Total lactate dehydrogenase activity of tail muscle is not cold-adapted in nocturnal lizards from cool-temperate habitats

The dependence of metabolic processes on temperature constrains the behavior, physiology and ecology of many ectothermic animals. The evolution of nocturnality in lizards, especially in temperate regions, requires adaptations for activity at low temperatures when optimal body temperatures are unlikely to be obtained. We examined whether nocturnal lizards have cold-adapted lactate dehydrogenase (LDH). LDH was chosen as a representative metabolic enzyme. We measured LDH activity of tail muscle in six lizard species (n = 123: three nocturnal, two diurnal and one crepuscular) between 5 and 35 °C and found no differences in LDH-specific activity or thermal sensitivity among the species. Similarly, the specific activity and thermal sensitivity of LDH were similar between skinks and geckos. Similar enzyme activities among nocturnal and diurnal lizards indicate that there is no selection of temperature specific LDH enzyme activity at any temperature. As many nocturnal lizards actively thermoregulate during the day, LDH may be adapted for a broad range of temperatures rather than adapted specifically for the low temperatures encountered when the animals are active. The total activity of LDH in tropical and temperate lizards is not cold-adapted. More data are required on biochemical adaptations and whole animal thermal preferences before trends can be established.     

From daily behavior to hormonal and neurotransmitters rhythms: Comparison between diurnal and nocturnal rat species

Mammalian species can be defined as diurnal or nocturnal, depending on the temporal niche during which they are active. Even if general activity occurs during nighttime in nocturnal rodents, there is a patchwork of general activity patterns in diurnal rodents, including frequent bimodality (so-called crepuscular pattern, i.e., dawn and dusk peaks of activity) and a switch to a nocturnal pattern under certain circumstances. This raises the question of whether crepuscular species have a bimodal or diurnal - as opposed to nocturnal - physiology. To this end, we investigated several daily behavioral, hormonal and neurochemical rhythms in the diurnal Sudanian grass rat (Arvicanthis ansorgei) and the nocturnal Long-Evans rat (Rattus norvegicus). Daily rhythms of general activity, wheel-running activity and body temperature, with or without blocked wheel, were diurnal and bimodal for A. ansorgei, and nocturnal and unimodal for Long-Evans rats. Moreover, A. ansorgei and Long-Evans rats exposed to light-dark cycles were respectively more and less active, compared to conditions of constant darkness. In contrast to other diurnal rodents, wheel availability in A. ansorgei did not switch their general activity pattern. Daily, unimodal rhythm of plasma leptin was in phase-opposition between the two rodent species. In the hippocampus, a daily, unimodal rhythm of serotonin in A. ansorgei occurred 7 h earlier than that in Long-Evans rats, whereas a daily, unimodal rhythm of dopamine was unexpectedly concomitant in both species. Multiparameter analysis demonstrates that in spite of bimodal rhythms linked with locomotor activity, A. ansorgei have a diurnally oriented physiology.     

Plasticity of Circadian Activity and Body Temperature Rhythms in Golden Spiny Mice

Most animals can be categorized as nocturnal, diurnal, or crepuscular. However, rhythms can be quite plastic in some species and vary from one individual to another within a species. In the golden spiny mouse (Acomys russatus), a variety of rhythm patterns have been seen, and these patterns can change considerably as animals are transferred from the field into the laboratory. We previously suggested that these animals may have a circadian time‐keeping system that is fundamentally nocturnal and that diurnal patterns seen in their natural habitat reflect mechanisms operating outside of the basic circadian time‐keeping system (i.e., masking). In the current study, we further characterized plasticity evident in the daily rhythms of golden spiny mice by measuring effects of lighting conditions and access to a running wheel on rhythms in general activity (GA) and body temperature (Tb). Before the wheel was introduced, most animals were active mainly during the night, though there was considerable inter‐individual variability and patterns were quite plastic. The introduction of the wheel caused an increase in the level of nighttime activity and Tb in most individuals. The periods of the rhythms in constant darkness (DD) were very similar, and even slightly longer in this study (24.1±0.2 h) than in an earlier one in which animals had not been provided with running wheels. We found no correlation between the distance animals ran in their wheels and the period of their rhythms in DD. Re‐entrainment after phase delays of the LD cycle occurred more rapidly in the presence than absence of the running wheel. The characteristics of the rhythms of golden spiny mice seen in this study may be the product of natural selection favoring plasticity of the circadian system, perhaps reflecting what can happen during an evolutionary transition as animals move from a nocturnal to a diurnal niche.     

Photic and nonphotic inputs to the diurnal circadian clock

Diurnal animals occupy a different temporal niche from nocturnal animals and are consequently exposed to different amounts of light as well as different dangers. Accordingly, some variation exists in the way that diurnal animals synchronize their internal circadian clock to match the external 24-hour daily cycle. First, though the brain mechanisms underlying photic entrainment are very similar among species with different daily activity patterns, there is evidence that diurnal animals are less sensitive to photic stimuli compared to nocturnal animals. Second, stimuli other than light that synchronize rhythms (i.e. nonphotic stimuli) can also entrain and phase shift daily rhythms. Some of the rules that govern nonphotic entrainment in nocturnal animals as well as the brain mechanisms that control nonphotic influences on rhythms do not appear to apply to diurnal animals, however. Some evidence supports the idea that arousal or activity plays an important role in entraining rhythms in diurnal animals, either during the light (active) or dark (inactive) phases, though no consistent pattern is seen. GABAergic stimulation induces phase shifts during the subjective day in both diurnal and nocturnal animals. In diurnal Arvicanthis niloticus (Nile grass rats), SCN GABA_A receptor activation at this time results in phase delays while in nocturnal animals phase advances are induced. It appears that the effect of GABA at this circadian phase results from the inhibition of period gene expression in both diurnal and nocturnal animals. Nonetheless, the resulting phase shifts are in opposite directions. It is not known what stimuli or behaviours ultimately induce changes in GABA activity in the SCN that result in alterations of circadian phase in diurnal grass rats. Taken together, studies such as these suggest that it may be problematic to apply the principles governing nocturnal nonphotic entrainment and its underlying mechanisms to diurnal species including humans.     

Nocturnal lizards from a cool-temperate environment have high metabolic rates at low temperatures

Ectotherms from low-temperature environments have higher metabolic rates at low temperatures than those from warm-temperature environments. We predicted that nocturnal lizards, which are active at much lower environmental temperatures than diurnal lizards, would also have higher metabolic rates at low temperatures, and by association a lower thermal sensitivity (Q ₁₀) than diurnal and crepuscular lizards. We measured the rate of oxygen consumption ( [(V)\dot]\textO ₂ \dot{V}{\text{O}}_{ 2} ) of eight cool-temperate species of lizard (four nocturnal, three diurnal, and one crepuscular) at 13 and 26°C and analyzed log transformations of these data using log mass as a covariate. As expected, [(V)\dot]\textO ₂ \dot{V}{\text{O}}_{ 2} was positively correlated with temperature in all eight species, with [(V)\dot]\textO ₂ \dot{V}{\text{O}}_{ 2} being two to four times higher at 26°C than at 13°C. As predicted, at 13°C (but not 26°C) the [(V)\dot]\textO ₂ \dot{V}{\text{O}}_{ 2} was significantly higher in nocturnal than diurnal lizards. Species-specific differences and mass scaling factors explain the patterns of thermal sensitivity seen among these eight lizard species. Thermal sensitivity is strongly influenced by mass, with smaller species generally having higher thermal sensitivity of their metabolic rate, and this result deserves further exploration among other ectotherms. We conclude that, along with the previously reported lower cost of locomotion found in nocturnal lizards, they also partially offset the thermal handicap of activity at low body temperatures by having an elevated [(V)\dot]\textO ₂ \dot{V}{\text{O}}_{ 2} at lower temperatures.     

A comparative study on insemination dependency of circadian activity pattern in mosquitoes

Abstract. Two forms of mosquitoes in the Culex pipiens complex, C.p.pallens and C.p.molestus , were compared for the insemination-dependency of entrained female circadian activity in LD 16:8h. When virgin, Pallens is nocturnal and molestus is, in addition, diurnal. Insemination and injection of extracts of the male accessory gland affects virgin activity in that nocturnality is strengthened in both mosquitoes, and the diurnal activity of molestus is depressed. This was so whether the treatments were intra- or interspecific. These findings are compared with those already published for other mosquitoes. A coherent view is proposed that the mosquito, nocturnal/crepuscular, nocturnal and diurnal, or diurnal, has potentially a similar entrained waveform which is affected by insemination in a common way, and by lighting conditions in species specific ways.     

RHYTHMIC cFOS EXPRESSION IN THE VENTRAL SUBPARAVENTRICULAR ZONE INFLUENCES GENERAL ACTIVITY RHYTHMS IN THE NILE GRASS RAT,ARVICANTHIS NILOTICUS

Circadian rhythms in behavior and physiology are very different in diurnal and nocturnal rodents. A pacemaker located in the suprachiasmatic nucleus (SCN) of the hypothalamus is responsible for generating and maintaining circadian rhythms in mammals, and cellular and molecular rhythms within the SCN of diurnal and nocturnal rodents are very similar. The neural substrates determining whether an animal has a diurnal or nocturnal phase preference are thus likely to reside downstream of the SCN. The ventral subparaventricular zone (vSPVZ), a major target of the SCN that is important for the expression of circadian rhythmicity in nocturnal lab rats (Rattus norvegicus), exhibits different rhythms in cFos expression in diurnal Nile grass rats compared to lab rats. We examined the effects of chemotoxic lesions of the cFos-expressing cells of the vSPVZ on activity rhythms of grass rats to evaluate the hypothesis that these cells support diurnality in this species. Male grass rats housed in a 12:12 light:dark (LD) cycle were given bilateral injections of the neurotoxin n-methyl-D-L-aspartic acid (NMA) or vehicle aimed at the vSPVZ; cells in the SCN are resistant to NMA, which kills neurons in other brain regions, but leaves fibers of passage intact. vSPVZ-damaged grass rats exhibited highly unstable patterns of activity in constant darkness (DD) and in the LD cycle that followed. However, crepuscular bouts of activity could be seen in all animals with vSPVZ lesions. Damage to the vSPVZ reduced cFos expression in this area but not in the SCN. Using correlational analyses, we found that the number of cFos-ir cells in the vSPVZ was unrelated to several parameters of the activity rhythms during the initial post-surgical period, when animals were in LD. However, the number of cells expressing cFos in the vSPVZ was positively correlated with general activity during the subjective day relative to the subjective night when the animals were switched to DD, and this pattern persisted when a LD cycle was reinstated. Also, the number of cFos-ir cells in the vSPVZ was negatively correlated with the strength of rhythmicity in DD and the number of days required to re-entrain to a LD cycle following several weeks in DD. These data suggest that the vSPVZ emits signals important for the expression of stable diurnal activity patterns in grass rats, and that species differences in these signals may contribute to differences in behavioral and physiological rhythms of diurnal and nocturnal mammals. (Author correspondence: mschw009@umaryland.edu)     

SYNCHRONIZATION TO LIGHT AND RESTRICTED-FEEDING SCHEDULES OF BEHAVIORAL AND HUMORAL DAILY RHYTHMS IN GILTHEAD SEA BREAM (SPARUS AURATA)

Food is not continuously available in the wild, and so most animals show a wide variety of circadian rhythms that can be entrained to feeding time. The aim of this research was to evaluate the effect of time-restricted feeding on the daily rhythms of gilthead sea bream, with food being provided during the day or night under a 12:12?h light-dark (LD) cycle or constant light (LL) conditions. Self-feeding and locomotor activity, as well as daily rhythms of cortisol, glucose, and melatonin, were evaluated. Fish synchronized their feeding behavior to the feeding phase, so that in LD they displayed 78% nocturnal feeding activity under night-feeding and 81% diurnal feeding activity under day-feeding, while under LL-feeding they displayed 72% of their daily activity during the 12?h feeding phase. In contrast, locomotor activity was mostly diurnal (66–71%), regardless of the feeding schedule, and it became arrhythmic under LL. Cortisol showed daily rhythms that peaked at different times, depending on the light and feeding schedule: one peak several hours before feeding under day-feeding and night-feeding, and two peaks under LL-feeding. Glucose displayed low-amplitude variations, with no daily rhythms being detected. Melatonin, however, showed a nocturnal rhythm, regardless of the feeding schedule, while the rhythm became attenuated under LL. Taken together, these results highlight the role of feeding on endocrine and metabolic rhythms, suggesting that feeding behavior should be considered when studying these variables. (Author correspondence: jflopez@um.es)     

Body temperature and behaviour of golden hamsters (Mesocricetus auratus) and ground squirrels (Spermophilus tridecemlineatus) in a thermal gradient

Golden hamsters and thirteen-lined ground squirrels were maintained individually in a thermal gradient (14°C to 33°C) for several weeks under a 14L: 10D light-dark cycle. Animals of both species showed robust daily rhythms of body temperature and locomotor activity with acrophases consistent with the habits of the species (diurnal acrophases in the diurnal squirrels and nocturnal acrophases in the nocturnal hamsters). Hamsters showed a robust daily rhythm of temperature selection 180° out of phase with the rhythms of body temperature and locomotor activity. Squirrels did not show a daily rhythm of temperature selection. These results raise the hypothesis that a daily rhythm of temperature selection is exhibited by nocturnal but not by diurnal endotherms.     

Temperature Cycles as Zeitgeber for the Circadian Clock of Two Burrowing Rodents, the Normothermic Antelope Ground Squirrel and the Heterothermic Syrian Hamster

The circadian systems of two burrowing rodents, the normothermic diurnal antelope ground squirrel (Ammospermophilus leucurus) and the heterothermic nocturnal Syrian hamster (Mesocricetus auratus) were compared with respect to entrainment by temperature cycles. Both species were subjected to the same ambient temperature (Ta) cycles with amplitudes between 4 and 12ºC at constant illuminations (100 and 0.05 lux in squirrels; 1.0 lux in hamsters). Wheel running activity was continuously measured. There was considerable interindividual variation in the daily pattern of wheel-running activity and in the ability to entrain to Ta cycles of the same amplitude in both species. The activity rhythms of about 33 to 67% of the animals of the two species entrained to Ta cycles with amplitudes of 6 to 12ºC. One of six squirrels and one of nine hamsters even entrained to Ta cycles of 4ºC. In the antelope ground squirrels, activity occurred predominantly in the cooler phase of the Ta cycle, whereas hamsters were mainly active during the warmer phase. In some squirrels, the activity rhythms were split in two main components which were both entrained to the cooler fraction of the Ta cycle, sometimes with additional (masking) activity during the warmer fraction (above 30ºC). The results do not support the earlier view that temperature cycles affect the circadian systems of heterothermic mammals, including hibernators, more strongly than those of normothermic species. It is suggested that behavioral and physiological adjustments to the environmental conditions play an important role for mammalian circadian systems to respond to temperature changes as a zeitgeber.     

Temperature Cycles as Zeitgeber for the Circadian Clock of Two Burrowing Rodents,the Normothermic Antelope Ground Squirrel and the Heterothermic Syrian Hamster

The circadian systems of two burrowing rodents, the normothermic diurnal antelope ground squirrel (Ammospermophilus leucurus) and the heterothermic nocturnal Syrian hamster (Mesocricetus auratus) were compared with respect to entrainment by temperature cycles. Both species were subjected to the same ambient temperature (Ta) cycles with amplitudes between 4 and 12ºC at constant illuminations (100 and 0.05 lux in squirrels; 1.0 lux in hamsters). Wheel running activity was continuously measured. There was considerable interindividual variation in the daily pattern of wheel-running activity and in the ability to entrain to Ta cycles of the same amplitude in both species. The activity rhythms of about 33 to 67% of the animals of the two species entrained to Ta cycles with amplitudes of 6 to 12ºC. One of six squirrels and one of nine hamsters even entrained to Ta cycles of 4ºC. In the antelope ground squirrels, activity occurred predominantly in the cooler phase of the Ta cycle, whereas hamsters were mainly active during the warmer phase. In some squirrels, the activity rhythms were split in two main components which were both entrained to the cooler fraction of the Ta cycle, sometimes with additional (masking) activity during the warmer fraction (above 30ºC). The results do not support the earlier view that temperature cycles affect the circadian systems of heterothermic mammals, including hibernators, more strongly than those of normothermic species. It is suggested that behavioral and physiological adjustments to the environmental conditions play an important role for mammalian circadian systems to respond to temperature changes as a zeitgeber.     

Olfactory bulbectomy modifies photic entrainment and circadian rhythms of body temperature and locomotor activity in a nocturnal primate

Studies on rodents have emphasized that removal of the olfactory bulbs modulates circadian rhythmicity. Using telemetric recordings of both body temperature (Tb) and locomotor activity (LA) in a male nocturnal primate, the gray mouse lemur, the authors investigated the effects of olfactory bulbectomy on (1) the circadian periods of Tb and LA in constant dim light condition, and (2) photic re-entrainment rates of circadian rhythms following 6-h phase shifts of entrained light-dark cycle (LD 12:12). Under free-running condition, bulbectomized males had significantly shorter circadian periods of Tb and LA rhythms than those of control males. However, the profiles of Tb rhythms, characterized by a phase of hypothermia at the beginning of the subjective day, and Tb parameters were not modified by olfactory bulbectomy. Under a light-dark cycle, olfactory bulbectomy significantly modified the expression of daily hypothermia, especially by an increase in the latency to reach minimal daily Tb, suggesting a delayed response to induction of daily hypothermia by light onset. Reentrainment rates following both a 6-h phase advance and a 6-h phase delay of entrained LD were also delayed in bulbectomized males. Olfactory bulbectomy led to significant fragmentation of locomotor activity and increased locomotor activity levels during the resting period. The shortening of circadian periods in bulbectomized males could partly explain the delayed responses to photic stimuli since in control males, the longer the circadian period, the better the response to light entrainment. This experiment shows for the 1st time that olfactory bulbs can markedly modify the circadian system in a primate.     

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.     

Low cost of locomotion in lizards that are active at low temperatures

The nocturnality hypothesis of K. Autumn and coworkers states that nocturnal geckos have evolved a low energetic cost of locomotion (C(min)). A low C(min) increases maximum aerobic speed and partially offsets the decrease in maximum oxygen consumption caused by activity at low nocturnal temperatures. We tested whether a low C(min) is unique to nocturnal geckos or represents a more general pattern of convergent evolution among lizards that enables nocturnality and/or cold-temperature activity. We measured C(min) in four carefully selected lizard species from New Zealand (two nocturnal and two diurnal; n=5-9 individuals per species), including a nocturnal and diurnal gecko (a low C(min) is a gecko trait and is not related to nocturnality), a nocturnal skink (a low C(min) is related to being nocturnal), and a diurnal skink active at low temperatures (a low C(min) is related to being active at low body temperatures). The C(min) values of the four species measured in this study (range=0.21-2.00 mL O(2) g(-1) km(-1)) are lower than those of diurnal lizards from elsewhere, and the values are within or below the 95% confidence limits previously published for nocturnal geckos. A low C(min) increases the range of locomotor speeds possible at low temperatures and provides an advantage for lizards active at these temperatures. We accepted the hypothesis that nocturnal lizards in general have a low C(min) and provide evidence for a low C(min) in lizards from cool-temperate environments. The low C(min) in lizards living at high latitudes may enable extension of their latitudinal range into otherwise thermally suboptimal habitats.     

Acute Behavioral Responses to Light and Darkness in Nocturnal Mus musculus and Diurnal Arvicanthis niloticus

The term masking refers to immediate responses to stimuli that override the influence of the circadian timekeeping system on behavior and physiology. Masking by light and darkness plays an important role in shaping an organism's daily pattern of activity. Nocturnal animals generally become more active in response to darkness (positive masking) and less active in response to light (negative masking), and diurnal animals generally have opposite patterns of response. These responses can vary as a function of light intensity as well as time of day. Few studies have directly compared masking in diurnal and nocturnal species, and none have compared rhythms in masking behavior of diurnal and nocturnal species. Here, we assessed masking in nocturnal mice (Mus musculus) and diurnal grass rats (Arvicanthis niloticus). In the first experiment, animals were housed in a 12:12 light-dark (LD) cycle, with dark or light pulses presented at 6 Zeitgeber times (ZTs; with ZT0 = lights on). Light pulses during the dark phase produced negative masking in nocturnal mice but only at ZT14, whereas light pulses resulted in positive masking in diurnal grass rats across the dark phase. In both species, dark pulses had no effect on behavior. In the 2nd experiment, animals were kept in constant darkness or constant light and were presented with light or dark pulses, respectively, at 6 circadian times (CTs). CT0 corresponded to ZT0 of the preceding LD cycle. Rhythms in masking responses to light differed between species; responses were evident at all CTs in grass rats but only at CT14 in mice. Responses to darkness were observed only in mice, in which there was a significant increase in activity at CT 22. In the 3rd experiment, animals were kept on a 3.5:3.5-h LD cycle. Surprisingly, masking was evident only in grass rats. In mice, levels of activity during the light and dark phases of the 7-h cycle did not differ, even though the same animals had responded to discrete photic stimuli in the first 2 experiments. The results of the 3 experiments are discussed in terms of their methodological implications and for the insight they offer into the mechanisms and evolution of diurnality.     

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.     

Seasonal variation of temporal niche in wild owl monkeys (Aotus azarai azarai) of the Argentinean Chaco: a matter of masking?

Among the more than 40 genera of anthropoid primates (monkeys, apes, and humans), only the South American owl monkeys, genus Aotus, are nocturnal. However, the southernmostly distributed species, Aotus azarai azarai, of the Gran Chaco may show considerable amounts of its 24-h activity during bright daylight. Due to seasonal changes in the duration of photophase and climatic parameters in their subtropical habitat, the timing and pattern of their daily activity are expected to show significant seasonal variation. By quantitative long-term activity recordings with Actiwatch AW4 accelerometer data logger devices of 10 wild owl monkeys inhabiting a gallery forest in Formosa, Argentina, the authors analyzed the seasonal variation in the temporal niche and activity pattern resulting from entrainment and masking of the circadian activity rhythm by seasonally and diurnally varying environmental factors. The owl monkeys always displayed a distinct bimodal activity pattern, with prominent activity bouts and peaks during dusk and dawn. Their activity rhythm showed distinct lunar and seasonal variations in the timing and daily pattern. During the summer, the monkeys showed predominantly crepuscular/nocturnal behavior, and a crepuscular/cathemeral activity pattern with similar diurnal and nocturnal activity levels during the cold winter months. The peak times of the evening and morning activity bouts were more closely related to the times of sunset and sunrise, respectively, than activity-onset and -offset. Obviously, they were better circadian markers for the phase position of the entrained activity rhythm than activity-onset and -offset, which were subject to more masking effects of environmental and/or internal factors. Total daily activity was lowest during the two coldest lunar months, and almost twice as high during the warmest months. Nighttime (21:00-06:00 h) and daytime (09:00-18:00 h) activity varied significantly across the year, but in an opposite manner. Highest nighttime activity occurred in summer and maximal daytime activity during the cold winter months. Dusk and dawn activity, which together accounted for 43% of the total daily activity, barely changed. The monkeys tended to terminate their nightly activity period earlier on warm and rainy days, whereas the daily amount of activity showed no significant correlation either with temperature or precipitation. These data are consistent with the dual-oscillator hypothesis of circadian regulation. They suggest the seasonal variations of the timing and pattern of daily activity in wild owl monkeys of the Argentinean Chaco result from a specific interplay of light entrainment of circadian rhythmicity and strong masking effects of various endogenous and environmental factors. Since the phase position of the monkeys' evening and morning activity peaks did not vary considerably over the year, the seasonal change from a crepuscular/nocturnal activity pattern in summer to a more crepuscular/cathemeral one in winter does not depend on a corresponding phase shift of the entrained circadian rhythm, but mainly on masking effects. Thermoregulatory and energetic demands and constraints seem to play a crucial role.     

Effects of environmental factors on circadian activity in the flesh fly, Sarcophaga crassipalpis

Abstract.The diel locomotor activity patterns of wandering larvae in the flesh fly, Sarcophaga crassipalpis Macquart (Diptera: Sarcophagidae), were examined using a novel apparatus and shown to be primarily diurnal, but with a minority (37%) showing nocturnal activity. In response to the environmental stress of heat shock, a significantly larger proportion (72%) of the larvae became nocturnal. In comparison, adult circadian activity also was predominantly diurnal, but not correlated with the larval activity patterns. In addition, adult patterns showed age-related changes in entrainment and free running period. Finally, the phase of circadian-gated adult eclosion was shown to be entrained by a 3-day exposure to light–dark cycles delivered prior to pupariation, with the phase maintained throughout pupal–adult metamorphosis under constant dark conditions. These results demonstrate that environmental changes may have profound effects on the expression of 24-h activity patterns and circadian rhythms during different life stages throughout development.     

Crepuscular activity of adult Sericesthis geminata (Coleoptera: Scarabaeidae): Influence of circadian rhythmicity and light intensity

Subterranean Sericesthis geminata (Boisduval) beetles emerge from the soil daily during the flight season. Emergence and flight are initiated within 15 min after sunset. The beetles are active above the soil surface as the intensity of illumination decreases from 240 lux to 0.7 lux.

If young adult beetles still in their pupal cells in the soil are placed under constant, low‐intensity illumination, they first emerge at any time of day, showing no cyclical diurnal activity pattern. When such beetles are exposed to diurnal fluctuations in illuminance, the first emergence and flight occur at dusk. These beetles subsequently show a diurnal rhythm in their activity, even when exposed to constant environmental conditions. This endogenous activity cycle has a period of about 22 h, and is synchronised with diurnal fluctuations in light intensity. It is reset in response to a change in the photoperiod.

The data suggest that, after initial emergence from the soil, daily crepuscular activity is initiated by an endogenously controlled activity rhythm which causes the beetles to burrow to the soil surface shortly before dusk. At the soil surface they are exposed to fluctuations in the duration and intensity of illumination which may reset the endogenous rhythm, affecting the time of subsequent emergence.