure energy expenditure during exercise.

"4( 4 . . . . . .

OUTEIOE EXPIRED

AIR GAE

Figure 2. Face-mask used to collect gases during exercise.

and the horse's muzzle with inflated blood pressure cuffs. The mask contained two oneway intake valves (V1 and V2) and one oneway outlet valve (V3). Outside air was drawn through the mask with a vacuum pump (figure 1) and all expired gases were directed through a 10-cm diameter flexible hose into the calorimeter, as indicated by the arrows in figures 1 and 2. A continuous aliquot of the gas passing through the calorimeter was collected and stored for analysis of oxygen and carbon dioxide concentration. The total volume of gas passing through the calorimeter also was measured, along with its temperature and relative humidity. The gas volume was adjusted to standard temperature and pressure of dry gas, and multiplied by the change in composition of the collected gas from that of inspired (outside) air to obtain the amount of oxygen consumed and carbon dioxide produced during a measure-

824

PAGAN

AND

HINTZ
80

ment. Energy expended was calculated by multiplying the number of liters of oxygen consumed during a measurement by the oxygen's thermal equivalent (kcal/liter) at the respiratory quotient (RQ) calculated for each measurement (Brody, 1945). The calorimeter was carried on a wagon pulled by a tractor. Electricity to power the pumps and clock on the calorimeter was supplied by a portable gasoline generator, also carried on the wagon. The horse being measured walked or ran alongside the wagon and was attached to the calorimeter by a plastic flexible hose connected to the face-mask. The hose was 2.4 m long and 10 cm in diameter. During each trial the horse wearing the facemask was led by a rider on another horse to ensure that it maintained a constant speed. This was the case even when a rider was on the horse being measured because the face-mask precluded the use o f bit and bridle. The horse and wagon were brought to the speed at which the measurement was to be taken in a particular trial about 1 min before the 5-min measurement period began. Total distance traveled during the 5 min was measured with an electronic bicycle odometer, and this distance was used to calculate speed. Step frequency was recorded by an observer with a hand held counter.
Results and Discussion

433-kg

Quarter (no rider)

Horse
9

65 .,A a 5O &2 9 2

2 35 9 A 6J 9 &~uL 9 Y: e 2 . 1 8 + .0063X

~6
2O G Ul 9u, A 5 J62 A

e 0

e 100

i 200

i 300

i 400

i 500

Speed

(meterslmln)

Figure 3. Relationship between energy expended and speed in a 433-kg Quarter Horse with no rider. Triangles indicate one value, numbers indicate number of identical values.

b o d y mass. They also found that oxygen consumption increased in direct proportion to mass supported by the muscles. A horse sustaining a load at rest increased its energy expenditure over resting without a load to a degree proportional to the weight of the added load (Armsby, 1908). While traveling at

Energy expenditures of the horses with and without riders are shown in figures 3 to 10. The relationship between energy expenditure and speed was best described as an exponential function. The data were transformed by taking the natural log (In) of energy expenditure regressed against speed. Linear regressions of the data from each horse with and without a rider are shown in table 1, along with the exponential equations. More energy was expended by each horse when carrying a rider than when unweighted. The amount of energy expended by the horses was related to speed and also appeared to be proportional to the body weight of the riderless horse or the combined weight of the horse plus the rider. In other words, a 450-kg horse carrying a 50-kg rider would expend about the same amount of energy as a 500-kg horse carrying no weight. Taylor et al. (1980) measured the energetic cost of carrying loads in rats, dogs, humans and horses. The loads ranged between 7 and 27% of

100433~kg Quarter Horse

c E I o b~

80-

(with rider)

60-

dl&

| ~ x LU 40& 9 Y: 9 ~j 202 . 4 5 + .0062 X

&&A

O-

I
70

t
140

I
210

I
280

J
350

I
420

Speed

(meters/rain)

Figure 4. Relationship between energy expended and speed in a 433-kg Quarter Horse carrying a 59-kg rider. Triangles indicate one value, numbers indicate number of identical values.

EQUINE ENERGETICS-EXERCISE 90 m/min, a load of 127 kg caused about an 8%

825

increase in the amount of energy expended (per kg of total mass). Because the energy expended was proportional to the body weight of the riderless horse, or combined weight of the horse plus rider, all
data could be combined. Linear and exponential equations for the combined data (figure 11) are shown in table 1. Stride length was highly correlated (P<.01)

The equations with and without rider were not significantly different. The energy expenditure measurements made during exercise in the present study are net energy expenditure values. They represent the a m o u n t of heat produced from the oxidation

with speed, with or without a rider (table 2).

= 100 490-'kg (no Grade Horse I 65

506.kg

Appaloosa

444A& & 9 9A 9

(no

rider)

9 & 50
-o

2 2

9 9

c E m o

80

rider)
4& 3 9 2

22 9149
o Q. x ul

&9
35

~t
9 v : e2 " t r x

80 *o 9 x uJ 40 a& 2

a I11

20

9 4& 2

2 4& 9

,tA 9149 uJ 20 2 0 i 0

,t 0

"~4 9
23

y s2"30

..

,.
:Speed

2,.
(maters/rain)

~;

,~

Speed

Figure 7. Relationship between energy expended and speed in a 506-kg Appaloosa with no rider. Triangles indicate one value, numbers indicate number of identical values.
(mat ers/caln)

Figure 5. Relationship between energy expended and speed in a 490-kg grade horse with no rider. Triangles indicate one value, numbers indicate number of identical values.
100 506*kg E 100 4~)0.kg grade
e o

Appaloosa

(with
80

rider)

horse
2 60 9 3
w

Iwith r iderl
E a o v 80

60 x uJ 40

40

g
20.

9 4A

2AA 9 y: 2.43 + .0063X

20 60 120 180 8Dead 240 (metarslmla) 300 360

ilO

180

240

320
(meters/rain)

~o

Speed

Figure 6. Relationship between energy expended and speed in a 490-kg grade horse carrying a 59-kg rider. Triangles indicate one value, numbers indicate number of identical values.

Figure 8. Relationship between energy expended and speed in a 506-kg Appaloosa carrying a 59-kg rider. Triangles indicate one value, numbers indicate number of identical values.

826

PAGAN AND HINTZ in zero energy balance (Pagan and Hintz, 1985). Total energy expended by the four horses walking, trotting and cantering on the racetrack was best described by the equation: Y = e 3.02+.0065X where Y = energy expended ( c a l ' k g - l ' m i n -1) and X = speed (m/min). The amount of net energy (cal'kg -l"min-1) expended above maintenance at each speed can be calculated as (e3.02 +.0065x ) _ 13.92. The efficiency of utilization of digestible energy (DE) by the horse for work must be known before DE requirements can be calculated. Assuming that body fat is the primary substrate for energy generation in horses during low intensity work, then the efficiencies of utilization of DE for fat production can be used. Respiratory quotients measured in this experiment indicated that fat was the major substrate for energy generation. The efficiency of utilization of DE for gain (fattening) in horses was estimated to be 55% in horses fed a 75% alfalfa meal-25% oat diet (Pagan and Hintz, 1986). Hoffman et al. (1967) reported an efficiency of 59% for fat production in horses from DE in diets consisting of 60% meadow hay, 20% oats, 10% wheat bran and 10% flaxseed meal. It therefore appears that a value of 57% might be reasonable for the efficiency of utilization of DE for submaximal work. This efficiency value will depend on the type of diet fed, and will tend to be lower on high roughage diets and higher in diets containing high levels of concentrate or fat (Blaxter, 1962; Kane et al., 1979). Digestible energy (cal,kg -1.min -1) required above maintenance can be calculated as e 3"02+'0065X - 13.92

of substrate to supply energy for locomotion and the maintenance of body functions. The amount of net energy expended by the animal for locomotion alone can be calculated by subtracting the amount of energy expended by the animal while standing at rest. A 500-kg horse would be expected to expend 13.92 cal'kg -1. min -1 while standing with minimal movement

80. 520~kg A G E i o v v 65

Thoroughbred

(no rider)

&9

50
4) cL x t~

35

92 9 A&

e c W

20

9 9

y: e2.30 ~ .0064 X

i 0 90

i 180

i 27O

J 36O

t 45O

Speed

(metersIml)

Figure 9. Relationship between energy expended and speed in a 520-kg Thoroughbred with no rider. Triangles indicate one value, numbers indicate number of identical values.

80 c E o v 520~kg Thoroughbred 9

65

(with rider)

CL X UJ y:e2-66+.0060X
~ o c ui 36

.57 To express this in the units of k c a l ' k g - l " h -1, this equation is multiplied by .06. Calculations using this equation at various speeds are shown in table 3. An example of how this equation combined with the maintenance DE equation [DE (kcal/ d) = 1,375 + 30.0 (W)] proposed by Pagan and Hintz (1986) can be used to calculate energy requirements for exercise is as follows: A 450-kg horse is ridden at a medium trot (250

20 i i i i t i

Speed ( m e t e r s l m l n )

mad speed

Figure 10. Relationship between energy expended in a 520-kg Thoroughbred carrying a 59-kg rider. Triangles indicate one value, numbers indicate number of identical values.

EQUINE ENERGETICS--EXERCISE
&

827

1:I

I:::l 0

,,.-~ ,~.

m~r

m ' ., ~

ed II

II

II

II

II

II

II

II

II

Z <

I:l,,

8
1.1

z~
Z~
~ 247
~

~
~

I::I

II

II

II

II

II

II

II

II

ii

5
Z~ Z~

i,d

Ill

~z
N
Z < <
z
M

+
II l::l o~

0
M

i.,,

< [..

i,.,

i-,

"T

~L~ "~

,-;

.7

0 ..c:l

ill 0

828

PAGAN AND HINTZ TABLE 2. RELATIONSHIP OF STRIDE LENGTH TO SPEED

All D a t a

c m

4uk A A AA A 9

Horse Quarter Horse NR a WRb Grade NR WR

Equation

Rz

tO 0

U433S 9 ~6,1 323

2255 &&

SLc -- 1.14 + .O060Xd SL = 1.27 + .0051X SL = 1.30 + .0049X SL = 1.09 + .0057X SL = 1.03 + .0063X SL = 1.16 + .0057X SL -- 1.03 + .O070X SL = 1.17 + .O060X SL -- 1.17 + .0058X

.97 .95 .92 .97 .98 .96 .98 .97 .94

'0 C

IlO0 C W

~ 40.

y . e3.O2 + .0085 x

Appaloosa NR WR Thoroughbred NR WR Combined aNR = No rider. bWR -- With rider.

7 o.
I o I Ioo
Speed

I 2OO

I 30o

(meterslmin)

Figure 11. Data representing 304 energy expenditure measurements from four horses weighing 433 kg, 490 kg, 506 kg and 520 kg. Each observation is from a 5-min measurement. The measurements were made on the horses both with and without riders, Measurements without riders were divided by the horse's body weight, while those taken with riders were divided by the horse's plus the rider's weight. Numbers represent the number of observations falling on the same location on the graph. (+) represents more than nine observations in the same location. Triangles indicate one value, numbers indicate number of identical values.

cSL = Stride length, m. dx = speed (m/min).

m/min) by a 75okg rider for 2 h. The horse's maintenance DE requirement equals 14,875 kcal/d [1,375 + 30.0 (450)]. The extra DE required for exercise would equal 9.49 kcal" kg - ~ ' h -1. The weight of the horse plus rider equals 525 kg, so the total DE required above maintenance equals 9.49 x 525 x 2 = 9,965 kcal. The horse's total daily DE requirement would equal 24,840 kcal. NRC (1978) energy recommendations for work are also shown in table 3. These recommendations were based primarily on the estimates given by Hintz et al. (1971). They measured feed intake and the kind and amount o f activity in two groups of horses. One group consisted of nine polo ponies used by the Cornell Polo Team; the other group was seven horses used in equitation courses. Estimates o f DE requirements for walking were calculate d by Hintz et al. (1971) using reports by Brody (1945) and Armsby (1903) that horses required

1.33 times as much energy for walking (59 m / min) as for standing. Digestible energy for walking was therefore calculated as maintenance DE times 1.33 minus maintenance DE. This resulted in a DE requirement above that for maintenance for walking of .5 k cal 'k g -1" h -1. However, this interpretation was not correct. Brody (1945) showed that horses walking at 59 m/min had about a 133% increase in metabolism in addition to standing, or a 2.34fold increase in energy expenditure. The 1.33 increase was a net increase over standing with the maintenance requirement already accounted for. This magnitude o f increase in energy expenditure at a 59 m / m i n walk was 1.8 k c a l ' D E k g - l " h -1 above that required for maintenance. This value is close to the 1.7 kcal D E . k g - l . h -1 requirement predicted from the present study (table 3). A walking speed of 59 m / m i n (3.5 km/h) is a speed that might be used by a team of draft horses plowing a field or pulling a heavy wagon. We observed, however, that riding horses tended to walk at a much faster pace, at speeds closer to 95 m / m i n (5.8 km/h). At these speeds the horses would be expected to require 2.5 kcal DE~ - l " h -1 above maintenance. Walking at this speed for 8 h would increase the horse's

EQUINE ENERGETICS--EXERCISE

829

T A B L E 3. DIGESTIBLE E N E R G Y (DE) REQUIREMENTS ABOVE M A I N T E N A N C E A T VARIOUS SPEEDS SUGGESTED IN THIS PAPER COMPARED WITH N A T I O N A L R E S E A R C H COUNCIL (NRC) RECOMMENDATIONS
DE.h -l

.kg -1

Gait
Present s t u d y

Speed, m/min

b o d y wt a, kcal

Slow walk Fast walk Slow trot Medium trot Fast trot/slow canter Medium canter NRC Walking Slow trotting, s o m e cantering

59 95 200 250 300 350

1.7 2.5 6.5 9.5 13.7 19.5

Fast trotting, cantering, some jumping Cantering, galloping,jumping Strenuous effort (polo, racing at full speed)
aweight o f horse plus and tack.

.5 5.0 12.5 23.0 39.0

DE requirement by 62% over maintenance. The NRC prediction for walking 8 h would only be a 12% increase over maintenance. The NRC divided activity into five categories: 1) walking; 2) slow trotting, some cantering; 3) fast trotting, cantering, some jumping; 4) cantering, galloping, jumping and 5) strenuous effort (polo, racing at full speed). These are arbitrary classifications of activity and make comparisons with the present study difficult. However, by classifying the activities from the present study into slightly less arbitrary categories, a comparison can be made. Table 3 contains DE requirements from the present study divided into six speeds. These are a slow and fast walk, slow, medium, and fast trot, and a medium canter. These divisions represent the gaits maintained at each speed by the four horses in the study. Obviously, different breeds will vary from these averages. For instance, some Standardbreds can trot at speeds up to 800 m/rain. At a slow trot in the present study (200 m/ min) the horses would be expected to require 6.5 kcal DE'kg -1 ~ -1 above maintenance. This is fairly close to the 5.0 kcal D E . k g - l ' h -1 recommended by the NRC for slow trotting with some cantering. A fast trot/slow canter (300 m/min) would require 13.7 kcal DE-kg -1h -1 in the present study compared with the 12.5 kcal D E ' k g - l - h -1 recommended by the

NRC for fast trotting, cantering and some jumping. A horse at a medium canter (350 m/ min), which is as fast as could be accurately measured in the present study, would be expected to require 19.5 kcal D E ' k g - l - h -1. The NRC recommends 23.0 kcal DE.kg-l.h -~ for horses cantering, gaUoping and jumping. The DE requirements from the present study are in reasonable agreement with the NRC values (except for walking) when the differences in classifying activity are taken into account. The equations from the present study allow for much more flexibility in the calculation of requirements at various speeds. It should be re-emphasized, however, that these equations can only be used for the range of speeds measured in the present experiment. Furthermore, our measurements were made on a smooth, level surface. Terrain, type of surface, temperature, relative humidity and ability of rider would likely influence energy expenditures. The horses in this study were selected to provide a wide range of temperament and conformation because these factors are often considered to influence energy requirements during work. In this study, body weight was found to be the primary factor influencing energy expenditure at these slow speeds. When the energy expenditures were corrected for body weight, no differences were found among the horses. Perhaps temperament is more likely

830 to influence m a i n t e n a n c e requirements for exercise.

PAGAN AND HINTZ requirements than

L iteratu re Cited

Armsby, H. P. 1908. The Principles of Animal Nutrition. (3rd Ed.). John Wiley and Sons, Inc., New York, NY. Blaxter, K. L. 1962. The Energy Metabolism of Ruminants. Hutchinson and Co., Ltd., London. Brody, S. B. 1945. Bioenergetics and Growth. Reinhold Pub. Corp., New York, NY. Fedak, M. A., L. Rome and H. J. Seeherman. 1981. One-step N~-dilution technique for calibrating open-circuit VO2 measuring systems. J. Appl. Physiol. Respir. Environ. Exercise Physiol. 51:
772.

Hintz, H. F., S. J. Roberts, S. W. Sabin and H. F. Schryver. 1971. Energy requirements of light horses for various activities. J. Anita. Sci. 32:100. Hoffman, L., W. Klippel and R. Schiemann. 1967.

Untersuchungen uber den Energieumsarz beim Pferd unter besonderer Berucksichtigung der Horizontal hewegung. Arch. Tierernachr. 17: 441. Kane, E., J. P. Baker and L. S. Bull. 1979, Utilization of a corn oil supplemented diet by the pony. J. Anita, Sci. 48:1379. Kleiber, M. 1961. The Fire of Life. John Wiley and Sons, Inc., New York, NY. NRC. 1978. Nutrient Requirements of Horses. No. 6. (4th Ed.). National Academy of SciencesNational Research Council, Washington, DC. Pagan, J. D. and H. F. Hintz. 1986. Equine energctics. I. Relationship between body weight and energy requirements in horses. J. Anita. Sci. 63:815. Reid, J. T. and O. D. White. 1978. Comparative energetic efficiency of farm animals. Univ. Ark. Agr. Exp. Sta. Special Rep. 72. Taylor, C. R., N. C. Heglund, T. A. McMahon and T. R. Looney. 1980. Energetic cost of generating muscular force during running. J. Exp. Biol. 86:9.

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