Development of hamster circadian rhythms. I. Within-litter synchrony of mother and pup activity rhythms at weaning

The circadian wheel-running activity rhythms of individual hamster pups raised and maintained in constant dim light were measured beginning at 18 days of age. Records of the postweaning free-running activity rhythm were used to determine the phase of a pup's rhythm on the day of weaning and its phase relationship to its mother's rhythm. Although raised in constant light, the rhythms of pups within a litter were approximately synchronous and in phase with their mother's activity rhythm. These results indicate that the circadian oscillator underlying the activity rhythm is functional prior to weaning and is entrained by some as yet unidentified aspect of maternal rhythmicity. Furthermore, the results suggest that even in the absence of external entraining cycles, behavioral rhythms, and perhaps physiologic rhythms as well, of a mother and her offspring are normally synchronized.     

Development of hamster circadian rhythms: Role of the maternal suprachiasmatic nucleus

Summary During development, the circadian rhythms of rodents become entrained to rhythmicity of the mother. Rhythms in behavior and in neuroendocrine function are regulated by a circadian pacemaker thought to be located within the suprachiasmatic nucleus (SCN) of the hypothalamus. Evidence indicates that this pacemaker begins to function and to be entrained by maternal rhythms before birth. Although the maternal rhythms which mediate prenatal entrainment of the fetal circadian pacemaker have not been identified, it is likely that they are regulated by the maternal SCN.The role of the maternal SCN in entrainment of the offspring was examined in Syrian hamsters (Mesocricetus auratus) by measuring the activity/rest rhythms of pups. Using the synchrony among the rhythms of pups within a litter as an indication that the pups had been entrained, the effect on entrainment of ablating the maternal SCN was determined. Lesions of the maternal SCN which were performed early in gestation (day 7) and which destroyed at least 75% of the SCN were found to disrupt the normal within litter synchrony among pups, indicating interference with the normal mechanism of entrainment.The effect of lesions on day 7 of gestation could mean that the maternal SCN is important for entrainment of the pups before birth, after birth, or during both of these times. To determine if the maternal SCN is specifically important for prenatal entrainment, lesions were performed two days before birth on day 14 of gestation. Lesions of the maternal SCN on day 14 were not as disruptive as were lesions on day 7. This suggests that the maternal SCN is important between days 7 and 14 of gestation and that the synchrony normally observed at weaning is already established, in part, on or before day 14 of gestation. This further suggests that an entrainable circadian pacemaker is present in the fetus only two weeks after fertilization.Abbreviations SCN suprachiasmatic nucleus - L:D light:dark - LL constant light - r mean vector length - 2DG 2-deoxyglucose - NAT N-acetyltransferase     

Maternal entrainment of tau mutant hamsters.

Maternal entrainment of the circadian wheel-running activity rhythm was examined in Syrian hamsters heterozygous for a single gene mutation (tau) that affects the free-running period of circadian rhythms. Heterozygous tau pups were born to and raised by wild-type mothers under constant dim light. The pups' wheel-running activity was recorded after weaning on postnatal day 18 or 24. Pups weaned on day 18 had an average free-running period of 21.70 hr, demonstrating that the tau phenotype was fully expressed at this age. Using the activity onset of the postnatal free-running rhythms as a phase reference, we estimated the phase relationships between the pups and their mothers on days 18 and 24. In contrast to results with wild-type pups, the activity rhythms of tau pups were not in phase with the rhythms of their wild-type mothers; that is, activity onsets of mothers and pups did not coincide. The pups did, however, show synchrony among themselves, indicating that they had been exposed to a synchronizing signal sometime during development. It is likely that this synchronizing signal was provided by the mothers, since pups from different litters showed phase relationships similar to those of their mothers. Thus the mothers provided a signal that was sufficient to cause entrainment, despite the 2-hr difference in free-running period between the mothers and pups. Although the pups' activity rhythms appeared to have been entrained by the mothers, they were clearly free-running by postnatal day 18. The mechanism for entrainment is lost during the course of development, despite continued interaction between the mothers and pups.     

Development of hamster circadian rhythms: prenatal entrainment of the pacemaker

Individual hamster pups were maintained in constant dim light from just prior to birth, and their circadian wheel-running activity rhythms were recorded beginning at 18 days of age. Records of the postweaning free-running activity rhythm were used to determine the phase of a pup's rhythm on the day of weaning. Two groups of pups (LD and DL) were born to mothers that had been entrained before birth to light-dark cycles 12 hr out of phase. Both groups of pups were raised in constant dim light by foster mothers that had been entrained to only one of the prenatal cycles (LD). Thus pups born to mothers from different cycles were exposed to identical rhythmic environments postnatally. Despite the similar postnatal treatment, the two groups of pups showed activity rhythms at weaning with very different phases. The LD pups, born to and raised by LD mothers, were approximately synchronous with one another and with their foster mothers. The DL pups, born to DL mothers, but raised by LD mothers, were not synchronous with one another or with their foster mothers. Half of the DL pups showed phases predicted by their prenatal treatment, but the other half showed scattered phases. The results demonstrate that phase at weaning is affected by prenatal rhythmicity, and suggest that the circadian pacemaker underlying the activity rhythm is already functional and entrained at, or before, birth.     

Circadian and photoperiodic time measurement in male Syrian hamsters following lesions of the melatonin-binding sites of the paraventricular thalamus.

Autoradiographic studies using [125I]iodomelatonin in several species, including the Syrian hamster, have revealed that the rostral region of the anterior paraventricular nucleus of the thalamus (aPVT) contains a very high density of binding sites for melatonin. In two studies, small or large bilateral electrolytic lesions of the aPVT were made in adult male hamsters maintained on long days (LD 16:8). The hamsters were then transferred to short days (LD 8:16) to test whether testicular regression could occur in response to a decrease in photoperiod. Serum prolactin concentrations were measured as a second photoperiodic response. All unoperated control hamsters showed the typical short-day photoperiodic response: A decrease in serum luteinizing hormone (LH) and prolactin concentrations and testicular regression all occurred within 6 weeks in short days, followed by the development of scotorefractoriness. Lesions of the aPVT did not significantly affect the rate or the degree of the short-day-induced decline in serum levels of LH or prolactin, nor the pattern of testicular regression and the subsequent expression of refractoriness. To enable us to determine whether the aPVT might be involved in the entrainment or the expression of circadian rhythms, locomotor activity was monitored continuously in lesioned and control groups in Experiment 2, prior to and following the switch to short days. The reduction in photoperiod (involving an 8-hr advance in the time of lights-off and an 8-hr extension of the dark phase) caused a decompression of the nocturnal activity bout of control animals, so that after 2 weeks in short days, activity onset had also advanced to regain its phase relationship to the timing of lights-off. A similar pattern of reentrainment was observed in lesioned animals, and no differences were observed between treatment groups in the rate of entrainment and decompression. In addition, both intact controls and animals bearing large bilateral lesions of the aPVT exhibited robust free-running circadian rhythms of locomotor activity when held under constant dim red light. In summary, the integrity of the aPVT is not necessary for the seasonal response of the reproductive axis and prolactin secretion to photoperiod, nor for photic entrainment of activity rhythms, in the Syrian hamster.     

Circadian rhythm of locomotor activity in the Japanese honeybee, Apis cerana japonica

Abstract.  To reveal circadian characteristics and entrainment mechanisms in the Japanese honeybee Apis cerana japonica , the locomotor-activity rhythm of foragers is investigated under programmed light and temperature conditions. After entrainment to an LD 12 : 12 h photoperiodic regime, free-running rhythms are released in constant dark (DD) or light (LL) conditions with different free-running periods. Under the LD 12 : 12 h regime, activity offset occurs approximately 0.4 h after lights-off transition, assigned to circadian time (Ct) 12.4 h. The phase of activity onset, peak and offset, and activity duration depends on the photoperiodic regimes. The circadian rhythm can be entrained to a 24-h period by exposure to submultiple cycles of LD 6 : 6 h, as if the locomotive rhythm is entrained to LD 18 : 6 h. Phase shifts of delay and advance are observed when perturbing single light pulses are presented during free-running under DD conditions. Temperature compensation of the free-running period is demonstrated under DD and LL conditions. Steady-state entrainment of the locomotor rhythm is achieved with square-wave temperature cycles of 10 °C amplitude, but a 5 °C amplitude fails to entrain.     

Photoperiod differentially modulates photic and nonphotic phase response curves of hamsters

Circadian pacemakers respond to light pulses with phase adjustments that allow for daily synchronization to 24-h light-dark cycles. In Syrian hamsters, Mesocricetus auratus, light-induced phase shifts are larger after entrainment to short daylengths (e.g., 10 h light:14 h dark) vs. long daylengths (e.g., 14 h light:10 h dark). The present study assessed whether photoperiodic modulation of phase resetting magnitude extends to nonphotic perturbations of the circadian rhythm and, if so, whether the relationship parallels that of photic responses. Male Syrian hamsters, entrained for 31 days to either short or long daylengths, were transferred to novel wheel running cages for 2 h at times spanning the entire circadian cycle. Phase shifts induced by this stimulus varied with the circadian time of exposure, but the amplitude of the resulting phase response curve was not markedly influenced by photoperiod. Previously reported photoperiodic effects on photic phase resetting were verified under the current paradigm using 15-min light pulses. Photoperiodic modulation of phase resetting magnitude is input specific and may reflect alterations in the transmission of photic stimuli.     

Prenatal Entrainment of Rat Testicular Malate Dehydrogenase Activity Circadian Rhythm

In mammals the photoperiodic synchronization of circadian system starts before birth. During fetal and neonatal period mothers relay the photoperiodic information to their litter. The maternal pineal melatonin 24 h cycle acts as a synchronizing signal. We have studied the effect of pineal maternal sympathetic denervation and administration of melatonin to mothers denervated during gestation on the prenatal synchronization of testicular malate dehydrogenase (MDH) activity circadian rhythm of the offspring 25 days after birth. When mothers were denervated at the 7th, 10th or 11th day of gestation, pups showed disruption of testicular MDH activity circadian rhythms. In contrast, no disruptive effect was observed when the mothers were denervated on the 12th or 14th day of gestation. When denervated mothers (7th day of gestation) were treated with a daily dose of melatonin from the 11th to the 14th day of gestation, pups showed a MDH activity circadian rhythm. The hormone failed to impose a daily phase when administered from the 9th to the 12th day of gestation. Results suggest that prenatal synchronization in the rat occurs very early in the development, before suprachiasmatic nuclei morphologic arrangement and functional activity begin.     

Prenatal Entrainment of Rat Testicular Malate Dehydrogenase Activity Circadian Rhythm

In mammals the photoperiodic synchronization of circadian system starts before birth. During fetal and neonatal period mothers relay the photoperiodic information to their litter. The maternal pineal melatonin 24 h cycle acts as a synchronizing signal. We have studied the effect of pineal maternal sympathetic denervation and administration of melatonin to mothers denervated during gestation on the prenatal synchronization of testicular malate dehydrogenase (MDH) activity circadian rhythm of the offspring 25 days after birth. When mothers were denervated at the 7th, 10th or 11th day of gestation, pups showed disruption of testicular MDH activity circadian rhythms. In contrast, no disruptive effect was observed when the mothers were denervated on the 12th or 14th day of gestation. When denervated mothers (7th day of gestation) were treated with a daily dose of melatonin from the 11th to the 14th day of gestation, pups showed a MDH activity circadian rhythm. The hormone failed to impose a daily phase when administered from the 9th to the 12th day of gestation. Results suggest that prenatal synchronization in the rat occurs very early in the development, before suprachiasmatic nuclei morphologic arrangement and functional activity begin.     

Entrainment of 2 subjective nights by daily light:dark:light:dark cycles in 3 rodent species

Recent work with exotic 24-h light:dark:light:dark (LDLD) cycles indicates surprising flexibility in the entrainment patterns of Syrian hamsters. Following exposure to an LDLD cycle, hamsters may adopt a form of rhythm splitting in which markers of subjective night (e.g., activity, melatonin) are expressed in each of the twice daily scotophases. This pattern contrasts markedly with that of conventionally entrained hamsters in which markers of subjective night are expressed once daily in only 1 of the 2 dark periods. The "split" entrainment pattern was examined further here in Syrian and Siberian hamsters and in mice exposed to LDLD 7:5:7:5, a condition that reliably induces split activity rhythms in all 3 species. The phase angle of entrainment and activity duration were generally similar comparing the 2 daily activity bouts in each species. The stability of this split entrainment state was assessed by deletions of photophases on individual days, by exposure to skeleton photoperiods, and by transfer to constant darkness. As in Syrian hamsters, the one-time substitution of darkness for one 7-h photophase did not grossly alter activity patterns of Siberian hamsters but acutely disrupted the split rhythms of mice. Skeleton light pulses of progressively shorter duration did not significantly alter split entrainment patterns of either Syrian or Siberian hamsters. Both species continued to exhibit stable entrainment with activity expressed in alternate scotophases of an LD 1:5 cycle presented 4 times daily. In contrast, the split activity rhythms of mice were not maintained under skeleton pulses. In constant darkness, rhythms of Siberian hamsters remained distinctly split for a minimum of 2 cycles. Split entrainment to these novel LDLD and 4-pulse skeleton lighting regimes demonstrates a marked degree of plasticity common to the circadian systems of several rodent species and identifies novel entrainment patterns that may be reliably elicited with simple environmental manipulations. Inter- and intraspecific differences in the stability of split activity rhythms likely reflect differences in coupling interactions between the component circadian oscillators, which, adopting separate phase relations to these novel LD cycles, yield a split entrainment pattern.     

Parametric and nonparametric entrainment of circadian locomotor rhythm in the Japanese honeybee Apis cerana japonica

The circadian rhythm of locomotor activity in the Japanese honeybee Apis cerana japonica was studied to determine the involvement of parametric and/or nonparametric entrainment. The rhythm was entrained to a skeleton photoperiod in which a 1-h first light pulse was imposed in the morning along with a second light pulse in the evening, as well as to a complete photoperiodic regime (LD 12:12). However, the timing of peak activity relative to the lights-off in the evening in the skeleton photoperiod was earlier than that in the complete photoperiod. A single daily light pulse in the evening entrained the rhythm, whereas a daily light pulse in the morning allowed free-running as in constant darkness. The free-running period (τ) of locomotor activity in constant light became longer as the light intensity increased. A Winfree's type I phase response curve of the locomotor activity rhythm was obtained using a single 1-h light pulse. The results suggest that both parametric and nonparametric entrainment are involved in the circadian rhythm of individual locomotor activity in this honeybee.     

Light-dark entrainment of proestrous LH surges and circadian locomotor activity in female hamsters

The temporal relationships of the proestrous LH surge and the circadian locomotor activity rhythm were compared in hamsters entrained to four different 24-hr light-dark (LD) cycles. Animals were housed in cages equipped with running wheels to obtain continuous activity records. Stable entrainment of locomotor activity was complete within 3 weeks of exposure to each photoperiod at which time hamsters were randomly assigned to hourly sample groups. Serum was obtained by cardiac puncture under light ether anesthesia on the day of proestrous and was assayed by RIA for LH. A computer-based least-squares sine wave-fitting technique determined a single objective phase reference point for the time of the hormone maximum. In each photoperiod, precise temporal relationships were maintained between the LH surge and activity onset, whereas the phase relationship between the LH surge and the LD cycle was more variable. These data indicate that the environmental LD cycle entrains the circadian timing system which, in turn, provides temporal information to the rhythms of proestrous gonadotropin and locomotor activity.     

Exotic photoperiods induce and entrain split circadian activity rhythms in hamsters

The split circadian activity rhythm that emerges in hamsters after prolonged exposure to constant light has been a theoretical cornerstone of a multioscillator view of the mammalian circadian pacemaker. The present study demonstrates a novel method for splitting hamster circadian rhythms and entraining them to exotic light:dark cycles. Male Syrian hamsters previously maintained on a 14-h day and 10-h night were exposed to a second 5-h dark phase in the afternoon. The 10-h night was progressively shortened until animals experienced two 5-h dark phases beginning 10 h apart. Most hamsters responded by splitting their activity rhythms into two components associated with the afternoon and nighttime dark phases, respectively. Each activity component was entrained to this light:dark:light:dark cycle. Transfer of split hamsters to constant darkness resulted in rapid joining of the two activity components with the afternoon component associated with onset of the fused rhythm. In constant light, the nighttime component corresponded to activity onset of the fused rhythm, but splitting emerged again at an interval characteristic for this species. The results place constraints on multi-oscillator models of circadian rhythms and offer opportunities to characterize the properties of constituent circadian oscillators and their interactions.     

Resilience of circadian pacemaker development in hamsters

Disruptions of circadian rhythms have been linked to a wide range of pathologies from sleep disorders to cancer. The extent to which disruptions of circadian rhythms during development contribute to later conditions is not known. The present study tested the hypothesis that functional properties of the central circadian pacemaker, the suprachiasmatic nucleus (SCN), are affected by abnormal entrainment during development. The SCN is specialized for the generation of robust rhythms, for direct and indirect output to physiological and behavioral systems, and for entrainment to light/dark cycles via direct retinal input. It consists of thousands of neurons and glia with distinct phenotypes and has subdivisions delineated by both anatomical and functional criteria. In rodents, SCN rhythms develop within days after SCN cells are produced and before many other aspects of differentiation, such as synaptogenesis, are complete. We demonstrated that around the time of birth, the hamster SCN in vivo can undergo repeated phase shifts by a dopamine D(1) receptor agonist (SKF-38393). For 2 days before and 2 days after birth, one group of hamsters received regular exposure to the drug at the same time of day, while another group was exposed at varying times to induce repeated phase shifts. Free-running and entrained activity rhythms were compared between the groups at different ages after weaning. Repeated phase shifts during SCN development had a significant effect on free-running period measured immediately after weaning. This effect was eliminated by subsequent entrainment to a light/dark cycle, indicating that the effect was not permanent. These and other results suggest that SCN development required for functional properties such as free-running period is resilient to perturbation.     

Pineal-independent regulation of photo-nonresponsiveness in the Siberian hamster (Phodopus sungorus)

The pineal hormone melatonin influences circadian rhythms and also mediates reproductive responses to photoperiod. The authors tested whether pinealectomy influences circadian oscillators responsible for induction of nonresponsiveness to short day lengths by preventing normal short-day patterns of circadian entrainment. Adult male Siberian hamsters were pinealectomized or sham operated, maintained in either 18 h light per day (18L) or 15L for 10 weeks, and then tested for responsiveness to 10L. Because pinealectomized hamsters do not show gonadal regression in short day lengths, responsiveness was assessed by measuring phase angle of entrainment and the length of the nightly activity period following transfer to 10L. The incidence of nonresponsiveness was significantly higher in 18L hamsters than in 15L hamsters but was unaffected by pineal status. Fully 88% of 18L hamsters failed to entrain to 10L in the normal short-day manner; the duration of nightly activity remained compressed, and the phase angle of entrainment was large and negative relative to lights off. The 15L hamsters entrained normally to 10L. Exposure to constant light after 10L treatment was equally effective in inducing arrhythmicity in pinealectomized and intact hamsters. Changes in the period of morning and evening circadian oscillators subsequent to 18L treatment did not predict circadian responsiveness to short photoperiod. Long-day induction of photo-nonresponsiveness, which prevents winter responses to short day lengths, occurs independently of pineal melatonin feedback on the circadian system.     

Suppression of circadian rhythms of pineal and testicular hormones following lithium treatment in normal and reversed light-dark cycles, constant light and constant dark in rats

The aim of the current investigation was to study the effect of lithium on circadian rhythms of pineal - testicular hormones by quantitations of pineal and serum serotonin, N-acetylserotonin and melatonin, and serum testosterone at four time points (06.00, 12.00, 18.00 and 24.00) of a 24-hr period under normal photoperiod (L:D), reversed photoperiod (D:L), constant light (L:L) and constant dark phase (D:D) in rats. Circadian rhythms were observed in pineal hormones in all the combinations of photoperiodic regimens, except in constant light, and in testosterone levels in all the photoperiodic combinations. Pineal and serum N-acetylserotonin and melatonin levels were higher than serotonin at night (24.00 hr), in natural L:D cycle, in reversed L:D cycle or similar to normal L:D cycle in constant dark phase, without any change in constant light. In contrast, testosterone level was higher in light phase (12.00 hr through 18.00 hr) than in the dark phase (24.00 hr through 06.00 hr) in normal L:D cycle, in reversed L:D cycle, similar to normal L:D cycle in constant dark (D:D), and reversed to that of the normal L:D cycle in constant light (L:L). Lithium treatment (2 mEq/kg body weight daily for 15 days) suppressed the magnitude of circadian rhythms of pineal and serum serotonin, N-acetylserotonin and melatonin, and testosterone levels by decreasing their levels at four time points of a 24-hr period in natural L:D or reversed D:L cycle and in constant dark (D:D). Pineal indoleamine levels were reduced after lithium treatment even in constant light (L:L). Moreover, lithium abolished the melatonin rhythms in rats exposed to normal (L:D) and reversed L:D (D:L) cycles, and sustained the rhythms in constant dark. But testosterone rhythm was abolished after lithium treatment in normal (L:D)/reversed L:D (D:L) cycle or even in constant light/dark. The findings indicate that the circadian rhythm exists in pineal hormones in alternate light - dark cycle (L:D/D:L) and in constant dark (D:D), but was absent in constant light phase (L:L) in rats. Lithium not only suppresses the circadian rhythms of pineal hormones, but abolishes the pineal melatonin rhythm only in alternate light - dark cycles, but sustains it in constant dark. The testosterone rhythm is abolished after lithium treatment in alternate light - dark cycle and constant light/dark. It is suggested that (a) normal circadian rhythms of pineal hormones are regulated by pulse dark phase in normal rats, (b) lithium abolishes pineal hormonal rhythm only in pulse light but sustains it in constant dark phase, and (c) circadian testosterone rhythm occurs in both pulse light or pulse dark phase in normal rats, and lithium abolishes the rhythm in all the combinations of the photoperiod. The differential responses of circadian rhythms of pineal and testicular hormones to pulse light or pulse dark in normal and lithium recipients are discussed.     

Entrainment of split circadian activity rhythms in hamsters

Hamsters that showed splitting of their circadian rhythms of wheel-running activity following long-term exposure to constant illumination (LL) were exposed to light-dark (LD) cycles with 2-hr dark segments, and with periods of 24.00, 24.23 or 24.72 hr. For comparison, hamsters showing nonsplit rhythms were also studied. In all cases of split rhythms, at least one of the two split components entrained to the LD cycles. In some animals, the second component continued to free-run until it merged with the entrained component, while in others, the second component also entrained to the LD cycle but maintained a stable phase angle of 6-14.5 hr relative to dark onset. These results were obtained in cases where the period of the LD cycle was shorter than that of the split rhythms and in cases where it was longer, implying that split components can be phase-advanced as well as phase-delayed by 2 hr of darkness. Three hamsters that showed stable entrainment of split rhythms were allowed to free-run in LL. The LD cycles were then reinstated, but instead of overlapping with the first component, as it did before, the dark segment was timed to overlap with the second. The entrainment patterns that ensued were similar to the ones obtained during the first LD exposure, indicating that the two split components respond to darkness in a qualitatively similar fashion. These results are further evidence that the pacemaker system underlying split circadian activity rhythms in hamsters is composed of two mutually coupled populations of oscillators that have similar properties, including a bidirectional phase response curve. Such a dual-oscillator organization may also underlie normal, or nonsplit, activity rhythms, as suggested by Pittendrigh and Daan (1976c), but the data are also compatible with the alternative view that the circadian pacemaker consists of a large number of coupled oscillators, which only dissociate into two separate populations in some animals under conditions of moderate LL intensity.     

Light-Dark Entrainment of Circadian Activity Rhythms of the Diurnal Indian Palm Squirrel (Funambulus pennanti)

A recent focus of chronobiological studies has been to establish diurnal models as alternatives to the more frequently used nocturnal rodents. In the present study, light-dark (LD) entrainment characteristics were examined in one diurnal species, the Indian palm squirrel (Funambulus pennanti). Palm squirrels showed strongly diurnal locomotor activity rhythms (? 88 percent) under light-dark (LD) cycles, with activity bimodally distributed during the L phase. In comparison to a dim LD cycle, exposure to a bright LD cycle caused a phase advance in the onset of activity, an increase in daily activity levels and an increase in the duration of activity. Percentage diurnality, however, did not vary between bright and dim LD cycles. Activity rhythms reentrained in significantly fewer days after an 8 hour phase delay of the LD cycle compared to an 8 hour phase advance. In both cases, the direction of reentrainment followed the direction of the LD shift. When exposed to single light pulses (1 hour) presented at the same time each day, 6/7 squirrels entrained. Under a skeletal photoperiod cycle (2 x 1 hour light pulses each day), 6/8 squirrels showed stable entrainment. The remaining squirrels exhibited rhythm splitting, with each component synchronising in an unstable manner with one of the light pulses. Under entrainment to single light pulses and to the skeletal photoperiod cycle, the phase angle of entrainment was negatively correlated with t. Finally, when exposed to a skeletal scotoperiod cycle (2 x 1-hour dark pulses each day), only 3/8 squirrels entrained, while the others free-ran. Two of the entrained squirrels showed spontaneous phase reversals during entrainment. As with other species, the activity rhythm of palm squirrels appears to be controlled by two separate self-sustaining oscillators. The strongly diurnal nature of palm squirrels make them a promising diurnal model for studies examining endogenous and exogenous influences on circadian functioning.     

Light-Dark Entrainment of Circadian Activity Rhythms of the Diurnal Indian Palm Squirrel (Funambulus pennanti)

A recent focus of chronobiological studies has been to establish diurnal models as alternatives to the more frequently used nocturnal rodents. In the present study, light-dark (LD) entrainment characteristics were examined in one diurnal species, the Indian palm squirrel ( Funambulus pennanti ). Palm squirrels showed strongly diurnal locomotor activity rhythms (~ 88 percent) under light-dark (LD) cycles, with activity bimodally distributed during the L phase. In comparison to a dim LD cycle, exposure to a bright LD cycle caused a phase advance in the onset of activity, an increase in daily activity levels and an increase in the duration of activity. Percentage diurnality, however, did not vary between bright and dim LD cycles. Activity rhythms reentrained in significantly fewer days after an 8 hour phase delay of the LD cycle compared to an 8 hour phase advance. In both cases, the direction of reentrainment followed the direction of the LD shift. When exposed to single light pulses (1 hour) presented at the same time each day, 6/7 squirrels entrained. Under a skeletal photoperiod cycle (2 x 1 hour light pulses each day), 6/8 squirrels showed stable entrainment. The remaining squirrels exhibited rhythm splitting, with each component synchronising in an unstable manner with one of the light pulses. Under entrainment to single light pulses and to the skeletal photoperiod cycle, the phase angle of entrainment was negatively correlated with t. Finally, when exposed to a skeletal scotoperiod cycle (2 x 1-hour dark pulses each day), only 3/8 squirrels entrained, while the others free-ran. Two of the entrained squirrels showed spontaneous phase reversals during entrainment. As with other species, the activity rhythm of palm squirrels appears to be controlled by two separate self-sustaining oscillators. The strongly diurnal nature of palm squirrels make them a promising diurnal model for studies examining endogenous and exogenous influences on circadian functioning.     

Freerunning and entrained circadian rhythms in activity,eating and drinking in the cat

Abstract

Because cats with pontile lesions exhibit an abnormal behavior that is under photoperiodic control, and because circadian rhythms are implicated in photoperiodic control mechanisms, an effort was made to detect circadian rhythms in the cat. Cats were isolated from all extraneous stimuli in soundproof chambers for extended periods of time. Photocells were used to monitor activity, eating and drinking in different LD cycles, in constant light at two intensities, and in constant dark. Freerunning circadian rhythms were found in the constant conditions, and entrained nocturnal patterns occurred in most of the LD cycles. The higher intensities of ambient illumination disrupted the freerunning rhythms. The freerunning rhythms were always greater than 24 h, ranging from 24.2 to 25 h. Measurements of food intake of cats living in a large colony room, obtained by weighing the food, revealed that a nocturnal pattern of entrainment was not present in the majority of the cats. Instead, most cats in the colony exhibited a random pattern of eating throughout the light and dark period of the LD cycle. However, the variation among the cats in the colony was considerable, extending from nocturnal to diurnal patterns of eating. A diurnal pattern of human activity was present in the colony and may account for the disruption of a basic nocturnal pattern. The presence of circadian rhythms in the cat leads us to consider the coincidence models for photoperiodic induction as possible explanations of the photoperiodic control of the lesion‐induced abnormal behavior.