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T H E R M O R E G U L A T I O N AND TOTAL BODY INSULATION IN THE NEONATAL FOAL
J. C. OUSEY, 1 A. J. MCARTHUR, 2 P. R. MURGATROYD, 3 J. H. STEWART I a n d P. D. ROSSDALEl t Beaufort Cottage Stables, High Street, Newmarket, Suffolk CB8 8JS, 2Department of Physiology and Environmental Science, University of Nottingham School of Agriculture, Sutton Bonington, Loughborough LEI2 5RD and 3Dunn Clinical Nutrition Centre, 100 Tennis Court Road, Cambridge CB2 IQL, England
(Received 26 January 1991; accepted in revised form 15 October 1991)
Abstract--1. The metabolic rates of pony foals (Equus caballus), age 2-4 and 7-9 days, were measured at different air temperatures between 2 and 40°C in order to identify their lower critical temperature (Ttc) and the environmental demand for heat when the air temperature (Ta) was below this value. In addition, respiratory quotient, rectal temperature and heart rate were measured. 2. The mean values of Tic were about 22 and 19°C for the 2-4 and 7 9 day old foals, respectively. The wide variation (about 10°C) in Tic within each group was attributed to differences in thermal insulation. Below thermoneutrality, metabolic rate, heart rate and respiratory quotient increased with decreasing Ta,
but rectal temperature remained fairly constant at about 38.4°C. 3. The foals thermoregulated effectively under the environmental conditions studied.
Key Word Index: Neonatal foal; thermoregulation; thermal insulation; lower critical temperature; metabolic rate
The importance of maintaining neonates at environmental temperatures within their thermoneutral zone has been demonstrated for many species. For example, Silverman (1959) reported that the mortality rate of premature babies decreased significantly when they were maintained in a warm (31.8°C) environment; and Alexander (1962) found that mortality in lambs due to hypothermia was more likely to occur when they were exposed to wet and windy conditions than when kept dry in still air. The literature contains much information about the thermal requirements of infants and the neonates of domestic livestock (e.g. Sinclair, 1978; Poczopko, 1981). The lower critical temperature T1¢(°C) for these species, defined as the air temperature below which a homeotherm must increase its metabolic rate to meet the environmental demand for heat, ranges from about 36°C for infants (Hill and Rahimtulla, 1965) to about 9°C for calves (Webster, 1974). In contrast, there is a paucity of data describing thermoregulation in foals. In the U.K. most Thoroughbred foals are born at night between January and April, when daily mean temperatures outdoors are usually below 10°C. Although most foaling boxes have radiant heaters, the thermal environment into which foals are delivered is usually poorly controlled. The optimum range of environmental temperatures for the young foal, and its ability to thermoregulate at temperatures below the lower critical value, are not known. The aim of this study was to measure the metabolic rates of young foals over a wide range of air temperatures Ti(°C), to identify their thermoneutral zone (zone of minimal metabolism) and lower critical temperature, and to determine their capacity to increase metabolic
rate in response to air temperatures below the lower critical value.
For homeotherms, the relationships between the rates of heat production, evaporative and non-evaporative heat loss, and environmental temperature can be described by the "metabolic diagram" (Mount, 1979). The lower critical temperature, which defines the lower end of an animal's thermoneutral zone, depends on its thermoneutral rate of heat production Mt,(W m -2) and on its components of thermal insulation. This temperature can be determined from Tie = Tb-- [(rs + re + re)(M,n -- 2E,m)/pCp]
-}-[rc+re]AEsm/pC p (1)
where body-core temperature (°C), thermal resistance of body tissue (s m - l ) , thermal resistance of coat (s m - 1), thermal resistance of environment (s m - l), volumetric specific heat of air (1290Jm -3 K-l), 2 = latent heat of vaporization of water (J g- 1), Esm = minimum rate of water loss per unit surface area by evaporation from skin ( g m -2 s - l ) , E~ = minimum rate of water loss per unit surface area by evaporation from respiratory system
Tb = rs = rc = re = pep =
Equation (1) is similar to that given by Blaxter (1989) for the air temperature at which the heat emission from an animal with fully vasoconstricted skin, and with the skin and lungs losing minimal amounts of
The mean body mass (±SD). Steady state measurements of metabolic rate were made at air temperatures between 2 and 40°C. The steady flow rate of air through the mask was between 40 and 80 1. Values of r~ are usually considered to be at a maximum (vasoconstriction) when air temperature is at or below the lower critical value. respectively. volume) over periods of about 4 rain. ) (4) water vapour. The wind speed varied with position in the chamber. min ~ . acts between the body-core and skin surface.. 1989).g. Similarly.9) kg.M x b ] / p c p .1 mob '67. equation (2) can be rearranged to give riot = p c p ( T b . respectively.: + r. M e t a b o l i c chamber The foals were studied individually in an insulated metabolic chamber which was 3. to a reasonable approximation. the rate of heat production per unit surface area M (W m 2) required for homeothermy can be calculated from M = pcp[Tb -.1) and 1.7) and 35. When the rate of Metabolic rate was determined by open-circuit indirect calorimetry. based on the formula of Meeh (1879).5 m s D e t e r m i n a t i o n o f metabolic rate (3) The value of r~o~ for an animal can be determined from the slope of a straight line fitted to measurements of its rate of heat production at different environmental temperatures below thermoneutrality.]/[r~+ r. but close to the foal it was usually less than 0. healthy pony foals were studied at ages 2 4 and 7-9 days. = Tb. The rates of oxygen consumption and carbon dioxide production were derived according to the assumption that nitrogen and other inert gases are conserved in the ventilating air (Dauncey et al. The surface area (m 2) of each foal was estimated as 0. This quantity depends on the thickness of the skin. Twenty-four spontaneously delivered.e."F~.. A thermal resistance of 100 s m ~is equivalent to an insulation of 0.. according to the size of the foal. The slope of the line relating M to 7~ below thermoneutrality is determined by the total thermal resistance of the body. + r~.m) and respiratory system (2Erm) are minimal. McArthur (1991a) showed that. using a ventilated mask placed over the foal's muzzle.C. The first.1 (_+0. In these conditions. The value of r~ depends mainly on the size of the animal and on the wind speed to which it is exposed (Monteith and Unsworth. the resistance rto~can be evaluated from equation (2) provided that figures can be obtained for the two terms on the right-hand side involving the latent heat losses ). the environmental resistance r~.Esm and 2E~m. 1990). Samples of the inspired and ventilating air were collected into Douglas bags (101. The third.Th -.078 K m 2W 1 (Monteith and Unsworth. is equal to its heat production in the thermoneutral zone (resting value). equation (1) can be approximated by fl~-. The ventilation rate of the mask was corrected to give the dry flow rate at STP. in accordance with equation (2).6 × 2. The final values of oxygen consumption and carbon dioxide production are estimated to be within + 2% of the true values. the coat resistance r~.+4. 1991a). on the wind speed to which the animal is exposed and on the gradients of temperature and humidity within the coat (Cena and Clark. Below thermoneutrality. the flux density of heat (total) from the body exceeds M.6 × 3. convection and thermal radiation exchange). OUSEVet al. mb (kg). of each group was 31. walls.m/(r~+r~+r~)]+)~Erm.r~o~(Mtn . These samples were analysed within 2 h of collection for oxygen and carbon dioxide content by means of Servomex (Model 137) and PK Morgan (model 801) gas analysers. The equation can be derived by manipulating the heat flow equations. floor). with both usually below 1 0 W i n 2 and nearly all of the heat loss from the body is by non-evaporative routes (i.~E. and this line usually extrapolates to intercept the temperature axis at a value above body-core temperature (McArthur. However. . the tissue resistance r~. Here. 1978).. The mean ( + S D ) values of surface area for the two groups were 1. and assumes that conduction of heat to the ground and convective heat losses from the respiratory system are small enough to ignore.4 m in size and in which the air temperature was controlled thermostatically to within + I~C. where Mro is the metabolic rate predicted by equation (2) when T. unless air temperatures are less than 0 C when cold-induced vasodilation may occur.1) m 2. Air temperature was monitored by two mercury-in-glass thermometers (accuracy _+0.MT~.1 C) situated close to the foal.2 J. The second. and was measured by a Rotameter flow meter (type 200/24). the flux densities of latent heat from the skin surface (. 1978). We can define the total thermal resistance of the body r~ot (sm -1 ) by rto~ = r~ + r. and thermal resistance is in units of s mrather than the traditional unit of insulation (K m2W t) often used by animal scientists. respectively. the heat flux densities are expressed in watts per unit area of body surface. on the amount of subcutaneous fat and on peripheral blood flow (Blaxter.0 (-+0.1 (+5.] +[(r~+r~))~E. depends on the depth and structure of the hair layer. governs the non-evaporative heat losses from the coat surface by convection to the surrounding air and by thermal radiation exchange with surrounding surfaces (e. For most species. Respiratory quotient (RQ) and metabolic rate (Win -2) were calculated from these rates of gaseous exchange. The total body resistance comprises three insulating layers acting in series. 1990).n.3 (.T a ) / ( M . MATERIALS AND METHODS (5) Animals (2) Equations (I) and (2) show that lower critical temperature and metabolism in the cold depend largely on the components of thermal resistance between an animal's body-core and its environment. heat production M is measured at a single air temperature below Tic. this notional metabolic rate is about 10 W m -2.
foals of a given age were studied at two air temperatures. some of the 24 day old foals shivered at air temperatures up to 26”C. as indicated by equation (1). The foal lay with its legs partially extended and this posture was maintained while respiratory gases were collected. the maximum 171 W m-* for a 2-4 day old foal and 153 W m-* by a 7-9 day old foal at air temperatures of 6. RESULTS Figures l(a) and (b) show metabolic rate plotted against air temperature for foals age 2-4 and 7-9 days..7”C. The depth of the pelage was measured on a small number of foals (n = 9). respectively. The total body resistance r. Figures 3(a) and (b) show that the RQ for all the foals was between about 0. observed in both age groups.. The lines drawn on Fig.. for each age group were 333 (+ 66) and 343 ( f 59) s m-‘.. Figures 4(a) and (b) show that the heart rate was positively correlated with metabolic rate (P -C0. the number of heart beats was counted over a period of 15-30 s before and during the gas collection. 1 are discussed below. respectively). rectal temperature was measured with a small clinical mercury-in-glass thermometer. of each foal at air temperatures < 20°C was calculated from equation (4). The mean (+ SD) values of rto. Below about 20°C the metabolic rate of both groups increased progressively with decreasing air temperature and shivering was observed in all foals.62 and r =0. This procedure was repeated until the recalculated values of h4. 1 and an improved value of T. Pilo-erection enabled the foals to increase their coat depth in the cold. The large variation in metabolic rates between individual foals at a given temperature in the cold can be accounted for by differences in the total body resistance (Fig. using a graduated probe held normal to the skin surface. respectively.. Above 20°C the metabolic rate of each group was relatively constant.25-2 h by bottle or stomach tube. on the total body resistance of foals in each age group. is attributable to an increased non-evaporative heat loss from the body surface. consistent with the thermoneutral zone (zone of minimal metabolism) described by Mount (1979). 5) which are likely to be a consequence of differences in coat depth and/or in the amount of subcutaneous fat. it was placed in left lateral recumbency in the metabolism crate and restrained until it relaxed and (usually) fell asleep. Sweating was observed when foals from both age groups were exposed to air temperatures above 38°C.5 and 2.3) “C for the 2-4 and 7-9 day old foals. mean values of M.3-l . Statistical procedures in the neonatal foal 3 the range of air temperature studied. 2 20°C.7 and 0.53 for 24 and 7-9 day old foals. Mean values + 1 SD are given where appropriate. were estimated from the data (Fig. The increase in metabolic rate with decreasing air temperature below this zone. under constant supervision. Using this value of T.. Table 1 presents these recalculated . The foal was fed every 1. typically from about 0. Fig. Just before the next feed. However. respectively.001. 5 were used in a simple iterative procedure to estimate the dependence of q. At an air temperature of 5°C the metabolic rates were almost double the thermoneutral values. It was noted whether the foal was shivering or sweating by visual/tactile observations..66 for 2-4 and 7-9 day old foals. Figures 2(a) and (b) show that the rectal temperatures of both groups remained almost constant over The results presented in Fig. Furthermore. There were no consistent changes with age in the total body resistance values for individual foals. The heart rate of both groups of foals increased when air temperature decreased below 20°C. At the end of the gas collection.9. r = 0.3 (kO. r =0. respectively.4 (f0. These graphs indicate that the lower critical temperature for both groups of foals is about 20°C. Each foal was separated from its dam and placed in a metabolism crate within the metabolic chamber. Figures 5(a) and (b) shows these values of r.59 and 0. However.001. A value of T. were recalculated from the data in Fig. The highest metabolic rates were observed in the younger age recorded values being group. there were considerable differences in body resistance between individuals: the smallest values of rtoL were about 250 s m-’ and the largest values nearly 500 s mm‘. If the foal became excited during the collection of respiratory gases. respectively. These differences in thermal insulation will also affect the lower critical temperature of individual foals. At the start of this calculation.c. Heart rate was monitored by hand. using dam’s milk or milk replacer. was then determined from equation (5).4 cm when air temperature was reduced from a thermoneutral value to about 10°C. 1) obtained when T. and some 7-9 day old foals shivered at air temperatures up to 22°C. differing by no more than 5°C. the procedure was abandoned and repeated when the foal was calm. with mean (*SD) values of 38. the mean values of M1. DISCUSSION Linear regression analysis was performed and the correlation coefficient r was calculated as described by Armitage (1971). 1 demonstrate that the metabolic rates of recumbent pony foals were fairly constant and minimal at air temperatures between about 20 and 35”C.3) and 38. at about 70 and 85 W m-*.c for each group was then calculated by means of equation (5) using the mean total body resistance values of 333 s m-’ (foals 2-4 days) or 343 s m-l (foals 7-9 days). It stood on a rubber mesh mat on the base of the crate which was raised off the floor of the metabolic chamber. to ensure that the foal was in a “resting” state.. and Tk for each group were insignificantly different from the previous values. and kept in the chamber for at least 3 h to ensure thermal equilibrium was established. respectively). 5 reveals evidence of a significant decrease in the total body resistance of some foals with decreasing air temperature below thermoneutrality.Thermoregulation Experimental protocol Within the two age groups. The slight negative correlation with air temperature was statistically significant (P ~0. plotted against air temperature for the respective age groups. The values of rtat presented in Fig.
Table 1 presents these m a x i m u m and m i n i m u m values of T~c for each age group.o S' o• %0 o 435 s m1 ~ 6o 40 20 i i .. 15 I . . . . and 260 and 435 s m ~ for foals age 7-9 days). . v 140 120 ~ I rtot= (b) rtot = 260 s m -1 IO0 o o o o o O o ..1 0 60 4O 2O 0 0 rio t I Io o . . The m i n i m u m and m a x i m u m values of T. values for the two age groups. . 1. . i .1 and 18. 255 and 4 6 5 s m -1 for foals age 2-4 days. . ! . and the metabolic rates in the cold for foals of high and low total body resistance r. l .. .o. . Metabolic rate plotted against air temperature for pony foals. . 20 25 30 35 40 45 Air temperature (°C) 220 200 180 160 ~. . This calculation was done using the values of Mr.e. The solid lines on Figs l(a) and (b) show the mean values of metabolic rate for each group of .Q 8o oo '. the values of T~c being 22. . t 5 10 15 20 25 30 35 40 45 Air temperature (°C) Fig. The solid lines show the mean values of metabolic rate. The dashed lines indicate the mean and range of values for lower critical temperature.J. t . . . 10 i. ... .6°C.. . . given in Table 1 and the average of the three lowest and three highest values of total body resistance obtained for each age group at air temperatures below the respective mean values of TLc (i. .c for each group were also evaluated from equation (5). respectively. m . OUSEY et al. . 5 i .. 220 (a) 200 ~ rtotp=255 s m 1 180 'E v 160 140 - - o o o o 120 m 100 o o o °o o 0 m "d 80 o o o 465 s m. . age (a) 2 4 days and (b) 7-9 days. C.
. . . . . . ..  ~ .. .. The mean values of T~ for the 2--4 and 7-9 day old foals are above the mean daily air temperatures .=.5 o 37.. Table 1 also indicates that the mean value of T~¢for the older pony foals was about 4°C lower than that for the younger foals. . i . i 0 5 10 15 20 25 30 35 40 45 Air temperature (°C) Fig.. . . i . i . A gradual rise in resting metabolic rate (expressed per unit surface foals at and below therrnoneutrality. 2. . ... . .. . . . 38. . . 0 .0 .0 temperature (b) 39. . the value of Tic for a well insulated (r. .1) foal.   ~__a_~.K. . .. i . . and the maximum and minimum values as calculated above according to differences in insulation. . . . i 5 10 15 20 25 30 (°C) 35 40 45 Air 40. i . . . a consequence of the increase of about 12 W m-2 (16%) in resting metabolic rate with age.. .... ..t. .0 5 (a) 39.5 A U o 39.. i . i . . . ... The dashed vertical lines indicate the mean value of T~ for each group. . .. . mean value o 0  o n37.5 U o 39. . . . I . i . i .. .=_~_. . .0 --'a . and the metabolic rates in the cold for foals of high and low (constant) total body resistance. . i . I . i .0 P o o   o o 0  13 0  o 13  o Q.5 3 7 . usually encountered outdoors during the early part of the foaling season in the U.. mean value E o m ca G) 38. . .0  o O E e~ 38. . Table 1 indicates that there is a wide range (approximately 10°C) in values for T~ within each group of foals.... . .. . .. i . . . . = 500 s m ._____.5 a Q UD  Q QQ D ~ O O _. . . . However. . a ~ .. ...     D q. . . . 0 1 l ... .5 E 38.o. . Rectal temperature plotted against air temperature for pony foals. ...0 o o 37. age (a) 2--4 days and (b) 7-9 days.Thermoregulation in the neonatal foal 40. . . . . . ... i . .l ) foal could be as low as II°C compared with 26°C for a poorly insulated (rtot = 250 s m .. . . . .
. . .5 kg d -1 . . . . i . Mount. . . .C. . . .6 0. the results presented in Fig.85 & • • A• • & .75 A A A A ~ A 0. . i . . and is usually associated with increases in insulation (i. The values of T~ presented in Table 1 apply only to dry foals which are recumbent. 1986). . . .90 A '~ 0. . t . . . i . . .= & • • • 4= & A • • • • 0. resting and in an environment where the wind speed is low and there . . fat deposition).60 0 . .95 (b) 0. The mass of the foals studied here increased at a rate of about 0. .75 & & & & t~ 0. t 5 ~0 ~5 Air 20 25 30 35 40 45 temperature (°C) 0. i .65 0.95 J. in body size (reducing the surface area to mass ratio of the individual) and in plane of nutrition (Alexander. . . . . O 0. . Dauncey and Ingram.65 0. Respiratory quotient plotted against air temperature for pony foals. e 0 5 10 15 20 25 30 35 40 45 Air temperature (°C) Fig. 1983). = . . . e . i . age (a) 2-4 days and (b) 7-9 days.85 & & & A A A &A & A A A A &&A & & A & A A 0. (a) 0. coat depth. t . ~ . i .. in contrast to infants who lose mass during the first few days post partum. . . t . This increase in mass of the foals is consistent with an increase in mares' milk production during the first few weeks of lactation (Oftedal et al.e. OUSEYet al. area) with age has been reported for other species. 3. . . . . | . 1974.90 '• (~ A & & & 0. . .80 • & • • O 0. . did not contribute significantly to the decrease in lower critical temperature during the first week post partum. . .80 A =>. therefore.70 0. . .70 0. . i .60 i a i i i . 5 indicate that the total body insulation of the foals remained almost constant between days 2-4 and 7-9 post partum and. . However. . 1976. i e . . .
Ousey et al. even when the air temperature was 20°C.2) Fig. I 70 . healthy young foals spend much of their time outdoors. Heart rate plotted against the metabolic rate for pony foals. Rossdale (1968) reported that the rectal temperatures of Thoroughbred foals suffering from prematurity. I 90 = I 110 M e t a b o l i c r a t e (W m. Also. an increase which is likely to lower the value of T~¢. 100 90 ~ m 80 7o Z @ 60 s a I 130 i I 150 I f 170 I I 190 I 50 50 . the critical air temperature for foals outdoors may differ considerably from the values given in Table 1.Thermoregulation in the neonatal foal 130 7 (a) 120 110 @: 100 @ @ @ @ @ ~• • @ • 90 ~ @ @ • @ (IB @ 0 @ @ • • @ • • t~ -I- so • 70 @ • @ @ 60 50 50 I 70 i I 90 .) affect the thermal status of animals (McArthur. etc. solar radiation. For comparison. For example. (1991) observed that the metabolic rates of standing foals (dry) were about 50% above the rates when recumbent. standing or physically active. Values of Tic for these wet foals are likely to have been much higher than those reported here for dry foals. consequently. age (a) 2 4 days and (b) 7-9 days. Ousey et al. Furthermore. I 110 i I 130 i I 150 l I i 170 I 190 M e t a b o l i c rate (W m" 2) 130 @ 120 (b) 110 E 0. asphyxia and convulsions were different . (1991) reported that newborn foals which were wet with amniotic fluid and shivering during the first 2 h post partum had metabolic rates which were above 200 W m -2. the thermoregulatory responses of the neonate can be altered by its clinical status. Other climatological variables (wind. 1991b) and. 4. is no sunlight.
The solid lines connect thermal resistance values for the same individual. / r~  0 300 N D F- 250 200 I I i I I I i I I i I . The metabolic rates of the older foals were relatively constant up to about 40°C. Total body resistance rtot plotted against air temperature (~<20°C) for pony foals.2°C (Fig. the highest air temperature which could be attained in the chamber. I . Clearly. I i I I 2 4 6 8 10 12 14 16 18 20 Air t e m p e r a t u r e (°C) Fig..C.0°C suggests that the upper critical temperature had been reached for this foal.~ 300 I'" 250 200 I . The warm limit (upper critical temperature) of the zone of minimal metabolism could not be estimated from the measurements reported here. . Figure 5 indicates that the thermal insulation of individual foals can alter below thermoneutrality. In the younger group.. age (a) 2 4 days and (b) 7 9 days. l(a)). the corresponding increase in rectal temperature to 39. determined at different air temperatures.. I i I i I i I i I . health must also be considered when choosing the thermal environment for equine neonates. from those of healthy foals during the first 48 h post partum.8 500 J. "0 0 . I . one foal increased its metabolic rate by 20% above the mean thermoneutral value when the air temperature reached 38. (a) 450 0 400 t~ 350 L. I 2 4 6 8 10 12 14 16 18 20 Air t e m p e r a t u r e (°C) 500 (b) 450 / / ra U t" (8 400 350 I. 5. OusEY et al.
surface area.1 73 85 333 343 22. In contrast.0 30. The lower critical temperature (Tic). the foal can withstand a n air temperature as low as 2°C without a c o n c o m i t a n t fall in rectal temperature. .1 kJ. A coefficient o f 0. Blaxter et al. 1961. they have a wider zone of minimal metabolism a n d Tic can be as m u c h as 30°C below n o r m a l body-core t e m p e r a t u r e (e. precocious or i m m a t u r e neonates.5 F o r some foals..10 was used in the present analysis.Thermoregulation in the neonatal foal Table 1.1 18. Pilo-erection in response to cold was observed in foals o f b o t h age groups a n d it is likely t h a t the foals h a d vasoconstricted when the air temperature was close to Tic.93/ 224 Hey and O'Connell 0970) Naked Human 3-10 28 ~7. resting rate of oxygen consumption. foals were exposed to each air temperature for between 4 and 5 h.1 35. the results presented here suggest that shivering is a n i m p o r t a n t mechanism for Table 2. a decrease in rtot below t h e r m o n e u t r a l i t y may be attributable in p a r t to a decrease in the external insulation (re + re) caused by the rapid body m o v e m e n t s during shivering. The presence of b r o w n adipose tissue a n d N S T has not been observed in foals. such as the human. The significant increase in R Q with decreasing air temperature.5 h. resting metabolic rate and total body resistance of newborn animals Oxygen* consumption Metabolic* Thermalt Age Tz (ml kg. Altricial newborn. The coefficient 0. tValues based on equation (4) in text (the value of MTb= 5 W m. Table 2 also presents values of oxygen c o n s u m p t i o n ( t h e r m o n e u t r a l rate) expressed in ml kg t min -~. rat a n d dog. a value which is consistent with o u r o w n m e a s u r e m e n t s (unpublished) of the relationship between surface area a n d mass of the foal. Like the lamb a n d calf. total body resistanceand range of lower critical temperature values for pony foals (age 2-4 and 7-9 days) Foal Mean body Mean surface Resting Mean value Mean value M i n i m u m Maximum age mass area metabolic rate Mtn of rioI of Tz value of Tic value of Tic (d) (kg) (m2) (W m 2) (s m. the period over which foals can m a i n t a i n this high level of metabolism is not known. (1959) noted that the body tissue resistance of clipped sheep decreased w h e n they shivered. suggests t h a t the foals increased their utilization of muscle glycogen with shivering in the cold. tend to have p o o r thermal control and their lower critical temperatures are just a few degrees below their n o r m a l body-core temperature. respectively ( M o u n t . Values of rtot are based in part o n an animal's surface area. However.i ) (°C) (°C) (°C) 2--4 7-9 31. during which time its metabolic rate remained consistently elevated at a b o u t 100 W m -2. In the present study.93/ 598 Hey and O'Connell (1970) Cot-nursed Monkey < 1 35 9-10 28. 1979).8~ 448 Hey and O'Connell (1970) Clothed Human 3-10 25 ~7. 1983). which include the foal.43/ 381 Crighton and Pownall (1974) Sheep < 1 25 -60-70 235 Alexander (1974) Varies with breed Cow <1 9 -100 430 Webster (1974) Varied with breed Horse 2-4 22 7. Alexander a n d Williams. Mean body mass. However. resting metabolic rate. 02 ---20. Table 2 c o m p a r e s m e a n values of the resting metabolic rate for the foals studied here with values for the n e w b o r n of o t h e r species. which is usually estimated from M e e h ' s formula as 0.3 1. F o r c o m p a r a t i v e purposes. The decreases in the foals' insulation in the cold reported here were associated with strong or severe shivering a n d are consistent with these suggestions. 3/Values based on 1 I.5°C for 8.0 30. and is likely to be too low for the foal which has longer legs t h a n the " a v e r a g e animal".g. Precocious newborn. the third of these groups (e. The Table also shows the corresponding values (where available) of Tic.8 13.0 1.5 ~7. (1959) suggested t h a t this decrease in r s below thermoneutrality m a y be caused by a n increase in blood supply to the shivering muscles. observations m a d e o n lambs a n d calves indicate that.6 15. thus opening the capillary beds within the muscle. or to the release of heat from peripheral muscles. marsupials) having virtually no thermal control at birth. by increasing metabolic heat p r o d u c t i o n to a rate which is a b o u t twice that in the thermoneutral zone. Vermorel et al.2 was used to calculate rtot for infants).5:~ 209 Dawes (1968) Pig < 1 34 11 40. are more developed at birth t h a n altricial n e w b o r n a n d can m a i n t a i n h o m e o t h e r m y over a wider range o f air temperatures.0 30. one foal remained in 12. Pilo-erection a n d vasoconstriction enable a n animal to increase re a n d rs.094 is a n average for different species. the t h e r m o n e u t r a l range a n d values of total body resistance rtot (estimated from e q u a t i o n (4)) for these species.t rate resistance rtot Species (d) (°C) rain-] ) (Wm -2) (S m -] ) Reference Comments Human 3-10 32.g.0 73 333 Present study Varies between individuals *Resting values in thermoneutral range. calves).0 23. 1968. reported here for b o t h age groups. a n d M c A r t h u r (1987) presented similar evidence for cattle. Alexander (1975) classified n e w b o r n animals on the basis of their t h e r m o r e g u l a t o r y capacity as altricial.63/ 194 Mount (1959) Dog 3 30 14. in these species. there is a cold-induced decrease in R Q associated with non-shivering thermogenesis (NST) and oxidation of fatty acids (Alexander.094 m 0"67.7 34. the resistance rtot decreased by nearly 30% when air temperature was reduced below T~c.1 26. Biaxter et al.
14.C. 251---276. Saido and Demigne C. Alexander G. Cambridge University Press. J. III. 382 389. Pitmans. These figures apply only to foals which are dry. (1976) Energy expenditure during the growing period. Nutr. 624i8. Alexander G. Fert. (1986) Acclimatization to warm or cold temperatures and the r.) 180. Biol. (1983) Lactation in the horse: milk composition and intake by foals. 1152 1174. Mount L. Cambridge. Cambridge. 173 203. Biol. (1978) Temperature Regulation and Energy Metabolism in the Newborn.. In Heat Loss from Animals and Man (Edited by Monteith J. OUSEYet al. (Camb) 52. 23. (I 959) The metabolic rate of the new-born pig in relation to environmental temperature and age.. Alternatively. N.l0 J. pp. Like other precocious newborn. A.. L. 198.. Year Book Medical Inc. L. Alexander G. Alexander G. pp. Blackwell. 18 22. pp. 39. Figure 4 shows a positive correlation between heart rate and metabolic rate for the foals. Arch. and Williams D.\. L. L. 205 231. J. and Clark J. 31.). REFERENCES Alexander G. 45. J. F. Arnold. (1974) tteat loss m cattle with particular emphasis on the effects of cold.L theor. Bull. re-breathing o f some carbon dioxide in the ventilated mask could account for these low values of RQ but the flow rate through the mask was considered high enough to prevent this. 21.). (1968) Oxygen consumption and temperature regulation in the newborn. E. body temperature and respiratory quotient. and Mount L. Dawes G. Butterworths. II. (1959) Environmental temperature. Newmarket. 236 265.! 123. Wiley. G. Effect of environmental temperature on metabolic rate. Crighton G. McArthur A. or exposure to o u t d o o r conditions. therm. Med. Three values o f RQ in Fig. C. Chicago. Aust. Di~'. J Reprod. Meeh K. 335 343. (1968) Shivering and nonshivering thermogenesis during summit metabolism in young lambs. R. . In conclusion.7. Phys. and Rossdale P. a b o u t 20°C. Our results indicate that healthy p o n y foals can thermoregulate effectively at air temperatures as low as 2°C for periods o f several hours. In Early Nutrition and Later Development (Edited by Wilkinson A. and Cole 1". 147. Paediatrics 23. McArthur A. A change in posture to standing. London. (1959)The physical environment and the premature infant. Ousey J. J. E. 156 163. C. lI. J. J. Res. The authors thank Dr J. A. (Edited by Monteith J.. and Unsworth M. J. The . J. In Environmental . 113. (1879) Oberfl/ichemnessungel~ des mcnschlichen K6rpers. Aust. 142 144 Dauncey M. and Rahimtulla K. 2096 2106. J. In Foetal and Neonatal Physiology. Values o f T~ range from about 26°C (low insulation) to about I Y C (high insulation). Silverman W. However. 1961). 565 591. because o f differences in body insulation. Armitage P. Hintz H. (1979) Adaptation to the Thermal Environment. Oftedal O. (1981) The environmental physiology of juvenile animals. J. Butterworths. pp. (1978) Thermal insulation of animal coats and human clothing. Murgatroyd P. 425 458 Monteith . (1962) Temperature regulation in the newborn lamb. Med. il99(I) Principles ~)l Environmental Physics. Br. and Pownall R (1974) The homeothermic status of the neonatal dog. J. (1961) The Fire q/'L!/~. I~!. V~t. McC.. Ann./ therm Biol. Cambridge University and Dunn Clinical Nutrition Centre. Webster (1967) reported differences in the regression equations between heart rate and metabolic rate derived for individual sheep during cold exposure. Physiol. The scatter o f points about the lines fitted in Fig. (1978) A human calorimeter for the direct and indirect measurement of 24 h energy expenditure. Biol. Hill J.. E. These unusually low values could be the consequence of carbohydrate p r o d u c t i o n from fat (Kleiber.). However. (1975) Body temperature control in mammalian young.l~peets ~!/ Housing fi*r Animal Production (Edited by ('lark ! . Cena K.. 769 785. A.1. F. (1974) Heat loss from sheep. Clark for his comments on the manuscript and the numerous people who assisted with the study. Br. Dardillat C. New York. 4 is likely to be a consequence o f biological variability between individuals (data for up to 24 animals are shown on the same graph). D. the foal can increase its metabolic rate markedly in response to cold. J. Childh. J. E. 25~10. 89 93 Dauncey M. (1987) Thermal interaction between animal and microclimate: a comprehensive model. Physiol. Vernet J. 1979). London. (1983) Energy metabolism and thermoregulation in the newborn calf. Mount L. J. J Nutr. Hey E. Blaxter K. Summit metabolism. Butterworths. Sci. (1965) Heat balance and the metabolic rate of new-born babies in relation to environmental temperature. L. Grune & Stratton. W. J. (1967) Continuous measurement of heart rate as an indicator of the energy expenditure of sheep. energy metabolism and heat regulation in sheep. 12. (1961) Temperature regulation in the newborn lamb. ( 1991 b) Thermal intcrtlction between animal and microclimate: specification of a "'standard environmental temperature" for animals outd~ors J. Nutr.. Arnold. 557 566. and Ingrain D. J Physiol. on average. Suppl. agrie.)251. W. R. (Lond. 561-570. Oxford. theor Biol. (1968) Clinical studies on the newborn Thoroughbred foal: thermal stabilit\ B~ ~ur . and the effect of age and of weight on basal metabolic rate. (1991a) Metabolism of homeothemas in the cold and estimation of lhermal insulation . Z. 100 121. Kleiber M. 331 343. W. agric. 148. 149 155. Nature (Lond. 126. Webster A. 203 238. 44. S. Acknowledgements--The Horserace Betting Levy Board and Darley Studfarm Management provided financial support for this project and facilities were made available by the Animal Health Trust. Vermorel M.345. 333. New York. (1970) Oxygen consumption and heat balance in the cot-nursed baby. Graham N. 2nd edn. and Schryver H F. T~ can differ considerably between individual foals. Br. London Rossdale P D. McArthur A. 191 209. A.. Res. (1989) Energy Metablism in Animals and Man. London. Biol. Poczopko P. Reeh. and Mount L. T. V. F.. may alter the values o f TI~ considerably. 166 171 Sinclair J. 16. McArthur A. Man and his Productive Animals. E. J. we report that the lower critical temperature for healthy young pony foals is. Wainman F. 3 are below 0. London. J. (1971 ) Statistical Methods for Medical Research. In Heat Loss from Animals and Man. and Armstrong D. A linear relationship between these two quantities has been established for other species (Mount.)le of food intake.). partition of heat losses in closely clipped sheep. 13. pp 109 130. agric. recumbent and exposed to still or slowly moving air (no sunlight). 15. and O'Connell B. London heat production in foals exposed to cold. Webster A. Blaxter K. Mount L. (1991) Metabolic changes in Thoroughbred and pony foals during the first 24 h post partum.
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