Daily rhythms of nonshivering thermogenesis and responses to photoperiod manipulations in Apodemus mystacinus from two different ecosystems

Comparative physiology is an important tool for understanding adaptation to environment. Regulatory nonshivering thermogenesis (NST) is an essential mechanism in small mammals for coping with low ambient temperatures. Because of its geographical location and its sharp climatic changes across short distances, Israel offers conducting comparative studies between different populations of the same species. The aims of this study were: (1) to compare NST-capacity daily rhythms, between the two populations of the broad-toothed field mouse Apodemus Mystacinus—that of Mount Carmel (Mediterranean) and that of Mount Hermon (Subalpinic), (2) to compare the thermoregulatory daily rhythms response to photoperiod manipulations of mice from the two populations.

Mice were acclimated for at least 3 weeks to long scotophase (16D:8L-LS) and then to long photophase (16L:8D-LP) at a constant ambient temperature (T_a) of 24±1 °C. The following variables were compared at four different times of the day (06, 12, 18, 24 h): minimal body temperature (T_bMin), minimal oxygen consumption (VO₂Min) measured at the lower critical point and their response to noradrenaline (NA), 1.5 mg/1 kg.W_b injected s.c., (VO₂NA and T_bNA). NST-capacity was calculated as the ratio between VO₂NA and VO₂Min. The measurements were carried out at T_a of 28 °C and in light conditions matching those of acclimation. NA was injected after establishing VO₂Min and T_bMin. The values mean±SD are given for each measurement for n=6.

A three-way ANOVA revealed a significant difference between the two populations. Significant differences were revealed at different hours within populations under the different photoperiod regimes for different measured variables.     

Daily activity and body temperature

Body temperature varies between 36 and 39° C in states ranging from sleep to high levels of sustained exercise, but it is not known whether this continuum of body temperature is related to a continuum of activity. Calorimetric studies of sedentary days were undertaken with four levels of food intake, men doing mild sustained exercise, and men and women walking and cycling vigorously. Steady states of metabolism were followed by slow exponential changes to steady states of heat loss (Q), followed in turn by changes in rectal temperature (T _re). Regression analysis showed a continuous, curvilinear relationship between Q andT _re from the low end of the activity spectrum (50 W) to progressively higher levels of exercise (600 W). These related continua of activity and body temperature appear to be the result of heat regulation.     

Pineal and retinal melatonin is involved in the control of circadian locomotor activity and body temperature rhythms in the pigeon

Summary Although pinealectomy or blinding resulted in loss of the clarity of the free-running rhythm of locomotor activity and body temperature and reduced the peak level of circulating melatonin rhythms to approximately a half in intact pigeons, neither pinealectomy nor blinding abolished any of these rhythms. However, when pinealectomy and blinding were combined, the rhythms of locomotor activity and body temperature disappeared in prolonged constant dim light, and melatonin concentration was reduced to the minimum level of detection. In order to examine the role of melatonin in the pigeon's circadian system, it was administered either daily or continuously to PX + EX-pigeons in LLdim. Daily administration of melatonin restored circadian rhythms of locomotor activity which entrained to melatonin injections, but continuous administration did not induce any remarkable change of locomotor activity. These results suggest that melatonin synthesized in the pineal body and the eye contributes to circulating melatonin and its rhythmicity is important for the control of circadian rhythms of locomotor activity and body temperature in the pigeon.Abbreviations LD Light-dark - LLdim constant dim light - LLbright constant bright light - PX pinealectomy - EX blinding - SCN suprachiasmatic nucleus     

Daily torpor in relation to photoperiod in a subtropical blossom-bat, Syconycteris australis (Megachiroptera)

Daily torpor in many temperate-zone mammals is affected by photoperiod. As little is known about the effects of photoperiod on torpor in subtropical species, we investigated whether, and if so how, torpor use, duration, and depth are affected by acclimation to three photoperiods (short, intermediate, long) in the blossom-bat Syconycteris australis. In contrast to many other studies, torpor occurrence, duration, and depth did not significantly respond to photoperiod acclimation in S. australis. Interestingly, the trend of a decline in torpor use under long photoperiod was the opposite of that observed previously in S. australis, which had been captured from the wild in summer and winter. Our study suggests that some species living in low latitude areas with unpredictable weather like S. australis may not use photoperiod for seasonal adjustments in physiology because it is not a reliable cue for food availability.     

Daily and seasonal cycles of body temperature and aspects of heterothermy in the hedgehog Erinaceus europaeus

Summary Intra-abdominal temperature-sensitive radio transmitters were used to collect more than 350 sets of body temperature (T _b ) data from 23 captive adult hedgehogs over a 3-year period. Each data set comprised measurements made every 1/2 h for 24-h periods. Between 20 and 60 such data sets were recorded every calendar month, and a total of 17400 measurements of T _b were collected. The hedgehogs were exposed to natural environmental conditions at 57°N in NE Scotland. Hedgehogs showed seasonal changes in mean daily euthermic T _b ,with a July maximum of 35.9±0.2°C, a September minimum of 34.7±0.9°C, and a marked circadian T _b cycle that correlates closely with photoperiod. Maximal T _b occurred within 2 h of midnight and this pattern of nocturnal maximum and diurnal minimum T _b was most marked between April and September. The circadian T _b cycle was least correlated with photoperiod during winter. Hibernal T _b during winter correlated with ambient temperature (T _a ),it was maximal in September (17.7±1.0°C) and minimal in December (5.2±0.9°C). Apart from the tracking of T _a and T _b during hibernal bouts, with a time-lag of 4–6 h, circadian rhythmicity of hibernal T _b was not evident. However, the T _b of hibernating hedgehogs rose significantly when T _a fell below — 5°C, although the animals did not neccessarily arouse. Although hibernal bouts occurred between September and April, 89.5% of such bouts were recorded between November and February. The mean time of entry into hibernation was 01:45±5.1 h GMT while the mean time of the start of spontaneous arousal from hibernation was 11:53±4.8 h GMT. Therefore, during hibernation hedgehogs were either fully aroused at night, when euthermic hedgehogs have maximalT _b ,or in deep hibernation around midday, when euthermic hedgehogs have minimal T _b .Since wild hedgehogs will feed during spontaneous arousal from hibernation, these timings are probably adaptive, and suggest that entry into, and arousal from, hibernation may be extensions of circadian cyclicity. Spontaneous bouts of transient shallow torpor (TST) were recorded throughout the year, with nearly 80% of observations occurring during August and September, at the start of the hibernal period. TST bouts lasted for 4.9±2.9 h, with T _b falling to 25.8±3.1 °C. Only 20% of TST bouts immediately preceded hibernation and their duration did not correlate with T _a or body mass. TST bouts started at 06:51±4.7 h GMT, significantly later than entry into hibernation, and ended at 13:04±5.4 h GMT. The function of TST bouts is unclear, but they may be preparation for the hibernation season or a further energy conservation strategy. When arousing from hibernation hedgehogs warmed at a rate of 1.9±0.4°C·h^-1, and when entering hibernation cooled at 7.9±1.9°C·h^-1. Warming rates were slightly higher during mid-winter when T _b and body mass were minimal, but cooling rates were 44% higher at the end of the hibernal period compared to the start. Cooling and warming rates were strikingly similar to those measured in hedgehogs at 31°N. These results demonstrate that thermoregulation in the hedgehog is closely regulated and changes on a seasonal basis, in meeting with requirements of surviving food shortages and low temperature during winter.Abbreviations T _a ambient temperature - T _b body temperature - CSD circular standard deviation - SWS slow wave sleep - TST transient shallow torpor     

Circadian rhythms of corneal mitotic rate,retinal melatonin and immunoreactive visual pigments,and the effects of melatonin on the rhythms in the Japanese quail

We investigated circadian ocular rhythms in the Japanese quail, Coturnix coturnix japonica. The birds were placed under light-dark cycles (LD 1212), constant light (LL) and constant darkness (DD), and the retinas were dissected out at four-hour intervals throughout 24 h. Following measurements were performed. (1) Melatonin content in the retina was measured by radioimmunoassay. It was low in light and several folds higher in darkness under LD 1212. The rhythm continued in DD, but disappeared in LL. (2) Mitotic figures in the corneal epithelium were counted. Similar rhythms to the melatonin content were observed in the corneal mitotic rate with a slight phase delay. (3) The retinas were fixed at 4-h intervals and immunostained with anti-bovine rhodopsin serum and anti-chicken iodopsin monoclonal antibodies. The outer segments of photoreceptor cells were stained intensively throughout 24 h in LD 1212, LL and DD. In contrast, the stainability of the locus close to the outer limiting membrane where the Golgi apparatus exists changed diurnally. Scores showing the ratio of cells with positive staining indicated high values from 4 h after the onset of light to the beginning of dark phase under LD 1212. The values were high throughout 24 h in LL and intermediate or low in DD. (4) To investigate the effect of melatonin on the corneal mitotic rate and visual pigments at the Golgi region, melatonin was injected into one eye and saline into the contralateral eye. Melatonin induced a phase advance in the corneal mitotic rate under LD 1212, but did not induce a rhythm under LL. The ratio of photoreceptor cells with positive staining to anti-visual pigment antibodies at the Golgi region was not affected by melatonin injection.Abbreviations ANOVA analysis of variance - BSA bovine serum albumin - DD constant darkness - Io-mAb monoclonal antibodies against chicken iodospin - LD light-dark - LL constant light - mRNA messenger ribonucleic acid - PBS phosphate buffer solution - Rh-As antiserum against bovine rhodopsin - SCN suprachiasmatic nucleus - T transducin - T transducin      

Impact of photoperiod manipulation on day/night changes in melatonin, sex steroids and vitellogenin plasma levels and spawning rhythms in Senegal sole, Solea senegalensis

Photoperiod and temperature are known as the main synchronizers of seasonal reproduction in fish. This paper studied the role of photoperiod on the synchronization of F1 Senegal sole reproduction rhythms. Fish were maintained under constant short-photoperiod (9L:15D) from the winter solstice onwards (experimental group) or under naturally-changing photoperiod (control group), and water temperature naturally oscillated in both groups. Blood samples were collected during the reproduction season at pre-spawning (March), spawning (April) and post-spawning (May) to determine the endocrine status. Spawning events and egg quality parameters were also monitored. The results revealed a significant increase in nocturnal melatonin concentration from March to May in the control group, while in the experimental group such seasonal change did not occur. As to plasma levels of vitellogenin, testosterone, estradiol and 11keto-testosterone, differences between groups were found mostly in March, while in April and May levels were often similar. Spawning was observed in both groups, although the experimental group started slightly earlier and also finished earlier than the control group, perhaps as a result of the increase in sex steroids and VTG observed at pre-spawning. Briefly, reproduction rhythms persisted in the absence of the natural lengthening of photoperiod, although photoperiod manipulation altered the seasonal modulation of melatonin, increased sex steroids and vitellogenin at pre-spawning, and slightly advanced the timing of spawning.     

Daily rhythms of body temperature and heat production of sibling Mastomys species from different ecosystems--the response to photoperiod manipulations

We compared body temperature (T_b) and metabolic rates, measured as oxygen consumption (VO₂), daily rhythms of two sibling species of the genus Mastomys. We also studied their responses to long day (16L: 8D, LD) and short day (8L: 16D, SD) photoperiod manipulations at a constant ambient temperature of 26 + 1 °C. We noted significant differences in T_b and VO₂ daily rhythm patterns, under SD and LD-acclimation between the sibling species. These differences explain adaptation to the climatic conditions that prevail in the different ecosystems where these species live. To the best of our knowledge, this is the first time that physiological differences between the two siblings are measured by using chronobiological methods.     

Thermal investigations of a honey bee colony: thermoregulation of the hive during summer and winter and heat production of members of different bee castes

Summary The temperature at the centre, the periphery and the entrance of a honey bee colony (Apis mellifera carnica) was continuously determined during the summer season and the broodless time in winter. During the summer season the temperature in the brood nest averages 35.5°C with brief excursions up to 37.0°C and down to 33.8°C. Increasing environmental temperatures resulted in linear increases in the temperature of the hive entrance, its periphery and its centre. The temperature in the centre of an overwintering cluster is maintained at an average value of 21.3°C (min 12.0°C, max 33.5°C). With rising ambient temperatures the central temperature of a winter cluster drops whereas the peripheral temperature increases slightly. With decreasing external temperatures the peripheral temperature is lowered by a small amount while the cluster's centre temperature is raised. Linear relationships are observed between the central and the ambient temperature and between the central temperature and the temperature difference of the peripheral and the ambient temperatures. The slopes point to two minimum threshold values for the central (15°C) and the peripheral temperature (5°C) which should not be transgressed in an overwintering cluster. Microcalorimetric determinations of the heat production were performed on the three castes of the honey bee: workers, drones and queens of different ages. Among these groups single adult workers showed the highest heat production rates (209 mW·g^–1) with only neglectible fluctuations in the heat production rate. Juvenile workers exhibited a mean heat production rate of 142 mW·g^–1. The rate of heat production of adult workers is strongly dependent upon the number of bees together in a group. With more than 10 individuals weight-specific heat dissipation remains constant with increasing group sizes at a level approximately 1/17 that of an isolated bee. Differences are seen between the rates of virgin (117 mW·g^–1) and laying (102 mW·g^–1) queens. Laying queens showed less thermal fluctuations than virgin queens. High fluctuations in heat production rates are observed for drones. In both groups (fertile, juvenile) phases of high and extremely low activity succeed one another. The heat production of juvenile drones was 68 mW·g^–1, that of fertile drones 184 mW·g^–1 due to stronger locomotory activities.     

Daily oxygen consumption rhythms of Senegalese sole solea senegalensis (Kaup, 1858) juveniles

Senegalese sole is known to be a species with pronounced nocturnal feeding behaviour. However, as for most fish species, there is a lack of knowledge concerning the influence of such biological rhythm on metabolic rate. The aim of this study was to determine whether individual variation in routine and fed metabolic rate was affected by daily light-dark rhythms in juveniles of Senegalese sole. The individual oxygen consumption measurements in Senegalese sole juveniles were determined by flow-through respirometry, at fasted conditions and after the fish were fed a single meal, the meal time started at 0930 h and fish fed ad libitum for 30 min. The measurements were made during 22 h, of which 8 h was in the light and 14 h in the dark, and started immediately after transfer to the respiratory chambers at 1100 h. The results suggest an influence of light-dark cycles in routine metabolism. It was observed that oxygen consumption increased during the dark phase in fasted fish (FAST) but was higher during the light phase in fed fish (FEED). However, when feed is provided during the light phase, juveniles are capable of shifting oxygen consumption rhythms to respond to the energetic demands of digestion and growth. These results suggest that routine metabolism varies according to the species natural habits as Senegalese sole is known to be nocturnal. The findings of this study underline the importance of understanding the biological rhythms of the species under study before metabolic data are interpreted.     

Endogenous angiotensin II and the regulation of oxygen consumption and colonic temperature in rats

Previous studies have suggested that angiotensin II, a hormone known to regulate water and salt balance and blood pressure, may also regulate oxygen consumption and body temperature. In this study we investigated the role of endogenous angiotensin in the regulation of oxygen consumption and colonic temperature in rats under low (control) and high (water deprivation, administration of isoproterenol and hemorrhage) peripheral angiotensin conditions. Peripheral administration of losartan, an AT₁ receptor antagonist or enalapril, an angiotensin converting enzyme inhibitor, did not alter oxygen consumption or colonic temperature in control or water deprived rats. Peripheral administration of losartan did not alter the oxygen consumption and colonic temperature responses to the administration of isoproterenol or hemorrhage. Endogenous peripherally generated angiotensin II does not play an important role in regulating either oxygen consumption or colonic temperature in rats under either low or high angiotensin II levels. The reductions in oxygen consumption and colonic temperature that accompany hemorrhage in rats are not mediated by angiotensin II.     

Circadian rhythms of oviposition and feeding activity in Japanese quail: effects of cyclic administration of melatonin

The aim of these experiments was to test the effect of a cyclic administration of melatonin, by mimicking the daily rhythm of hormone levels, on the circadian organization of two distinct functions in quail: oviposition and feeding activity. Laying and feeding rhythms under photoperiodic conditions and constant darkness (DD) were investigated. Under DD, where the two rhythms were free running, a daily rhythm of melatonin was administered. In LD 14h:10h, two different individual profiles of laying were established, with stable females laying at the same time each day and delayed females laying progressively later each day. For feeding activity, all birds were clearly synchronized to the photoperiodic cycle. In DD, the laying birds showed a free-running rhythm of oviposition with a period longer than 24 h for both profiles but the delayed profile females had a longer period than stable profile females. In comparison, the free-running period of feeding rhythm of the same birds was shorter than 24 h. A cyclic administration of melatonin had no effect on laying rhythm, which continued to free-run in DD, whereas feeding activity was synchronized as soon as the first cycle of melatonin was administered. From these results, it seems that two different circadian systems drive each of the two types of behavior separately. Melatonin could be the main synchronizer for the temporal control of feeding behavior, but it does not play a part in the control of oviposition in Japanese quail.     

Endogenous 24-hour cycle of core temperature and oxygen consumption in week-old Zucker rat pups

Summary Experiments were designed to test whether or not the 24-h core temperature fluctuations in week-old rat pups are of endogenous origin. Lean (Fa/-) Zucker rat pups born on the same day to mothers maintained in two different colonies with light/dark cycles 12 h out of phase with each other were mother-reared through the first 3–4 days of life and then artificially reared simultaneously in constant dim light. Continuous, automatic measurement of core temperature and oxygen consumption during artificial rearing showed clear 24-h rhythms in 5- to 8-day-old pups. Each rhythm reached a daily minimum at a time corresponding to the beginning of the light period in the colony of origin. The amplitude of these rhythms did not diminish during artificial rearing, nor did the phase difference between the rhythms of pups originating in the two colonies systematically change. The persistent 12-h phase differences between these two groups of pups prove that the observed rhythms are not caused by exogenous stimuli. We conclude that the rat pup possesses an endogenous time-keeping mechanism that permits the expression of overt rhythmicity at the age of 1 week.Abbreviations Tc core temperature - LD-pups born to a mother entrained to a 7:00 to 19:00 light cycle, then artificially reared in constant dim light - DL-pups born to a mother entrained to a 19:00 to 7:00 light cycle, then artificially reared in constant dim light - SCN suprachiasmatic nucleus     

Daily rhythm of oxygen consumption and thermoregulatory responses in some European winter- or summer-acclimatized finches at different ambient temperatures

The oxygen consumption of European finches, the siskin (Carduelis spinus), the brambling (Fringilla montifringilla), the bullfinch (Pyrhulla pyrhulla), the greenfinch (Carduelis chloris) and the hawfinch (Coccothraustes coccothraustes), was recorded continuously while ambient temperature was decreased stepwise from +30 down to-75°C. The oxygen consumption, body temperature (telemetrically), and shivering (integrated pectoral electromyography) of greenfinches were measured simultaneously at ambient temperatures between +30 and-75°C. Maximum heat production, cold limit, lower critical temperature, basal metabolic rate and thermal conductance (of the greenfinch) were determined. The diurnal variation of oxygen consumption of siskins and greenfinches was recorded at thermoneutrality and below the thermoneutral zone in winter- and summer-acclimatized birds. The diurnal variation of body temperature and thermal conductance of greenfinches were also determined. The diurnal variation of heat production was not seasonal or temperature dependent in the siskin and in the greenfinch. Nocturnal reduction of oxygen consumption saved 15–33% energy in the siskin and greenfinch. Body temperature of the greenfinch was lowered by 2.5–3.4°C. The nocturnal reduction of thermal conductance in the greenfinch was 39–48%. The basal metabolic rate was lowest in the largest bird (hawfinch) and highest in the smallest bird (siskin). The values were in the expected range. The heat production capacity of finches in winter was 4.7 times basal metabolic rate in the siskin, 4.2 times in the brambling, 3.5 times in the greenfinch and 2.9 times in the bullfinch and hawfinch. The heat production capacity of the siskin and greenfinch was not significantly lower in summer. The cold limit temperatures (°C) in winter were-61.2 in the siskin,-41.3 in the greenfinch,-37.0 in the bullfinch,-35.7 in the brambling and-28.9 in the hawfinch. The cold limit was 14.3°C higher in summer than in winter in the siskin and 8.7°C in the greenfinch. Thermal insulation of the greenfinch was significantly better in winter than in summer. The shivering of the greenfinch increased linearly when ambient temperature was decreased down to-40°C. Maintenance of shivering was coincident with season. In severe cold integrated pectoral electromyography did not correlate with oxygen consumption as expected. The possible existence of non-shivering thermogenesis in birds is discussed. It is concluded that the acclimatization of European finches is primarily metabolic and only secondly affected by insulation.Abbreviations AAT avian adipose tissue - bm body mass - BMR basal metabolic rate - C _t thermal conductance - EMG electromyogram - HP heat production - HP _max maximum heat production - MR metabolic rate - NST non-shivering thermogenesis - RMR resting metabolic rate - RQ respiratory quotient - T _a ambient temperature - T _b body temperature - T _c colonic temperature - T _1c lower critical temperature - TNZ thermoneutral zone - T _st shivering threshold temperature - V oxygen consumption     

Regulation of melatonin production by light,darkness, and temperature in the trout pineal

Summary The pineal gland of the rainbow trout, Salmo gairdneri, when kept under in vitro perifusion culture conditions, displays a consistently elevated level of melatonin production in darkness (Gern and Greenhouse 1988). Upon light exposure melatonin production falls and stabilizes at a new lower level that is dependent upon the irradiance of the stimulus. To achieve the maximal response for each irradiance, the duration of the stimulus must exceed 30 min. The response amplitude is maximally sensitive to photons presented over durations of 30–45 min; is very insensitive to shorter light exposures; and is maintained with no evidence of adaptation over longer exposures. Temperature plays a role in regulation of melatonin production both in darkness and during light exposure; increased temperature increases melatonin production in darkness and also increases the sensitivity of the response to light. The action spectrum for the response is best fit by the Dartnall nomogram for a vitamin A₁ based rhodopsin with peak sensitivity near 500 nm. The possible adaptive significance of control of melatonin synthesis by light and temperature is considered.Abbreviations LD lightdark cycle - RIA radioimmunoassay - I ¹²⁵ Iodine - HIOMT hydroxyindole-O-methyltransferase     

Comparative study of the activity/rest rhythms in young and old ringdove (Streptopelia risoria): correlation with serum levels of melatonin and serotonin

Aging is characterized by changes in the circadian rhythms of melatonin, serotonin, and sleep/wakefulness, alterations that affect sleep quality. The authors studied the circadian rhythms of serotonin and melatonin in young and old ringdoves (Streptopelia risoria) (2-3 and 10-12 yrs old, respectively), animals that are characterized by being monophasic and active by day, like humans. The aim was to correlate the indole rhythms with the animals' activity/rest periods. The animals were kept under a 12:12 h light/dark cycle, fed ad libitum, and housed in separate cages equipped for activity recording. Activity pulses were recorded with one actometer per animal (two perpendicular infrared transmitters) and were logged every 15 min by a computer program (DAS 16) throughout the experiment. Melatonin was measured by radioimmunoassay and serotonin by ELISA at intervals of 3 h (from 09:00 to 18:00 h) and 1 h (from 21:00 to 06:00 h), respectively. The results showed a reduction in nocturnal vs. diurnal activity of 89% and 61% in the young and old animals, respectively, with 100% considered to be the diurnal activity of each group. The amplitude of a cosine function fit to the melatonin concentrations of the old animals was half that of the young birds. The acrophase and nadir were at 02:00 and 14:00 h in the young and 01:00 and 13:00 h in the old animals, respectively. The amplitude of the corresponding cosine function fit to the serotonin concentrations in the old birds was one-third that of the young animals. The acrophase and nadir were at 15:00 and 03:00 h in the young and 16:00 and 04:00 h in the old animals, respectively. For both melatonin and serotonin, the concentrations in the young animals were significantly higher than in the old at most of the measurement times. There was a clear negative correlation between the circadian rhythms of activity and the serum melatonin levels in both young and old animals. The equivalent correlation for serotonin was positive, and stronger in the case of the young animals. The results suggest a possible relationship between the observed decline in the amplitude of the old animals' melatonin and serotonin rhythms and the lower percentage reduction in their nocturnal relative to diurnal activity pulses compared to the young animals. In conclusion, the circadian rhythms of melatonin and serotonin undergo alterations with age that could be involved in the changes in age-associated sleep.