The Circadian Rhythm of Body Temperature of the Horse

Rectal temperature of 10 female adult horses was recorded every 2 h for 10 consecutive days under a natural winter photoperiod (9 h of light and 15 h of darkness per day). A robust daily rhythm of body temperature was observed in all animals. The rhythm had a mean level of 38.3°C and a range of excursion of 1.0°C. Temperature started its daily ascent at dawn each day and reached a maximum 14 hours later. Body temperature of 5 of the horses was studied for 10 more days under constant illumination. The rhythm persisted under this condition, although with a slightly longer period of 24.2 h, which confirms the endogenous nature of the rhythm. Despite the fact that the body size of the horse is several orders of magnitude greater than that of rodents, the various parameters of the body temperature rhythm of the horse are similar to those of several species of rodents previously studied.     

Circadian modulation of starvation-induced hypothermia in sheep and goats

Prolonged food deprivation is known to cause a fall in the core body temperature of homeotherms. In various species of small birds and mammals (body mass up to 2-3 kg), it has been shown that starvation-induced hypothermia is modulated by the circadian system, in the sense that hypothermia is observed primarily during the inactive phase of the daily activity cycle (i.e., during the night for diurnal animals and during the day for nocturnal animals), whereas relatively normal temperatures are recorded during the active phase. To investigate whether this modulation occurs also in larger animals, we investigated the effects of 4d food deprivation on the body temperature rhythm of goats and sheep (body mass 30-40 kg). In goats, the body temperature rhythm was found to have a mean level of 39.0°C with a mean daily range of excursion of 0.42°C. The daily oscillation in body temperature persisted during the first day of fasting, but the rhythm was drastically damped, if not eliminated, over the next 3 d as body temperature descended from the baseline level of 39.0 to 38.2°C. In sheep, the rhythm was found to have a mean level of 39.3°C with a mean daily range of excursion of 0.34°C. The daily oscillation in body temperature persisted through the 4 d of food deprivation, even though the mean level of body temperature gradually fell. Temperature fell more during the third and fourth nights than during the third and fourth days. Thus, circadian modulation of starvation-induced hypothermia was observed in sheep but not in goats.     

Influence of fasting and exercise on the daily rhythm of serum leptin in the horse

The hormone leptin is secreted by white adipocytes and regulates food intake and energy expenditure in rodents and humans. The goal of the present study was to investigate the existence of a daily rhythm of serum leptin in horses and its dependence on fasting and physical exercise. A robust daily rhythm of leptin was found in both athletic and sedentary horses, with a daytime trough and a peak in the dark phase. While physical exercise never induced changes in circulating leptin, fasting reliably affected serum leptin levels. Food deprivation did not abolish the daily rhythm of serum leptin, but daily mean leptin levels in fasted horses were significantly lower than in regularly fed horses. This result indicates that leptin production is not a mere consequence of feeding behavior. The fact that in a large animal such as the horse a short fast decreases leptin without significantly changing the body weight demonstrates that changes in levels of circulating leptin associated with food restriction do not solely reflect changes in amount of body fat.     

Change of serum mitochondrial uncoupling protein 1 (UCP1) levels and daily rhythm of rectal and cutaneous temperatures in Equus caballus and Capra hyrcus

The aim of the present study was to monitor the daily rhythm of rectal and cutaneous temperatures together with the changes in the serum levels of mitochondrial uncoupling protein 1 (UCP1) in equine and goat species. On five gelding horses and five female goat rectal and cutaneous temperatures were recorded at 4 h intervals for a 48 h period. At the same time points blood samples were collected from each animal. Daily rhythms of rectal and cutaneous temperatures were observed in both species, in both day of monitoring. Mesor value of rectal temperature was statistically higher of about 4 °C than cutaneous temperature, and acrophase was postponed of about 2 h in both day of monitoring and in both species. UCP1 did not show daily rhythmicity in horses and goats. We can speculate that the thermogenesis due to thermal system is auxiliary to keeping the body temperature daily rhythm.     

Comparison of daily rhythm of rectal and auricular temperatures in horses kept under a natural photoperiod and constant darkness

The aim of this study was to assess the accuracy of auricular temperature (AT) recording in the determination of body temperature as well as rectal temperature (RT). For this purpose we compared RT and AT in five clinically healthy horses. Data collections were performed every 3 h over 2 different 24 h photoperiods, 13/11 light/dark cycle and constant darkness. Repeated measures multifactor analysis of variance (MANOVA) was applied to determine a statistical significant effect of time of day, side of temperature collection and photoperiods on AT and RT. Our finding showed a significant effect of time of day on the temperature values recorded and an influence of side of temperature collection. RT values were higher of about 4 °C than the AT values, and the pattern of the two temperatures was not comparable. A daily rhythmicity of rectal temperature was observed. Auricular temperature no showed daily rhythmicity in both periods of monitoring. On the basis of these findings AT does not reflect body temperature as well as RT in horse.     

Daily and estrous rhythmicity of body temperature in domestic cattle

Background

Rhythmicity in core body temperature has been extensively studied in humans and laboratory animals but much less in farm animals. Extending the study of rhythmicity of body temperature to farm animals is important not only from a comparative perspective but also from an economic perspective, as greater knowledge of this process can lead to improvements in livestock production practices. In this study in cattle, we investigated the maturation of the daily rhythm of body temperature in newborn calves, characterized the parameters of the daily rhythm in young cows, and studied the oscillation in body temperature associated with the estrous cycle in adult cows.

Results

We found that the daily rhythm of body temperature is absent at birth but matures fully during the first two months of life. The mature rhythm had a mean level of 38.3°C, a range of excursion of 1.4°C, and was more robust than that of any mammalian species previously studied (90% of maximal robustness). Sexually mature cows also exhibited a robust estrous rhythm of body temperature. An elevation of about 1.3°C was observed every 21 days on the day of estrus. Small seasonal variations in this pattern were observed.

Conclusion

In conclusion, calves exhibit a very robust daily rhythm of body temperature, although this rhythm is absent at birth and develops during the first two months of life. Adult cows exhibit also 21-day rhythmicity in body temperature reflecting the duration of the estrous cycle.

     

Daily rhythms of oxygen consumption, body temperature, activity and melatonin in the Norwegian lemming Lemmus lemmus under northern summer photoperiod

Rodents inhabiting high latitudes, close to the Arctic Circle or above it, are exposed to near 24 h daylight during the summer season. An example to such rodent species is the Norwegian lemming Lemmus lemmus, which is distributed in northern Fennoscandia. We measured daily rhythms of heat production (VO₂), body temperature (T_b), motor activity and melatonin secretion (measured from its metabolite 6-sulfatoximelatonin 6-SMT) in individuals exposed to natural day light, Oulu Finland, in August 1997 and at a controlled ambient temperature of 22 °C. Our results show a daily rhythm of VO₂ with an acrophase at 04:00 h and minimal values measured at 16:00 h, with a significant difference (p<0.001) between mean day and night values. 6-SMT also shows a daily rhythm with maximal secretion during the dark phase 24:00 and 06:00 h in which total 6-SMT values were 2.92±1.1 and 3.87±1.2 ng, respectively. The lowest values were recorded at 12:00 h; 0.86±0.63 ng. These results suggest that lemmings show a VO₂ and melatonin daily rhythms, which seem to correlate with each other and it appears that melatonin secretion increases heat production.     

Daily rhythm of body and auricle temperature in goats kept at two different ambient temperatures

In fluctuating environmental temperature, homeotherms are able to maintain stable their body temperature, which however reveals a rhythmic daily pattern as described in literature. Because of the importance of body temperature rhythmicity in the knowledge of thermal homeostasis and as a means to facilitate the study of biological rhythmicity in general, the aim of our study was to assess the influence of two different ambient temperature on the daily rhythmicity of body and auricle temperature in goat (Capra hircus). For our study 6 female adult Maltese goats (18 month old), not pregnant and clinically healthy were used. On each subject body and auricle temperature was recorded during two different periods, of 7 days each, every 3 hours for 24 hours from 07.00 to 04.00, at different environmental temperature (1^st period 18.25 ± 1.48°C, 2^nd period 28.25 ± 2.05°C). The analysis of the obtained result under two ambient temperature indicates the existence of a daily periodicity, with small differences in amplitude and acrophase between the two studied period, both in body and auricle temperature in Maltese breed goat so we can assume that environmental temperature can influence in a significant way the daily pattern of body and auricle temperature.     

Infrared methodologies for the assessment of skin temperature daily rhythm in two domestic mammalian species

To assess the accuracy of infrared methodologies for daily rhythm monitoring of skin temperature, five clinically healthy Italian Saddle gelding horses, and five not pregnant and not lactating Camosciata goats, were monitored every 4 h over a 48 h period. The horses were housed in individual boxes, while the goats in two indoor pens, under natural photoperiod and natural environmental temperature. In each animal, skin temperature was recorded with the use of a digital infrared camera and a non-contact infrared thermometer, in five regions: neck, shoulder, ribs, flank and croup. Recorded values were compared with the well-established daily rhythm of rectal temperature. Rectal temperature was recorded at the same time by means of a digital thermometer. In horses, a lower value of skin temperature was recorded using the infrared thermometer for the croup region compared to shoulder and flank; a lower value of skin temperature was recorded using thermography for the croup region compared to the shoulder. In goats, a lower value of skin temperature was recorded using the infrared thermometer for the croup region compared to the flank. In both species, higher values of rectal temperature were observed, compared to the temperature recorded at the skin regions using the other two methodologies. Cosinor rhythmometry showed a daily rhythm of rectal and skin temperature recorded using both methodologies in all the examined regions. General linear model (GLM) showed statistically significant effect of breed on all rhythmic parameters; of day of monitoring on amplitude; of site of recording (rectal vs skin regions) on mesor, amplitude and acrophase; and no effect of methodologies used. The results of this study show the differences in rhythmicity of various body regions temperature and their differences in comparison with daily rhythm rectal temperature. The use of infrared methodologies was inaccurate in assessing body core temperature, but its use could be considered for the evaluation of inflammation in the different body sites.     

Effects of temperature, photoperiod, and light intensity on the eclosion rhythm of the high-altitude Himalayan strain of Drosophila ananassae

Eclosion rhythm of the high-altitude Himalayan strain of Drosophila ananassae from Badrinath (altitude 5123 m) was temperature-dependent and at 21°C, it was entrained by cycles of 12 h light: 12 h darkness (LD 12:12) and free-ran in constant darkness, however, it was arrhythmic at 13°C or 17°C under identical experimental conditions (Khare, P. V., Barnabas, R. J., Kanojiya, M., Kulkarni, A. D., Joshi, D. S. (2002). Temperature dependent eclosion rhythmicity in the high altitude Himalayan strains of Drosophila ananassae. Chronobiol. Int. 19:1041-1052). The present studies were designed to see whether or not these strains could be entrained at 13°C, 17°C, and 21°C by two types of LD cycles in which the photoperiod at 100 lux intensity varied from 6 h to 18 h, and the light intensity of LD 14:10 cycles varied from 0.001 lux to 1000 lux. All LD cycles entrained this strain at 21°C but not at 13°C or 17°C. These results demonstrate that the entrainment of eclosion rhythm depends on the ambient temperature and not on the photoperiod or light intensity of LD cycles. Thus the temperature has taken precedence over the light in the entrainment process of eclosion rhythm of the high altitude Himalayan strain of D. ananassae. This may be the result of natural selection in response to the environmental temperature at Badrinath that resembles that of the sub-Arctic region but the photoperiod or light intensity are of the subtropical region.     

Rhythmic cortisol secretion in the equine: Analysis and physiological mechanisms

Abstract

Four Thoroughbred mares (no. 1–4) were maintained under constant temperature (24°C) and controlled light (L/D:12/12 with lights on at 06.00 hr) conditions. They were fed and watered ad libitum with fresh feed and water given at 09.00 hr. After a 45‐day pre‐conditioning period, blood samples were obtained by veinipuncture at 4‐hr intervals for 14 days to determine circadian and day‐to‐day variation. The horses exhibited a circadian rhythm with maximum values attained at about 12.00 hr, however, there are periods of days in which no rhythm is distinguishable. Ultradian rhythms with mean periods of 105 to 128 and 24 to 31 min are superimposed upon the circadian rhythm. The individual rhythms are quite variable from horse to horse and within the same horse. During periods of decline in plasma cortisol with metabolic half‐lives of approximately 70 min, secretion of cortisol was very low or had ceased. During periods of increasing plasma concentration, secretion was occurring at a faster rate than degradation. Rapid decreases in plasma concentration (metabolic half‐life of approximately 30 min) was accompanied by a rise in specific activity indicating cortisol with a high specific activity was entering the plasma pool from other storage pools.     

Different behavior of body temperature and total locomotor activity daily rhythms during light/dark cycle in stabled Oryctolagus cuniculus

In order to investigate the potential causal link between the rhythm of activity and body temperature, we simultaneously recorded rectal temperature and total locomotor activity in five clinically healthy female rabbits (blue Vienna breed), 12 week old and mean body weight 2.7 ± 0.3. Animals were housed in individual cages (90?×?50?×?35 cm) under natural 12/12 light/dark cycle. Total locomotor activity was monitored for 15 days by an activity data logger. On day 1, 5, 10, and 15 rectal temperature was recorded every 2 h for a 24-h period. Application of single cosinor method showed a nocturnal daily rhythm of rectal temperature with a range of oscillation of about 1 °C, acrophase after dusk and low robustness value. The daily rhythm of locomotor activity showed its acrophase in the middle of the scotophase and a high robustness value. This information improves the knowledge available on the circadian biology of rabbits useful in the evaluation of physiology of this species.     

Light and temperature cycles as zeitgebers of zebrafish (Danio rerio) circadian activity rhythms

Light and temperature cycles are the most important synchronizers of biological rhythms in nature. However, the relative importance of each, especially when they are not in phase, has been poorly studied. The aim of this study was to analyze the entrainment of daily locomotor activity to light and/or temperature cycles in zebrafish. Under two constant temperatures (20°C and 26°C) and 12:12 light-dark (LD) cycles, zebrafish were most active during the day (light) time and showed higher total activity at the warmer temperature, while diurnalism was higher at 20°C than at 26°C (87% and 77%, respectively). Under thermocycles (12:12 LD, 26:20°C thermophase:chryophase or TC), zebrafish daily activity synchronized to the light phase, both when the thermophase and light phase were in phase (LD/TC) or in antiphase (LD/CT). Under constant dim light (3 lux), nearly all zebrafish synchronized to thermocycles (τ=24 h), although activity rhythms (60% to 67% of activity occurred during the thermophase) were not as marked as those observed under the LD cycle. Under constant dim light of 3 lux and constant temperature (22.5°C), 4 of 6 groups of zebrafish previously entrained to thermocycles displayed free-running rhythms (τ=22.9 to 23.6 h). These results indicate that temperature cycles alone can also entrain zebrafish locomotor activity.     

Equine sweating responses to submaximal exercise during 21 days of heat acclimation.

This study examined sweating responses in six exercise-trained horses during 21 consecutive days (4 h/day) of exposure to, and daily exercise in, hot humid conditions (32-34 degrees C, 80-85% relative humidity). On days 0, 3, 7, 14, and 21, horses completed a standardized exercise test on a treadmill (6 degrees incline) at a speed eliciting 50% of maximal O(2) uptake until a pulmonary artery temperature of 41.5 degrees C was attained. Sweat was collected at rest, every 5 min during exercise, and during 1 h of standing recovery for measurement of ion composition (Na(+), K(+), and Cl(-)) and sweating rate (SR). There was no change in the mean time to reach a pulmonary artery temperature of 41.5 degrees C (range 19.09 +/- 1.41 min on day 0 to 20.92 +/- 1.98 min on day 3). Peak SR during exercise (ml. m(-2). min(-1)) increased on day 7 (57.5 +/- 5. 0) but was not different on day 21 (48.0 +/- 4.7) compared with day 0 (52.0 +/- 3.4). Heat acclimation resulted in a 17% decline in SR during recovery and decreases in body mass and sweat fluid losses during the standardized exercise test of 25 and 22%, respectively, by day 21. By day 21, there was also a 10% decrease in mean sweat Na(+) concentration for a given SR during exercise and recovery; this contributed to an approximately 26% decrease in calculated total sweat ion losses (3,112 +/- 114 mmol on day 0 vs. 2,295 +/- 107 mmol on day 21). By day 21, there was a decrease in sweating threshold ( approximately 1 degrees C) but no change in sweat sensitivity. It is concluded that horses responded to 21 days of acclimation to, and exercise in, hot humid conditions with a reduction in sweat ion losses attributed to decreases in sweat Na(+) concentration and SR during recovery.     

Effect of ambient temperature on the circadian activity rhythm in common marmosets, Callithrix j. jacchus (primates)

Whereas the (zeitgeber) effect of ambient temperature T_a and temperature cycles TaC's on circadian rhythmicity has been well documented for heterofhermic mammals, inconsistent results have been obtained for strictly homeothermic species. Hence, it might be inferred that the susceptibility of the mammalian circadian timing system (CTS) to Ta and TaC's depends on the range of the animals' core and/or brain temperature rhythm. This hypothesis was tested in the common marmoset (Callithrix j. jacchus, n=12), a small diurnal primate with an amplitude in body temperature rhythm that is larger than for other homeothermic primates studied so far. Within the range 20-30°C, no systematic effects of constant Ta on most parameters of the marmosets' light-dark (LD)-entrained and free-running circadian activity rhythm (CAR) were found. Significant differences could be established in the average amount of activity per circadian cycle. It was highest at Ta 25°C (LD) and 20°C (light-light, LL) and most probably reflected a temperature-induced masking effect. A 24h trapezoidal TaC of 20:30°C entrained the free-running CAR in two of six marmosets and produced relative coordination in all others. Accordingly, in all animals tested, it had an effect on the CTS. In marmosets free running in LL at a Ta of 20°C or 30°C, 3h warm and cold pulses of 30°C and 20°C, respectively, produced neither systematic phase responses nor period responses of the CAR. So, there is no evidence of a phase-response mechanism underlying circadian entrainment. The results show that large-amplitude TaC's function as a weak zeitgeber for the marmosets' CTS. Since this zeitgeber effect is significantly larger than that found in owl monkeys, the results are consistent with the starting hypothesis that the zeitgeber effect of a given T,C on the mammalian CTS may be related to the amplitude of the species' core and/or brain temperature cycle.     

Effects of feeding frequency and voluntary salt intake on fluid and electrolyte regulation in athletic horses.

The effect of feeding frequency and voluntary sodium intake (VSI) on fluid shifts and plasma aldosterone concentration (PAC) were studied at rest and after exercise in six athletic horses. The horses were fed twice a day (2TD) and six times a day (6TD) for 25 days for each protocol, according to a changeover design. VSI was measured by weighing each horse's salt block daily. Feeding 2TD or 6TD caused no major alterations in fluid shifts, but in the 2TD treatment there was a postprandial increase in plasma protein concentration and osmolality that lasted <1 h. PAC and VSI were not affected by feeding frequency. VSI ranged from 0 to 62 mg x kg body weight-1 x day-1 and caused significant alterations in PAC. At VSI <26 mg x kg body weight-1 x day-1, a diurnal rhythm for PAC was noted. Water intake, fecal concentrations of sodium and potassium, and packed cell volume during exercise were influenced by VSI. The response to exercise did not differ between treatments. In conclusion, VSI, but not feeding frequency, has significant effects on fluid and electrolyte regulation in athletic horses.     

The Effect of Activity on the Waking Temperature Rhythm in Humans

Nine healthy female subjects were studied when exposed to the natural light-dark cycle, but living for 17 “days” on a 27h day (9h sleep, 18h wake). Since the circadian endogenous oscillator cannot entrain to this imposed period, forced desynchronization between the sleep/activity cycle and the endogenous circadian temperature rhythm took place. This enabled the effects of activity on core temperature to be assessed at different endogenous circadian phases and at different stages of the sleep/activity cycle. Rectal temperature was measured at 6-minute intervals, and the activity of the nondominant wrist was summed at 1-minute intervals. Each waking span was divided into overlapping 3h sections, and each section was submitted to linear regression analysis between the rectal temperatures and the total activity in the previous 30 minutes. From this analysis were obtained the gradient (of the change in rectal temperature produced by a unit change in activity) and the intercept (the rectal temperature predicted when activity was zero). The gradients were subjected to a two-factor analysis of variance (ANOVA) (circadian phase/ time awake). There was no significant effect of time awake, but circadian phase was highly significant statistically. Post hoc tests (Newman-Keuls) indicated that gradients around the temperature peak were significantly less than those around its trough. The intercepts formed a sinusoid that, for the group, showed a mesor (±SE) of 36.97 (±0.12) and amplitude (95% confidence interval) of 0.22°C (0.12°C, 0.32°C). We conclude that this is a further method for removing masking effects from circadian temperature rhythm data in order to assess its endogenous component, a method that can be used when subjects are able to live normally. We suggest also that the decreased effect of activity on temperature when the endogenous circadian rhythm and activity are at their peak will reduce the possibility of hyperthermia.     

Effects of hair coat characteristics on radiant surface temperature in horses

Horse owners may lack knowledge about natural thermoregulation mechanisms in horses. Horses are managed intensively; usually stabled at night and turned out during the day. Some are clipped and many wear a blanket, practices which reduce the horse's ability to regulate heat dissipation. The aim of this study was to investigate the relationship between hair coat characteristics, body condition and infrared surface temperatures from different body parts of horses. Under standard conditions, the body surface temperature of 21 adult horses were investigated using infrared thermography. From several readings on the same body part, a mean temperature was calculated for each body part per horse. Detailed information on horse breed, age, management and body condition was collected. Hair coat samples were also taken for analyses. A mixed statistical model was applied. Warmblood horse types (WB) had lower hair coat sample weights and shorter hair length than coldblood horse types (CB). The highest radiant surface temperatures were found at the chest 22.5 ± 0.9 °C and shoulders 20.4 ± 1.1 °C and WB horses had significantly higher surface temperatures than CB horses on the rump (P < 0.05). Horses with a higher hair coat sample weight had a lower surface temperature (P < 0.001) and hind hooves with iron shoes had a significant lower surface temperature than unshod hind hooves (P = 0.03). In conclusion, individual assessment of radiant surface temperature using infrared thermography might be a promising tool to gather data on heat loss from the horses' body. Such data may be important for management advice, as the results showed individual differences in hair coat characteristics and body condition in horses of similar breeds.     

The effects of constant and changing tempertures on the development of eggs of the freshwater snail Lymnaea auricularia (L.)

The rate of development of Lymnaea auricularia eggs was studied at various constant temperatures between 10° and 36°C. Development was accelerated as the temperature increased and at 36°C the eggs failed to develop. Spring eggs showed differences in their rate of development when compared with summer eggs when measured at similar tempertures.

Both spring and summer eggs were more than 90% fertile. Hatching success was high at temperatures between 10° and 30° (100%–82/9%); while at 34°C it was reduced to 60.6% for spring eggs. It was above 87% at temperatures between 10° and 34°C but it dropped to 62.3% at 36°C for summer eggs.

In one regularly changing temperature experiment a significant acceleration (P < .05) was found. In two others there was no significant difference beween predicted and observed egg durations. In one suddenly changing temperature regime (1 day at 20°, 1 day at 30° and so on) a huge retardation of development was found. In the other suddenly changing experiment (1 day at 15°, 1 day at 25°) no significant difference was found.

The exposure of eggs to extreme temperature (4°C, freezing and 4°C caused a retardation in the race of subsequent development of eggs at 25°C.