Estimation of energy expenditure from heart rate measurements in cattle maintained under different conditions A. Brosh, Y. Aharoni, A. A.

Degen, D. Wright and B. Young J ANIM SCI 1998, 76:3054-3064.

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Estimation of Energy Expenditure from Heart Rate Measurements in Cattle Maintained Under Different Conditions1
A. Brosh*,2, Y. Aharoni*, A. A. Degen, D. Wright, and B. Young
*Department of Beef Cattle, Agricultural Research Organization, Institute of Animal Science, Newe Yaar Research Center, P. O. Box 1021, Ramat Yishay 30095, Israel; Desert Animal Adaptations and Husbandry, Jacob Blaustein Institute for Desert Research, Ben-Gurion University of the Negev, Beer Sheva 84105, Israel; Department of Companion Animal Medicine and Surgery, University of Queensland, St. Lucia 4067, Australia; and Department of Animal Production, University of Queensland, Gatton College 4345, Australia

ABSTRACT: We examined whether heart rate ( H R ) could be used to estimate energy expenditure ( E E ) in cattle. Six Hereford heifers (345 10.8 kg BW) 12 mo of age were implanted with HR radio transmitters and maintained in individual pens under the following treatments: 1 ) shade or sun exposure, 2 ) high- or low-energy diet, and 3 ) feeding in morning or afternoon. The HR of animals was measured every .5 h during 3 mo; measurements of oxygen consumption and HR were made simultaneously in the morning and in the afternoon while animals were resting and exercising. Average daily HR (52 4 beats/min) and average daily EE (380 9 kJ/kg.75) in animals on the low-energy diet were less than values in animals on the high-energy diet (94 4 beats/min and 653 9 kJ/ kg.75, respectively). For each animal and within each diet, linear regressions best described the relationship between HR and EE in resting animals, whereas

quadratic regressions best described this relationship for exercising animals. The quadratic equation for the exercising animals could also be used for resting animals. In addition, a constant value of EE per heart beat (EE pulse) for each individual resting animal was found and gave accurate estimations. This method was convenient because 1 ) no exercise equipment was needed to generate the regression equations and 2 ) EE pulse was less affected by diet than was EE estimated by regression equations. We conclude that HR, a relatively easy measurement, can be useful and accurate in estimating EE. To increase the accuracy of the estimation of EE by HR, the relationship of HR to EE should be established for each animal. In addition, the nutritional regimen for the animal in which EE is estimated should be used for the animal in establishing the relationship.

Key Words: Cattle, Feeds, Heart Rate, Oxygen Consumption, Energy Expenditure, Solar Radiation
1998 American Society of Animal Science. All rights reserved.

J. Anim. Sci. 1998. 76:30543064

Introduction
Energy expenditure ( EE) of farm animals has been determined mostly under controlled, confined conditions. These conditions do not necessarily reflect those of free-ranging animals, or of commercial cattle in feedlots. Environmental conditions, feeding level, time

1Contribution from the Agric. Res. Organization, Volcani Center, Bet Dagan, Israel, No. 2328-E, 1997 series. The authors wish to thank S. Fennell, G. Beneke, B. Hall, A. Goodwin, K. Rowan, J. McCosker, M. Josey, F. Gorbacz, R. Englebright, I. Williams, and T. Schoorl for their contributions. 2To whom correspondence should be addressed: phone: 972-4-9539523; fax: 04-9836936; E-mail: beefny@netvision.net.il. Received January 20, 1998. Accepted June 3, 1998.

spent eating and digesting, tissue and pelage conductance, production level, and season of the year may affect EE of animals (NRC, 1981). Double-labeled water ( DLW) can be used to determine CO2 production, a measure of metabolic rate of free-living animals, but DLW is expensive and the method contains errors of 8 to 11% (Nagy, 1989). Acute tracheal intubation for measurement of O2 uptake (Young and Webster, 1963) and infusion of [14C]bicarbonate (Corbett et al., 1971; Young and Corbett, 1972) have restricted field application and are expensive to use with large animals. In addition, the latter method interferes in the normal behavior of the animal. Estimation of EE of free-ranging large animals based on heart rate ( HR) has been examined by several workers (Webster, 1967; Yamamoto et al.,

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a

Table 1. The arrangement of the treatments during the experiment

Animal Days 14b 511 1218 1925b 2531 3238 3952b 5359 6066 6773b 7479 8085
aThe

1 L E A L E A L E M L P M L P M L P A H E M H E M H E A H P A H P M H P A

2 L P A L P A L P M L E M L E M L E A H P M H P M H P A H E A H E M H E A

3 H E A H E A H E M H P M H P M H P A L E M L E M L E A L P A L P M L P A

4 H P A H P A H P M H E M H E M H E A L P M L P M L P A L E A L E M L E A

5 H E A H E A H E M H P M H P M H P A L E M L E M L E A L P A L P M L P A

6 H P A H P A H P M H E M H E M H E A L P M L P M L P A L E A L E M L E A

Change of

Time fed Shade Time fed Diets Time fed Shade Time fed

letters represent the treatment conditions according to the following order. 1st: diet of low- ( L ) and high- ( H ) energy concentration, 2nd: exposure to sun ( E ) or protected by shade ( P ) , and 3rd: feeding in the morning ( M ) or afternoon ( A ) . bAcclimatization period.

1979; Richards and Lawrence, 1984; Renecker and Hudson, 1985; Yamamoto, 1989). This method could be most useful because recent developments in microelectronics allow the use of small HR radio transmitters to measure HR of animals in their natural habitat. The objective of this study was to determine whether HR in cattle can be used to estimate EE under different nutritional and environmental conditions.

Materials and Methods

Six trained 12-mo-old Hereford heifers (345 10.8 kg BW) were implanted with heart-rate radio transmitters (Telonics, Mesa, AZ) approximately 1 mo before commencement of measurements. During the 85-d study, HR for each animal was measured for 5 min every .5 h throughout every day. The six heifers were studied in a 2 2 2 arrangement (high- and low-energy feed; exposure to and protection from solar radiation; fed in morning and in afternoon) and underwent all treatments as outlined in Table 1. Measurements were made during the summer (JanuaryMarch) of 1993 in southeast Queensland, Australia, a subtropical region with summer temperatures commonly over 30C. Animals were kept individually in open feedlot pens, each 40 m2. Shade was

provided in about half the area of each of the pens: galvanized iron sheets, at a height of 2.2 m, covering 11.5 m2, and 70% shade cloth, at a height of 4 m, covering 12 m2. The animals were fed either a high-energy diet ( H) of 80% concentrate and 20% sorghum hay, or a lowenergy diet ( L) of only sorghum hay. The ME of the concentrate was calculated from its composition and known energy equivalents. The ME of the sorghum hay offered and left over was determined by nylon bag measurements of organic matter digestibility (Setala, 1983) and assuming a value of 15.06 MJ/kg digestible organic matter (Minson, 1982). The approximate analysis (AOAC, 1980) of the feed eaten and its calculated ME, taken as offered minus left over, are presented in Table 2. The quantity of feed given was regulated to reduce the refusals to less than 5% of that offered, and it was given either in the morning (0800 to 0830) or afternoon (1630 to 1700). Refusals were collected and weighed once weekly. Oxygen uptake was measured by the use of a face mask open-circuit respiratory system (Taylor et al., 1982). The accuracy was checked gravimetrically by injecting nitrogen into the mask (McLean and Tobin, 1990). The EE was calculated assuming 20.47 kJ/L O2 (Nicol and Young, 1990).

Table 2. Approximate analysis (% dry matter) and ME of the low- (L) and high- (H) energy diets consumed by the heifers
Ether extract 1.31 2.23 ME, MJ/kg 7.21 10.63

Diet L H
aCrude

OM 89.77 93.56 fiber.

CP 7.68 16.92

CFa 52.93 18.12

NDF 68.53 29.53

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Measurements of HR and O2 uptake were made at the same time for each animal on each treatment during two consecutive days when penned. The animals stood most of the time and, occasionally, lay down. Measurements were made over 15 to 20 min each time: between 0700 and 0830 before morning feeding and between 1400 and 1530 in the afternoon. Data were averaged every 5 s, recorded on a data logger (Mini-Logger, Mini-Mitter Co., Sunriver, OR), and transferred to a laptop computer for processing. For analysis, the data were pooled over 30-s intervals. For each such simultaneous measurement of HR and O2 uptake, the O2 pulse was calculated as the O2 uptake per heartbeat. There were four measurements per animal per treatment and 32 measurements per animal in total, composed of two measurements for each time (morning or afternoon) for each treatment. For analysis of the stability of O2 pulse, the difference between these two measurements was tested. Oxygen uptake and HR during exercise were measured simultaneously on four occasions for each animal on each diet. Measurements were made over 20 min on 2 d for each animal, each day, once in the morning and once in the afternoon. A circular walker was used to exercise the animals and the rate of walking was up to 6 km/h. The EE pulse was calculated as the energy expenditure per heart beat. Daily animal EE was calculated by multiplication of the total daily heart beats by the average EE pulse. Body weights of the heifers were determined every 2 wk and at the beginning and end of each treatment. Live weight ( LW) and daily LW gain ( LWG) were calculated from the regression slope of all the measurements of LW on each diet. Meteorological data (ambient temperature [Ta], black globe temperature [BG], and relative humidity [RH] ) were collected between 0700 and 0830 and between 1400 and 1530, and black globe humidity indices ( BGHI) (Buffington et al., 1981) were calculated.

effect of HR within diet. In the exercising model, a fixed effect was assigned to diet, and either a linear or a quadratic effect within diet. The equations of the model were as follows: Y = A + R + TF + TM + D/HR + e [1] Y = A + R + TF + TM + D/(HR + HR2) + e [2]

The equations of the models for exercising were as follows: Y = D/HR + e Y = D/(HR + HR2) + e [3] [4]

Statistical Analysis
Regression analysis was used to test whether a relationship existed between O2 uptake and HR and between O2 pulse and HR. The regressions were done separately for the data taken at rest and at exercise. For exercising animals, averages of O2 uptake and O2 pulse for each 5-beats/min interval of HR were taken for each animal at each state. Regressions were also done for all the animals, either at rest or exercising, with a random animal effect, to obtain 1 ) intercepts and slopes for each animal in each state and 2 ) a common slope that characterized the state. In the resting animal model, fixed effects were assigned for radiation conditions, time of feeding, time of measurement, and diet, with either a linear or a quadratic

where Y = O2 uptake or O2 pulse, A = random animal effect, R = radiation (exposed or protected) effect, TF = time of feeding (morning or afternoon) effect, TM = time of measurement (morning or afternoon) effect, D = diet ( L or H ) effect, HR, HR2 = linear and quadratic effects of heart rate, and e = error. The regression equations for each animal, linear and quadratic, based on the exercising data, were used to examine whether its O2 uptake at rest could be estimated from its HR. The O2 pulse was calculated in the same way except that only a linear regression was used. These estimated values were compared with the observed values, which were recorded simultaneously with the HR, by means of a paired t-test. The effect of time elapsed between a last given meal and measurement on the observed values of HR, O2 uptake, and O2 pulse during rest was estimated by linear regression of these variables on the elapsed time (hours). In this regression, a random effect was assigned to the animal, and fixed effects to conditions of radiation, time of feeding, time of measurement, and diet. The contribution of the variance between the pair of measurements for each situation to the residual variance was tested by means of the residual error when all the fixed effects and the linear effect of the elapsed time were accounted for by the regression. The absolute value of the difference between the residuals of the two replicates for each state for each animal ( n = 96) was calculated, and the mean value of this difference was averaged for each variable and compared with the mean value of this variable. All analyses were made with Genstat 5 Release 3.2 (Lawes Agricultural Trust, 1995).

Results and Discussion

Morning and afternoon air temperatures were 23.5 .8 and 30.2 .7C, whereas RH were 65 2 and 45 3%, respectively. Black globe temperatures were 25.2

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Figure 1. Average daily heart rates (SE) of heifers on high (n = 4; D) and low (n = 2; o) energy diets. Feed was given at 0830. .8 and 30.2 .7C under the shade and 37.6 1.3 and 45.0 1.7C in the sun in the morning and afternoon, respectively. Black globe humidity indices were 72.6 1.1 and 80.1 .7 in the shade and 84.9

1.3 and 92.1 1.6 in the sun, in the morning and in the afternoon, respectively. The HR was higher in heifers fed the H diet than the L diet and increased in all heifers during and after feeding (Figure 1). Previous studies that measured HR to estimate EE used linear or logarithmic regression equations to relate O2 uptake and HR (Webster, 1967; Yamamoto et al., 1979; Richards and Lawrence, 1984; Renecker and Hudson, 1985; Purwanto et al., 1990). In this study, within each diet, linear regression equations best described resting animals, whereas quadratic regression equations best described exercising animals (Tables 3 and 4). Generally, the slopes of the equations for exercising animals were steeper than those of resting ones (Figure 2). These data indicate that the dependence of O2 uptake on HR during rest is linear, and, hence, the O2 uptake per heartbeat (the O2 pulse) does not depend on HR under these conditions (Figure 3). In contrast to the resting conditions, a physical strain such as that imposed during exercise induced a O2 uptake increase that was not linearly related to the increase in HR. However, if the O2 uptake could be accurately described by a quadratic regression on HR, then the O2 pulse should be satisfactorily described by a linear regression, because the O2 pulse is given by O2 uptake divided by HR. The finding that the O2 pulse regression on HR was not linear suggests, therefore,

Table 3. Linear regression of O2 uptake (mL O2/(kg.75h)) and of O2 pulse (mL O2/ (kg.75beat) on heart rate (HR), with fixed effects of radiation (exposed or protected), time of feeding (morning or afternoon), time of measurements (morning or afternoon), and diet (L or H) at rest (df = 82) and at exercise (df = 102)
Rest Effect O2 uptake Radiation Time of feeding Time of measurement Diet HR slope on L dietb Difference H dietc r2 of regression O2 pulse Radiation Time of feeding Time of measurement Diet HR slope on L dietb Difference H dietc r2 of regression
aSignificance: bValue/beat. cDifference of P < .10.

Exercise Pa NS

Value 14.7 39.2 65.2 146 18.1 3.7 .933 3.77 8.14 17.98 2.4 .34 .37 .314

SE 20.0 19.8 21.0 172 2.5 2.8

Value 145 23.2 3.4 .742 14.3 1.24 .26 .202

SE

Pa

** NS *** NS

251 1.7 2.5

NS *** NS

4.36 4.33 4.58 37.5 .55 .61

NS

*** NS NS NS

39.8 .27 .39

NS *** NS

NS, not significant. slopes between diets L and H ( L H).

**P < .01. ***P < .001.

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Table 4. Quadratic regression of O2 uptake (mL O2(kg.75h) and of O2 pulse (mL O2/(kg.75beat) on heart rate (HR), with fixed effects of radiation (exposed or protected), time of feeding (morning or afternoon), time of measurements (morning or afternoon), and diets (L or H) at rest (df = 82) and at exercise (df = 102)
Rest Effect O2 uptake Radiation Time of feeding Time of measurement Diet HR slope on L dietb HR2 slope on L dietc Difference HR H diet2 Difference HR2 H diet2 r2 of regression O2 pulse Radiation Time of feeding Time of measurement Diet HR slope on L diet HR2 slope on L diet Difference HR H dietd Difference HR2 H dietd r2 of regression
aSignificance: bValue/beat. cValue/beat2. dDifference of

Exercise Pa NS * ** NS NS NS NS NS Value SE Pa

Value

SE

12.3 20.6 44.0 20.6 59.6 21.7 286 1,207 21.2 34.6 .387 .339 24.0 39.0 .33 .35 .932 3.35 .922 16.66 69 9.01 .092 5.57 .075 .319 4.49 4.48 4.72 262 7.51 .074 8.48 .077

2,880 810 43.5 7.8 .106 .040 44.2 14.6 .178 .064 .805 381 130 5.30 1.25 .021 .0065 5.12 2.34 .018 .010 .382

*** *** ** ** **

NS * *** NS NS NS NS NS

** ** ** *

NS, not significant; P < .10, *P < .05, **P < .01, ***P < .001. slopes between diets L and H ( L H).

that the regression of O2 uptake on HR during exercise could not be described satisfactorily, even with a quadratic polynomial. During exercise, three parameters that contribute to the O2 uptake are altered: 1 ) HR; 2 ) A-V difference, the difference between arterial and venous O2 in blood; and 3 ) stroke volume (Eckert et al., 1988; Jones et al., 1989). The negative slope on HR2 could have resulted from a reduction in the rate of increase of the A-V difference (Jones et al., 1989) and from a reduction of the stroke volume as the O2 uptake and HR increased as a result of exercise. This is probably why, when a polynomial of the third order was used to describe the relations of O2 uptake and O2 pulse on HR, neither of the regressions was improved, compared with the second-order polynomial (results of the third-order polynomial are not presented). During rest, the effect of time of feeding on the O2 uptake and the O2 pulse tended to be significant, and the effect of time of measurement on these parameters was significant. These parameters were lower in the afternoon than the morning feeding and in the afternoon than the morning measurement. No effect of radiation was detected on either O2 uptake or O2 pulse on HR. Individual regression equations for each animal on each diet (Figure 2 ) generated when exercising were

used to estimate EE at rest. The difference between estimated and measured values was significant only for the L diet, and only when the linear regression equations were used (Table 5). However, the standard errors of the mean were much lower when O2 pulse was used than O2 uptake for the linear and quadratic regressions. As can be seen in Figure 2, regression equations differed among animals and were affected by dietary ME. Therefore, to estimate EE from HR, a regression equation should be determined for each animal when it consumes a diet similar to the one in which the animals EE is to be estimated. In contrast to this, despite the significant effect of dietary ME on O2 pulse, the range of variations among animals and diets is minor (Figure 3), so O2 pulse (or EE pulse) to estimate EE by HR is less affected by diet than was EE estimated by regression equations. The HR increased after the meal and decreased thereafter (Figure 1). The correlation of O2 uptake and O2 pulse with HR (Tables 3 and 4 ) was affected by the time of feeding, morning or afternoon, and by the time of measurement, and it was postulated that these effects could also be attributed to the time that elapsed between the meal and the measurement. Therefore, we tested the effect of elapsed time on HR, O2 uptake, and O2 pulse (Table 6). We used all 192 individual measurements of HR and O2 uptake,

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comprising two measurements for each combination of diet, time of feeding, time of measurement, and radiation conditions for each heifer. Random effect of the animal and fixed effects of all the above-mentioned conditions were accounted for by the regression equations obtained in the first step, in addition to the linear effect, the elapsed time. In each of the regressions of HR, O2 uptake, and O2 pulse, all the fixed effects that were not significant in the first step

were excluded from the regression in the next step. Thus, only diet and time of measurement effects were included in the regression of HR, only diet effect in the regression of O2 uptake, and all the fixed effects but radiation in the regression of O2 pulse on the time elapsed since the last meal before the measurement. The HR and O2 uptake decreased with increasing time, at rates of .72 and .92% of the mean per hour, respectively, which left the O2 pulse relatively cons-

Figure 2. Oxygen uptake (mL/(kg.75h) SE on the ordinate and heart rate (beats/min) on the abscissa of six heifers on low- (L) and high- (H) ME diets. Measurements at rest in the pens (R, empty symbols) under all treatments and at exercise using a walker (W, filled symbols). Linear regression, broken line for resting animals and quadratic regression, full line when exercising. Note the different axes limits for each animal.
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Figure 3. Oxygen pulse (mL/(beatkg.75) SE and heart rate (beats/min) of six heifers on low- (L) and on high- (H) ME diets at rest in the pens, panels A and B, respectively, and at exercise using a walker on diet L and on diet H, panels C and D, respectively. Note the different axes limits for each panel.

tant with time. Even though the effect of diet was highly significant for all three parameters, the magnitude of this effect was only 10.5% of the mean for the O2 pulse, compared with 68.5 and 59.0% of the mean for HR and O2 uptake, respectively. The O2 pulse was relatively constant for each animal, compared with O2 uptake, for the range of conditions in the pens, provided that the animals were not under intensive physical strain. We suggest,

therefore, that O2 pulse is the most appropriate parameter to use in estimating EE from the HR of animals in rest conditions. In pens, the percentage change between daily minimum and maximum HR was 80 to 175% on the L diet and 70 to 149% on the H diet, with constant O2 pulse throughout this range (Table 3). Determinations of HR, O2 uptake, and O2 pulse were done twice for each animal on two separate days,

Table 5. The percentage difference between measured O2 uptake and O2 pulse of heifers at rest and those estimated from linear and quadratic regression equations at exercise. The regressions for exercising animals were calculated separately for each one (n = 6)
Low-energy diet Parameter O2 uptake O2 uptake O2 pulse
aNS,

High-energy diet Pa * NS NS % difference 1.0 .6 1.0 SE 6.2 7.0 4.0 P NS NS NS

Calculation Linear Quadratic Average

% difference 25.5 3.3 .6

SE 9.8 13.4 2.5

not significant; *P < .05.

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a

Table 6. The linear effect of time elapsed (h) from the previous feeding time to measurement, and fixed effects of treatments in the pens, on HR, O2 uptake, and O2 pulse (n = 192)
Fixed effects Dependent parameter HRc Slope % of mean SE Pd O2 uptakee Slope % of Mean SE Pd O2 pulsef Slope % of Mean SE Pd Time of measurement 4.41 6.3 1.29 *** g 17.81 7.1 3.67 *** Time of feeding 8.48 3.4 3.43 * Diet 48.2 68.5 1.16 *** 613.2 59.0 .34 *** 26.39 10.5 3.26 ***

Time from meal, hb .508 .72 .091 *** 9.54 .92 .024 *** .183 .07 .269 NS

of fixed and linear effects. Only significant fixed effects were included. cBeats/min. dNS, not significant; *P < .05, ***P < .001. emL O /kg.75. 2 fmL O /(kg .75beat). 2 gDash indicates that the fixed effect did not account for a significant amount of variance in an initial model and thus was dropped from the analysis used to generate the data in this table.

aVaried between 3.4 and 30.2 h. bUnits/h. Evaluated in a common analysis

under each set of conditions in the pens. The mean absolute values of the differences between the residuals of the two replicates for each state for each animal were .52 beat/min, .78 mL O2/(kg .75h), and .90 mL O2/ (kg .75beat) for HR, O2 uptake, and O2 pulse, respectively. These differences were equal to only .74, 3.90, and .36% of the mean values of HR, O2 uptake, and O2 pulse, respectively. It is suggested, therefore, that a single determination is sufficient. The O2 pulse of a number of cattle breeds under different conditions at rest is presented in Table 7.

The variance in each breed was small. It is suggested, therefore, that despite the individual character of the O2 pulse and its dependence on diet and environmental conditions, HR can provide an estimation of their EE. Moreover, provided that the cattle in pens and pasture are not exposed to exercise, day-to-day changes in HR provide a reliable estimation of changes in EE. It can be concluded that the EE of the heifers can be estimated from HR measurements, using quadratic equations that relate O2 uptake to HR or by using a

Table 7. The O2 pulse (mL/(kg.75beat) and CV (SE/mean100) of cattle at rest under different conditions
Animals and conditions Heifers, low-energy diet Heifers, high-energy diet Standing Hinterwaeelder oxen Standing Zebu oxen Swedish Red and White steers Simmental oxen Boran cows Heifers of various breeds Black-Pied dairy cattle bulls (16 a) Mean SE of the 9 references
aAverage of six bMeasurements

n 6 6 7 5 3 8 6

BW, kg 345 337 494 516 449 562 475 123177 280350 315 3.2

O2 pulse 265 238 347 300 377 338 429 252 286 315 3.2

CV, % 2.60 2.50 2.8 2.7 6.80 2.4 1.0

References Present study Present study Rometsch et al., 1997 Rometsch et al., 1997 Jones et al., 1989 Clar, 1991 Zerbini et al., 1992 Liang et al., 1997 Derno et al., 1997 CV of the 9 references

measurements of 16 animals. of animals at rest.

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constant value of O2 uptake per heartbeat ( O 2 pulse), which is less affected by the nutritional conditions. The relation of O2 uptake to HR is dependent on the causes for changes of HR rather than on HR itself. Use of HR to estimate energy retention and energy balance was evaluated. The ME intake ( MEI) , EE, and retained energy ( RE) of the heifers during the entire study are presented in Figure 4A, B, and C, respectively. The LW, gain, and the energy balance

parameters on each diet during the entire experiment are presented in Table 8. The MEI of the heifers was directly related to the ME concentration of the diet, and the EE and the RE followed the changes in the MEI. Before the start of the experiment, the heifers were fed the L diet; during the first days of the experiment, the intake of the heifers on the H diet greatly increased, resulting in similar increases in the EE and RE. Because the amount of feed given during each period was constant and the weight gain of the animals on the H diet was positive, the intake per metabolic weight decreased during each period. When the diets were switched, the MEI followed the changes in the diet energy density and, consequently, so did the EE and the RE. Calculation of EE by multiplication of the HR by the EE pulse, calculated from the O2 pulse, was less accurate immediately after the change in feed quality. For calculation of EE during that period, the EE pulse of the previous diet was used for the calculation. Because the difference between the EE pulses on the two diets was only 10.5% and the time taken for the heifers to reach stability in intake and EE after the switch of diets was short, the resulting inaccuracy in the determination of EE was small. Predicted EE according to the National Research Council (NRC, 1984) is 432 47 and 750 46 kJ/ (dkg .75) on the L and H diets, respectively (the individual MEI and the LWG of the heifers in the present study were used for the calculation). The NRC predictions were not significantly different from our estimation according to the HR and the EE pulse (Table 8). The calculated RE per LWG for the H diet (28.4 1.5 MJ/kg LWG) tended to be larger than the value obtained by use of the observed LW and LWG in the NRC (1984) equation (23.6 1.0 MJ/kg LWG). The LWG on the L diet was negligible, as was the RE, and, therefore, the calculation of RE per LWG from these values were not done. We conclude that HR, a relatively easy measurement, can be useful and accurate in estimating EE. To increase the accuracy of the estimation of EE by HR, the relationship of HR to EE should be established for each individual animal. In addition, the nutritional regimen used in establishing the relationship should be similar to that for the animal in which EE will be estimated. Measurements obtained using this method, combined with MEI, would allow estimations of body energy changes in animals losing or gaining body weight.

Figure 4. Metabolizable energy intake (A), energy expenditure (B), and retained energy (C) of six heifers. Four were fed a high-energy diet (10.62 MJ/kg DM of ME) during the first 45 d (thin lines) and a low-energy diet (7.21 MJ/kg DM of ME) subsequently. The other two (thick lines) were fed the low- and then high-energy diets.

Implications
Heart rate is relatively easy to measure, and this measurement seems to be a practicable method for estimating accurately the short-term and long-term

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Table 8. Body weight, live weight (LW) gain (LWG), energy balance parameters, and energy content in LW gain of the heifers (mean of six heifers SE) that were fed high- (H) and low- (L) energy diets. The animals were kept in individual pens during the summer
Diet H SE CV L SE CV Alli SE CV BWa 337 15 4.6 345 10 3.0 337 13 6.2 Gainb 1,460 120 8.0 157 97 61.9 810 50 6.3 LWGc 18.45 .88 .8 2.079 1.233 59.3 10.37 .64 6.2 MEId 1,190 25 2.1 468 26 5.6 843 13 1.6 EEe 670 10 1.4 394 19 4.8 541 8 1.4 REf 520 22 4.14 74 11 14.6 301 9 3.0 NEgg 28.4 1.5 5.20
h

29.6 2.1 7.2

akg. bg/d. cg/(dkg .75) . dMetabolizable energy intake, kJ/(dkg .75) . eEnergy expenditure, kJ/(dkg .75) . fRetained energy, kJ/(dkg .75) . gNE , energy content in live weight gain, kJ/g. g hNot calculated because of the negligible LWG. iPeriod of adaptation to the diets and period of diet

changeover in the middle of the experiment (about

2 wk for each period) are included.

variations of energy expenditure in free-range cattle. When metabolizable energy intake is also measured, energy retention in animals and the energy content of the gain or loss of body weight can be estimated. This would allow the determinations of energy requirements of animals and would be more accurate than simply using changes in body mass as a criterion. Under certain circumstances, it would also allow the calculation of the efficiency of utilization of energy intake for maintenance and for growth.

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