Genetic analyses of photoresponsiveness in the Djungarian hamster,Phodopus sungorus

Summary Endotherms living at temperate and arctic latitudes must adjust their physiology and behavior in order to survive seasonal change. The Djungarian hamster uses photoperiod to cue annual cycles of reproduction and thermoregulation, and its responses to short photoperiod include loss of body weight and change in pelage color. Some individuals do not exhibit these responses when exposed to short days.In this study individual variation in photoresponsiveness is quantified, and four lines of evidence for a genetic component to that variation are provided. First, two separate breeding stocks differed in both the percent of animals responding to a short-day lighting regimen (SD) and in the degree and timing of their response. Second, analysis of variance within and between families of full sibs for a photoresponsive index, PI (body weightloss +2 (molt index –1)) following 12 weeks in SD demonstrated a significant family resemblance (intraclass correlation of 0.36±0.03). Third, heritability estimates from regression of offspring scores on parent scores for body weight loss, molt index and PI after 12 weeks in SD were 0.340.13, 0.36±0.10 and 0.37±0.12, respectively, indicating a strong additive genetic component for the three characters. Finally, a significant response occurred after one generation of artificial selection for and against photoresponsiveness.Abbreviations PI photoresponsive index (body weight loss+2 (molt index –1)) - SD short day light regimen     

Ketamine-xylazine anaesthesia in the Djungarian hamster (Phodopus sungorus)

The combination of ketamine-xylazine was assessed as a surgical anaesthetic in Djungarian hamsters acclimatized to both long (16 h light: 8 h dark) and short (8 h light: 16 h dark) photoperiods. It was concluded that 50 mg/kg of ketamine with 10 mg/kg of xylazine or 100 mg/kg of ketamine with 5-10 mg/kg of xylazine when given together by intraperitoneal injection was a satisfactory general anaesthetic. Two hundred mg/kg of ketamine with 10 mg/kg xylazine caused death in 13 of 24 animals. There were no clinically significant effects on depth of anaesthesia due to photoperiod.     

Circadian rhythms of pineal N-acetyltransferase activity in the Djungarian hamster,Phodopus sungorus,in response to seasonal changes of natural photoperiod

Summary The aim of this study was to describe the regular annual pattern of the daily melatonin synthesis in Djungarian hamsters,Phodopus sungorus sungorus. The hamsters were maintained from birth in natural photoperiodic conditions and in bimonthly intervals the day/night rhythms of pineal N-acetyltransferase (NAT) were measured. Analysis of the circadian profiles of NAT activity showed that the duration of elevated melatonin synthesis closely reflects the duration of the scotophase throughout the seasons. Thus the duration of elevated melatonin seems to represent a direct humoral signal transmitting the photoperiodic message. The duration of the nightly melatonin pulse appears to be influenced mainly by the time of dawn rather than by the time of dusk. Additional information about the time of year might be encoded in the total amount of melatonin synthesized per day, whereas the amplitude of the nightly melatonin peak seems to be of minor importance.Abbreviation NAT N-acetyltransferase Dedicated to Dr. Klaus Hoffmann on the occasion of his 60th birthday     

Chromosome replication patterns in the Djungarian hamster (Phodopus sungorus)

Patterns of early and late replication in the individual chromosomes of the Djungarian hamster (Phodopus sungorus) have been studied using the techniques of Giemsa staining suppression when bromodeoxyuridine is incorporated into the DNA. — Late replicating autosome regions correspond to G-band regions, early replication regions are less clearly demarcated but correspond to R-band regions plus some G-band zones. In part this reduction in sharpness of early replication bands may be due to the fact that nearly all metaphase G-bands contain R-band material since they are compounded from blocks of sub-G bands. — The long arm of the X chromosomes in the female differ in the start time of synthesis but are rarely separable at the close of S. There are no differences between the short arms. In the male, Y starts very late but finishes about the same time as the X which behaves like the early replicating X of the female.Visiting worker from Department of Biological Sciences, Sambulpur University, Burla 768017, India     

Modulation of leptin sensitivity by short photoperiod acclimation in the Djungarian hamster, Phodopus sungorus

During seasonal acclimation, Djungarian hamsters spontaneously exhibit a reduction in food intake, body mass and body fat stores, which is externally cued by shortening of day length in autumn and controlled by a sliding set-point. We investigated the function of the leptin adipostatic feedback system in the photoperiodic control of seasonal acclimation. In response to mouse recombinant leptin injections for 10 days, long day photoperiod (LD) and short day photoperiod (SD)-acclimated hamsters decreased food intake and body mass. The reduction of body mass was due to the depletion of body fat, as revealed by carcass composition analysis. In SD hamsters, leptin caused a larger reduction of body fat mass than observed under LD conditions, whereas the anorectic effect was similar in both photoperiods. The serum leptin concentration was 9.3 ± 1.2 ng/ml in LD-acclimated hamsters and decreased significantly to 4.2 ± 0.8 ng/ml and 2.1 ± 0.6 ng/ml in hamsters exposed to SD for 66 days and 116 days, respectively (P < 0.001). A strong positive correlation between total body fat mass and serum leptin concentration was found (r _S=0.935, P < 0.0001, n=70). Despite the anorectic action of exogenous leptin, higher endogenous leptin levels in LD hamsters were paralleled by higher food intake in LD hamsters as compared to SD hamsters. This paradoxical finding further supports the increased leptin sensitivity in SD hamsters as judged from leptin treatment experiments. We tested the functional significance of leptin for the controlled down-regulation of food intake and body mass induced by short photoperiod. Food restriction for 10 days during the transition phase decreased body mass below the desired sliding set-point, which was recovered in control hamsters following ad libitum refeeding. Treatment with mouse recombinant leptin during ad libitum refeeding inhibited the recovery of body mass and blunted the increase of food intake observed in controls, indicating that the sliding set-point utilizes leptin as a signal for the adjustment of the appropriate body mass level. Accepted: 15 October 1999     

Photoperiod-dependent changes in exocytotic activity in the hypophyseal pars tuberalis of the Djungarian hamster,Phodopus sungorus

The pars tuberalis of the hypophysis of the Djungarian hamster, Phodopus sungorus, was investigated with regard to secretory activity by applying the tannic acid-Ringer perfusion technique. Two groups were maintained under long photoperiods (16 h light: 8 h dark) or short photoperiods (8 h light: 16 h dark), respectively. Perfusion with tannic acid showed that specific pars tuberalis cells release some of their secretory granules as indicated by typical exocytotic figures. The percentage of cells displaying exocytotic activity was significantly higher in the pars tuberalis of hamsters kept under long photoperiods. The number of exocytotic figures per single cell was not increased. These results provide further evidence for a secretory activity of the pars tuberalis and support the hypothesis of its involvement as a mediator between photoperiodic stimuli and the endocrine system.     

Long photoperiod restores the 24-h rhythm of sleep and EEG slow-wave activity in the Djungarian hamster (Phodopus sungorus)

Photoperiod influences the distribution of sleep and waking and electroencephalogram (EEG) power density in the Djungarian hamster. In an experimental procedure combining short photoperiod (SP) and low ambient temperature, the light-dark difference in the amount of sleep was decreased, and the changes in slow-wave activity (SWA) (mean EEG power density between 0.75 and 4.0 Hz) in nonrapid eye movement (NREM) sleep within 24 h were abolished. These findings, obtained in three different groups of animals, suggested that at the lower ambient temperature, the influence of the circadian clock on sleep-wake behavior was diminished. However, it remained unclear whether the changes were due to the photoperiod, ambient temperature, or both. Here, the authors show that EEG and electromyogram recordings in a single group of animals sequentially adapted to a short and long photoperiod (LP) at low ambient temperature (approximately 15 degrees C) confirm that EEG power is reduced in SP. Moreover, the nocturnal sleep-wake behavior and the changes in SWA in NREM sleep over 24 h were restored by returning the animals to LP and retaining ambient temperature at 15 degrees C. Therefore, the effects cannot be attributed to ambient temperature alone but are due to a combined effect of temperature and photoperiod. When the Djungarian hamster adapts to winter conditions, it appears to uncouple sleep regulation from the circadian clock.     

Fenestrated capillaries in the ventral sebaceous gland of the Djungarian hamster, Phodopus sungorus

The ventral sebaceous gland of the Djungarian hamster is a macroscopically visible organ situated in the midventral area of the abdominal wall. It consists of densely packed acini arranged in lobules with common excretory ducts. The rich vascular network of the gland is characterized by fenestrated capillaries. Fenestrated endothelium has not yet been reported as a characteristic and regular finding within sebaceous glands. Results are discussed with regard to proliferation rate of sebocytes and the demand of fluid and nutrient supply.     

Seasonal changes of heart weight and erythrocytes in the Djungarian hamster, Phodopus sungorus

Djungarian dwarf hamsters (Phodopus sungorus) show an annual cycle in weight-specific metabolic rate with a high level during winter. These seasonal changes in oxygen demand are met by hematological adjustments, primarily based on an increased number of erythrocytes, but a decreased erythrocyte volume during winter. Subsequently, the diffusion area for blood gas exchange is increased during this time of high metabolic capabilities. Blood oxygen capacity (hemoglobin, 2,3-diphosphoglycerate (2,3-DPG) does not change with the season. However, seasonal changes in heart weight suggest changes in cardiac output, causing an increased blood flow per unit tissue weight during winter. This increase in circulatory efficiency, as well as changes in erythrocyte surface, are primarily controlled by photoperiod, since it occurred in hamsters living indoors at thermoneutrality but subjected to seasonal changes in photoperiod to the same extent as in hamsters living outdoors.     

Maximum energy assimilation rates in the Djungarian hamster (Phodopus sungorus)

Summary Physiological limits to energy budgets were estimated in Djungarian hamsters (Phodopus sungorus) using food balance and respirometric methods. The summer acclimatized, reproductively inactive hamsters could balance their energy budget at-2° C, assimilating 91.1 kJ·animal^-1· day^-1 after gradual cold acclimation, whereas non-acclimated hamsters showed negative energy balance assimilating only 54.4 kJ·animal^-1·day^-1. At the same ambient temperature, multiparous females (although neither pregnant nor lactating at the time) maintained positive energy balance assimilating 81.6 kJ·animal^-1·day^-1. Hamsters are capable of rapid adjustments of their maximum assimilation rates to meet their current energy demands, but only up to the value of about 3.5xBMR. It is concluded, that the actual energy budgets of small mammals keep, all the time, fairly near the upper physiological limit, with body reserves ready to buffer short-term oscillations.     

Effects of temperature,steroids and castration on daily torpor in the Djungarian hamster (Phodopus sungorus)

1. In Phodopus sungorus the frequency and characteristics of daily torpor were not affected by a decrease in temperature. The seasonal cycle of daily torpor is essentially under photoperiodic control. 2. Testicular regression is a necessary condition for daily torpor occurrence. Testosterone and 5 alpha-dihydrotestosterone administration totally inhibited daily torpor in hamsters which were exposed to short days. The temporal inhibition of torpor by steroids had no effect on the seasonal time measuring mechanism governing daily torpor. 3. Castration influenced certain aspects of the daily torpor display. Castrated hamsters showed a delay in terminating torpor season lending further support to the hypothesis that recrudescence of the testis plays a role in terminating the daily torpor season. Castration, when performed before or on the day of transfer to short photoperiod led to an increase in the frequency of torpor bouts and reduce SP exposure duration which is necessary for the daily torpor season beginning.     

Photoperiod and ambient temperature as environmental cues for seasonal thermogenic adaptation in the Djungarian hamster,Phodopus sungorus

For their seasonal control of thermogenesis Djungarian hamsters rely on environmental cueing by both photoperiod and ambient temperature. Their total potential for adaptive improvements of nonshivering thermogenesis is constant in summer and winter. The shortening of photoperiod in fall is used to anticipate about half of the total improvement in thermogenesis, in advance of any experience of cold, as can be concluded from the photoperiodic control of thermogenesis, cold resistance, and the protein content, cyctochrome oxidase activity and content of mitochondria in brown adipose tissue. The remainder of the seasonal thermogenic adaptation is due to stimulatory responses to chronic exposure to cold.This research was supported by the Deutsche Forschungsgemeinschaft, Schwerpunktprogramm Mechanismen der Temperaturregulation und -Adaptation.     

Cardiac dynamics during daily torpor in the Djungarian hamster (Phodopus sungorus)

Djungarian or Siberian hamsters (Phodopus sungorus) acclimated to short photoperiod display episodes of spontaneous daily torpor with metabolic rate depressed by approximately 70% and body temperature (T(b)) reduced by approximately 20 degrees C. To study the cardiovascular adjustment to daily torpor in Phodopus, electrocardiogram (ECG) and T(b) were continuously recorded by telemetry during entrance into torpor, in deep torpor, and during arousal from torpor. Minimum T(b) during torpor bouts was approximately 21 degrees C, and heart rate, approximately 349 beats/min at euthermy, displayed marked sinus bradyarrhythmia at approximately 70 beats/min. Arousal was typically completed within approximately 40 min, followed by a sustained post-torpor inactivity tachycardia ( approximately 540 beats/min). The absence of episodes of conduction block, tachyarrhythmia, or other forms of ectopy throughout the torpor cycle demonstrates a remarkable resistance to arrhythmogenesis. The ECG morphology lacks a distinct isoelectric interval following the QRS complex, and the ST segment resembles the ECG pattern in mice, with a prominent fast transient outward K(+) current (I(to,f)) determining the early phase of ventricular repolarization. During low-temperature torpor, the amplitudes of the QRS complex substantially increased, suggesting that in the euthermic state the terminal portion of ventricular depolarization is fused with the beginning of repolarization, low T(b) acting to decorrelate the superposition between depolarization and repolarization by delaying the repolarization onset. Atrioventricular and ventricular conduction times were prolonged as function of T(b). In contrast, the QT vs. T(b) relationship showed marked hysteresis indicating the operation of nonlinear control mechanisms whereby the rapid QT shortening during arousal results from additional mechanisms (probably sympathetic stimulation) other than temperature alone.     

Nucleolus-like bodies in the pineal gland of the Djungarian hamster (Phodopus sungorus)

Summary Light- and electron-microscopic observations on the pineal gland of Phodopus sungorus revealed intracytoplasmic inclusions resembling nucleolus-like bodies similar to those found in other regions of the central nervous system. Bernhard's EDTA method was used to confirm that these inclusions were nucleolus-like bodies. These structures were rarely found in pinealocytes of sexually active longday animals, whereas large numbers of them were observed in pinealocytes of sexually quiescent short-day animals. Nucleolus-like bodies may therefore be involved in pineal secretion.     

Cold prevents the light induced inactivation of pineal N-acetyltransferase in the Djungarian hamster,Phodopus sungorus

Summary In the Djungarian hamster seasonal acclimatization is primarily controlled by photoperiod, but exposure to low ambient temperature amplifies the intensity and duration of short day-induced winter adaptations. The aim of this study was to test, whether the pineal gland is involved in integrating both environmental cues. Exposure of hamsters to cold (0 °C) reduces the sensitivity of the pineal gland to light at night and prevents inactivation of N-acetyltransferase (NAT). The parallel time course of NAT activity and plasma norepinephrine content suggests that circulating catecholamines may stimulate melatonin synthesis under cold load.Abbreviations NAT N-acetyltransferase - NE norepinephrine - T _a ambient temperature     

A high incidence of abdominal pregnancy in the Djungarian hamster (Phodopus sungorus).

Eight animals with abdominal fetuses were found in a relatively small colony of Djungarian hamsters. All the fetuses were macerated. Some had remains of an amniotic sac with an umbilical attachment to a placenta adherent to the peritoneum, but others were without placental remains. Gross and histological examinations failed to reveal any ruptures of the uterus or oviducts. It was not possible to decide whether the abdominal pregnancies were primary or secondary.     

The role of ozone/oxygen in clindamycin-associated enterocolitis in the Djungarian hamster (Phodopus sungorus sungorus)

Rectal and oral ozone-O2 gas insufflations prevented a fatal clindamycin-induced enterocolitis in Djungarian dwarf hamsters. In spite of no gross lesions in the colon of these species, the term 'colitis' is justified because of intracellular degenerative changes of colon epithelial cells. Dwarf hamsters are omnivores and are, therefore, more relevant to man than other animal species as an animal model for studying colitis.     

Hormonal regulation of the annual pelage color cycle in the Djungarian hamster, Phodopus sungorus. II. Role of prolactin

This study investigated whether photoperiod-induced changes in circulating prolactin levels, which have been observed in the Djungarian hamster ( Yellon and Goldman, '83; Duncan and Goldman, ' 83a ), might be involved in seasonal pelage color changes in this species. Injection of ovine prolactin (100 micrograms/day) inhibited the short photoperiod-induced winter molt. This finding indicated that the suppression of endogenous prolactin levels normally occurring in short photoperiod-housed hamsters (Duncan and Goldman, ' 83a ) may induce the winter molt. Suppression of prolactin secretion with bromoergocryptine (200 micrograms/day) strongly inhibited the spring molt, while concomitant treatment with ovine prolactin (100 micrograms/day) overcame this effect of bromoergocryptine. Injection of bromoergocryptine (200 micrograms/day) stimulated the winter molt in castrated hamsters housed in long photoperiod; concomitant injection of prolactin (100 micrograms/day) reversed this effect as well. These findings strongly suggested that an increase in endogenous prolactin levels may be necessary for the development and maintenance of the summer pelage.