Locomotor Activity of the Fiddler Crab,Uca subcylindrica (Stimpson), under Artificial Illumination

Specimens of the fiddler crab Uca subcylindrica (Stimpson) were captured in south Texas (USA) for locomotor rhythm studies. Actographic data were analyzed using Tau? sofware. Under constant illumination (LL) and darkness (DD), the semiterrestrial crabs express a circadian rhythm of locomotion. When exposed to illumination/darkness cycles (LD_12:12 or LD_14:10), their bouts of activity are entrained to the photoperiod. In LD, activity is generally bimodal with an initial burst about 0.5 h after illumination. A second burst of activity begins 1 to 2 h before the end of illumination. When transferred from LD to LL, a locomotor rhythm with an average period of 24.6 ± 1.0 h (n = 19) is expressed in 89 percent of the crabs. On the other hand, when placed in DD after LD (n = 8), the crabs are either arrythmic or weakly rhythmic (period = 23.8 ± 0.2 h; n = 2). If the onset of illumination is advanced by 6 h, a period of less than 24.0 h is detected in the actogram. If the onset of illumination is delayed by 6 h, a locomotor rhythm with a period greater than 24.0 h appears. The locomotor behavior of this species of fiddler crab, Uca subcylindrica, is not related to the tidal rhythmicities seen in other members of the genus Uca. Rather, it has strong circadian components.     

Locomotor Rhythms in the Fiddler Crab, Uca subcylindrica

The locomotor activities of individual specimens of Uca subcylindrica (Stimpson) collected from semi-arid, supratidal habitats in south Texas and northeastern Mexico were studied in the laboratory using periodogram analysis. When crabs were placed under constant darkness (DD) or constant illumination (LL), free-running circadian rhythms were observed in the activity recordings. The locomotor activity of strongly rhythmic crabs in LL has an average period length of 24.4 h. Crabs held in DD express motor rhythms with periods of approximately 24.0 h. In LL the most common wave form for activity is unimodal, while under DD it is bimodal. Recordings under natural illumination (NL) revealed that both period length and the time of maximum activity (phasing) varied through the year. During winter months, the crabs are primarily diurnal with peaks in activity occurring between 0900 and 2100 h and possess a circadian rhythm with a 23.9 h period. During summer, crabs were nocturnal with maximal activity between 1300 and 0600 and a circadian period closer to 24.0 h. In these experiments, the rhythmic locomotor activities of U. subcylindrica are best described as “circadian”. This is unusual for a genus known for its expression of circatidal and circalunidian rhythms.     

Presence of circadian rhythms in the locomotor activity of a cave-dwelling millipede Glyphiulus cavernicolus sulu (Cambalidae, Spirostreptida)

The locomotor activity of the millipede Glyphiulus cavernicolus (Spirostreptida), which occupies the deeper recesses of a cave, was monitored in light-dark (LD) cycles (12h light and 12h darkness), constant darkness (DD), and constant light (LL) conditions. These millipedes live inside the cave and are apparently never exposed to any periodic factors of the environment such as light-dark, temperature, and humidity cycles. The activity of a considerable fraction of these millipedes was found to show circadian rhythm, which entrained to a 12:12 LD cycle with maximum activity during the dark phase of the LD cycle. Under constant darkness (DD), 56.5% of the millipedes (n = 23) showed circadian rhythms, with average free-running period of 25.7h ± 3.3h (mean ± SD, range 22.3h to 35.0h). The remaining 43.5% of the millipedes, however, did not show any clear-cut rhythm. Under DD conditions following an exposure to LD cycles, 66.7% (n = 9) showed faint circadian rhythm, with average free-running period of 24.0h ± 0.8h (mean ± SD, range 22.9h to 25.2h). Under constant light (LL) conditions, only 2 millipedes of 11 showed free-running rhythms, with average period length of 33.3h ± 1.3h. The results suggest that these cave-dwelling millipedes still possess the capacity to measure time and respond to light and dark situations. (Chronobiology International, 17(6), 757-765, 2000)     

Locomotor Rhythms in the Fiddler Crab,Uca subcylindrica

The locomotor activities of individual specimens of Uca subcylindrica (Stimpson) collected from semi-arid, supratidal habitats in south Texas and northeastern Mexico were studied in the laboratory using periodogram analysis. When crabs were placed under constant darkness (DD) or constant illumination (LL), free-running circadian rhythms were observed in the activity recordings. The locomotor activity of strongly rhythmic crabs in LL has an average period length of 24.4 h. Crabs held in DD express motor rhythms with periods of approximately 24.0 h. In LL the most common wave form for activity is unimodal, while under DD it is bimodal. Recordings under natural illumination (NL) revealed that both period length and the time of maximum activity (phasing) varied through the year. During winter months, the crabs are primarily diurnal with peaks in activity occurring between 0900 and 2100 h and possess a circadian rhythm with a 23.9 h period. During summer, crabs were nocturnal with maximal activity between 1300 and 0600 and a circadian period closer to 24.0 h. In these experiments, the rhythmic locomotor activities of U. subcylindrica are best described as “circadian”. This is unusual for a genus known for its expression of circatidal and circalunidian rhythms.     

Circadian Locomotor Activity Under Artificial Light in the Freshwater Crab Pseudothelphusa americana

Long-term recordings of locomotor activity were obtained from intact freshwater crabs, Pseudothelphusa americana in constant darkness (DD), constant light (LL) and different light-dark (LD) protocols. Bimodal rhythms were typically observed in this crab when subjected to DD or LD, with bouts of activity anticipating lights-on and lights-off, respectively. Freerunning circadian rhythms were expressed in both DD and LL for longer than 30 days. In DD, we observed that some animals presented different period lengths for each activity component. During LL, activity was primarily unimodal, however spontaneous splitting of the rhythms were observed in some animals. When activity was recorded under artificial long days, the morning bouts maintained their phase relationship but the evening bouts changed their phase relationship with the Zeitgeber. Our results indicate that, bimodal locomotor activity rhythm in the crab Pseudothelphusa americana is variable among organisms. The characteristics of phase relationship with LD and responses to LL for morning and evening bouts, suggest that, locomotor activity could be driven by multiple oscillators, and that coupling between these oscillators may be regulated by light.     

Circadian Locomotor Activity Under Artificial Light in the Freshwater Crab Pseudothelphusa americana

Long-term recordings of locomotor activity were obtained from intact freshwater crabs, Pseudothelphusa americana in constant darkness (DD), constant light (LL) and different light-dark (LD) protocols. Bimodal rhythms were typically observed in this crab when subjected to DD or LD, with bouts of activity anticipating lights-on and lights-off, respectively. Freerunning circadian rhythms were expressed in both DD and LL for longer than 30 days. In DD, we observed that some animals presented different period lengths for each activity component. During LL, activity was primarily unimodal, however spontaneous splitting of the rhythms were observed in some animals. When activity was recorded under artificial long days, the morning bouts maintained their phase relationship but the evening bouts changed their phase relationship with the Zeitgeber. Our results indicate that, bimodal locomotor activity rhythm in the crab Pseudothelphusa americana is variable among organisms. The characteristics of phase relationship with LD and responses to LL for morning and evening bouts, suggest that, locomotor activity could be driven by multiple oscillators, and that coupling between these oscillators may be regulated by light.     

Circadian Rhythm in the Locomotor Activity of a Surface-Dwelling Millipede Syngalobolus sp.

Locomotor activity of the surface-dwelling millipede Syngalobolus sp. was recorded under laboratory conditions. Infra-red diodes were used to detect the locomotor activity in an oval shaped chamber, which was connected with an event recorder. The results of 11 individuals showed that the millipedes entrained to light/dark (LD12:12 h) conditions with negative phase angle difference (-83.2 ± 24.72 min). The millipedes showed a clear-cut free-running rhythm with a period (t) of 23.8 ± 1.0 h (n = 9) in constant darkness (DD). The period in continuous light (LL) was relatively greater (25.2 ± 0.1 h; n = 3) than that in DD.     

Entrainment of eclosion rhythm in Drosophila melanogaster populations reared for more than 700 generations in constant light environment

In this paper, we report the results of our extensive study on eclosion rhythm of four independent populations of Drosophila melanogaster that were reared in constant light (LL) environment of the laboratory for more than 700 generations. The eclosion rhythm of these flies was assayed under LL, constant darkness (DD) and three periodic light-dark (LD) cycles (T20, T24, and T28). The percentage of vials from each population that exhibited circadian rhythm of eclosion in DD and in LL (intensity of approximately 100 lux) was about 90% and 18%, respectively. The mean free-running period (τ) of eclosion rhythm in DD was 22.85 ± 0.87 h (mean ± SD). Eclosion rhythm of these flies entrained to all the three periodic LD cycles, and the phase relationship (ψ) of the peak of eclosion with respect to “lights-on” of the LD cycle was significantly different in the three periodic light regimes (T20, T24, and T28). The results thus clearly demonstrate that these flies have preserved the ability to exhibit circadian rhythm of eclosion and the ability to entrain to a wide range of periodic LD cycles even after being in an aperiodic environment for several hundred generations. This suggests that circadian clocks may have intrinsic adaptive value accrued perhaps from coordinating internal metabolic cycles in constant conditions, and that the entrainment mechanisms of circadian clocks are possibly an integral part of the clockwork.     

PRESENCE OF CIRCADIAN RHYTHMS IN THE LOCOMOTOR ACTIVITY OF A CAVE-DWELLING MILLIPEDE GLYPHIULUS CAVERNICOLUS SULU (CAMBALIDAE,SPIROSTREPTIDA)

The locomotor activity of the millipede Glyphiulus cavernicolus (Spirostreptida), which occupies the deeper recesses of a cave, was monitored in light-dark (LD) cycles (12h light and 12h darkness), constant darkness (DD), and constant light (LL) conditions. These millipedes live inside the cave and are apparently never exposed to any periodic factors of the environment such as light-dark, temperature, and humidity cycles. The activity of a considerable fraction of these millipedes was found to show circadian rhythm, which entrained to a 12:12 LD cycle with maximum activity during the dark phase of the LD cycle. Under constant darkness (DD), 56.5% of the millipedes (n = 23) showed circadian rhythms, with average free-running period of 25.7h ± 3.3h (mean ± SD, range 22.3h to 35.0h). The remaining 43.5% of the millipedes, however, did not show any clear-cut rhythm. Under DD conditions following an exposure to LD cycles, 66.7% (n = 9) showed faint circadian rhythm, with average free-running period of 24.0h ± 0.8h (mean ± SD, range 22.9h to 25.2h). Under constant light (LL) conditions, only 2 millipedes of 11 showed free-running rhythms, with average period length of 33.3h ± 1.3h. The results suggest that these cave-dwelling millipedes still possess the capacity to measure time and respond to light and dark situations. (Chronobiology International, 17(6), 757–765, 2000)     

Temperature dependent eclosion rhythmicity in the high altitude Himalayan strains of Drosophila ananassae

The circadian pacemaker controlling the eclosion rhythm of the high altitude Himalayan strains of Drosophila ananassae captured at Badrinath (5123 m) required ambient temperature at 21°C for the entrainment and free-running processes. At this temperature, their eclosion rhythms entrained to 12h light, 12h dark (LD 12:12) cycles and free-ran when transferred from constant light (LL) to constant darkness (DD) or upon transfer to constant temperature at 21°C following entrainment to temperature cycles in DD. These strains, however, were arrhythmic at 13 or 17°C under identical experimental conditions. Eclosion medians always occurred in the thermophase of temperature cycles whether they were imposed in LL or DD; or whether the thermophase coincided with the photophase or scotophase of the concurrent LD 12:12 cycles. The temperature dependent rhythmicity in the Himalayan strains of D. ananassae is a rare phenotypic plasticity that might have been acquired through natural selection by accentuating the coupling sensing mechanism of the pacemaker to temperature, while simultaneously suppressing the effects of light on the pacemaker.     

Circadian Rhythms of Locomotor Activity in Lycosa tarentula (Araneae, Lycosidae) and the Pathways of Ocular Entrainment

Lycosa tarentula is a ground-living spider that inhabits a burrow where it awaits the appearance of prey or conspecifics. In this study, circadian rhythms of locomotor activity were examined as well as the ocular pathway of entrainment. Thirty-three adult virgin females were examined under constant darkness (DD); all of them exhibited robust circadian rhythms of locomotor activity with a period averaging 24.1h. Fourteen of these spiders were studied afterwards under an LD 12:12 cycle; they usually entrained to in the first or second day, even when the light intensity was as low as 1 lx. During the LD cycle, locomotor activity was generally restrained to the darkness phase, although several animals showed a small amount of diurnal activity. Ten males were also examined under LD; they were also nocturnal, but were much more active than the females. Seven females were examined under constant light (LL); under this they became arrhythmic. Except for the anterior median eyes (OMAs), all the eyes were capable of entraining the locomotor activity to an LD cycle. These results demonstrate that under laboratory conditions and low light intensities locomotor activity of Lycosa tarentula is circadian and in accordance with Aschoff's 'rule'. Only OMAs are unable to entrain the rhythm; the possible localization of circadian clock is therefore discussed.     

About the Existence of Circadian Activity in Cave Crayfish

Evidence of a circadian clock mechanism was found in the cave crayfish Procambarus cavernicola. Analysis of motor activity recorded in this species during 12 consecutive days in either free running (constant darkness, DD or constant light, LL) or entrainment conditions (12 h of light alternated with 12 h of darkness, 12 : 12 LD) showed a well recognized circadian rhythm. In this rhythm however, the absence of synchronization by periodical external signals was notorious. The comparison between the motor circadian rhythm in cave crayfish and epigeous crayfish Procambarus clarkii (these last studied during juvenile and adult stages), evidenced strong similitude between the motor circadian rhythm of cave crayfish and juvenile epigeous crayfish.     

Effect of light on the development of the circadian rhythm of motor activity in the mouse

In previous experiments, we found that rats raised in constant light (LL) manifested a more robust circadian rhythm of motor activity in LL and showed longer phase shifts after a light pulse in constant darkness (DD) than those raised under constant darkness. In addition, we observed that the effects produced by constant light differed depending on the time of postnatal development in which it was given. These results suggest that both sensitivity to light and the functioning of the circadian pacemaker of the rat could be affected by the environmental conditions experienced during postembryonic development. Thus, the present experiment aimed to study whether postnatal exposure to light could also affect the circadian system of the mouse. Three groups of mice were formed: One group was raised under constant darkness during lactation (DD group), the second under constant light (LL group), and the third under light-dark cycles (LD group). After lactation, the three groups were submitted first to constant light of high intensity, then to LD cycles, and finally to constant darkness. In the DD stage, a light pulse was given. Finally, mice were submitted to constant light of low intensity. We observed that the circadian rhythm of the DD group was more disturbed under constant light than the rhythm of the LL group, and that, when light intensity increased, the period of the rhythm of the DD group lengthened more than that of the LL group. No significant differences among the groups were found in the phase shift induced by the light pulse. Therefore, it appears that DD mice are more sensitive to light than their LL counterparts. However, at present there is no evidence to affirm that the light environment experienced by the mouse during postnatal development affects the circadian pacemaker. (Chronobiology International, 18(4), 683-696, 2001)     

Multi-Oscillatory Control of Eclosion and Oviposition Rhythms in Drosophila melanogaster: Evidence from Limits of Entrainment Studies

The eclosion and oviposition rhythms of flies from a population of Drosophila melanogaster maintained under constant conditions of the laboratory were assayed under constant light (LL), constant darkness (DD), and light/dark (LD) cycles of 10:10 h (T20), 12:12 h (T24), and 14:14 h (T28). The mean (±95% confidence interval; CI) free-running period (τ) of the oviposition rhythm was 26.34 ± 1.04 h and 24.50 ± 1.77 h in DD and LL, respectively. The eclosion rhythm showed a τ of 23.33 ± 0.63 h (mean ± 95% CI) in DD, and eclosion was not rhythmic in LL. The τ of the oviposition rhythm in DD was significantly greater than that of the eclosion rhythm. The eclosion rhythm of all 10 replicate vials entrained to the three periodic light regimes, T20, T24, and T28, whereas the oviposition rhythm of only about 24 and 41% of the individuals entrained to T20 and T24 regimes, respectively, while about 74% of the individuals assayed in T28 regimes showed entrainment. Our results thus clearly indicate that the τ and the limits of entrainment of eclosion rhythm are different from those of the oviposition rhythm, and hence this reinforces the view that separate oscillators may regulate these two rhythms in D. melanogaster.     

Circadian Rhythms and time Measurement in Locomotor Activity of the Scorpion Leiurus Quinquestriatus (Buthidae)

The circadian rhythms of locomotor activity of the scorpion Leiurus quinqueslriatus were examined under different light-dark cycles and in free-running conditions. The circadian rhythm is bimodal in LD 12:12 with alternating cycles of temperature (35°-25°C) with high intensity (1300 lux) or in LD 12: 12 with constant temperature 35° C with 300 lux. In LD 12:12 (1300 lux), in long or in short light spans with constant temperature, the bimodal pattern is slightly changed with the appearance of a third minor peak of activity. In free-running conditions, the bimodal rhythm of locomotor activity persists in DD with T about 24 hr, but in LL the rhythm becomes unimodal with T about 24 hr. Cosinor and power spectrum analysis showed the presence of more than one periodic component. It seems that there is a correlation between the range of light regimens, temperature, light intensity and the coincidence of these components. These components are independently entrained by the environmental light cycle. The mechanism of entrainment of components is discussed.     

Possible evidence for shift work schedules in the media workers of the ant species Camponotus compressus

The locomotor activity rhythm of the media workers of the ant species Camponotus compressus was monitored under constant conditions of the laboratory to understand the role of circadian clocks in social organization. The locomotor activity rhythm of most ants entrained to a 24 h light/dark (12:12 h; LD) cycle and free-ran under constant darkness (DD) with circadian periodicities. Under entrained conditions about 75% of media workers displayed nocturnal activity patterns, and the rest showed diurnal activity patterns. In free-running conditions these ants displayed three types of activity patterns (turn-around). The free-running period (τ) of the locomotor activity rhythm of some ants (10 out of 21) showed period lengthening, and those of a few (6 out of 21) showed period shortening, whereas the locomotor activity rhythm of the rest of the ants (5 out of 21) underwent large phase shifts. Interestingly, the pre-turn-around τ of those ants that showed nocturnal activity patterns during earlier LD entrainment was shorter than 24 h, which became greater than 24 h after 6-9 days of free-run in DD. On the other hand, the pre-turn-around τ of those ants, which exhibited diurnal patterns during earlier LD entrainment, was greater than 24 h, which became shorter than 24 h after 6-9 days of free-run in DD. The patterns of activity under LD cycles and the turn-around of activity patterns in DD regime suggest that these ants are shift workers in their respective colonies, and they probably use their circadian clocks for this purpose. Circadian plasticity thus appears to be a general strategy of the media workers of the ant species C. compressus to cope with the challenges arising due to their roles in the colony constantly exposed to a fluctuating environment.     

Circadian Rhythm of Blood Ethanol Clearance Rates in Rats: Response to Reversal of the L/D Regimen and to Continuous Darkness and Continuous Illumination

Phase relationships of the circadian rhythms of blood ethanol clearance (metabolic) rates and body temperature were studied in rats successively exposed to 4 illumination regimens: LD (light from 0800-2000 hr), DL (light from 2000-0800 hr), constant darkness (DD) and, lastly, constant light (LL). After a 4-wk standardization to each regimen, body temperatures were taken at 9 × 4-hr intervals to establish baseline circadian profiles. One week later, groups (N = 8) received 1.5 g/kg ethanol (i.p.) at 6 equally spaced timepoints during a 24-hr span, when temperatures were again measured. Ethanol clearance rates were estimated from decreasing blood ethanol levels sampled every 20 min from 60-200 min after dosing, and the resultant elimination curves were subjected to cosinor analysis. These studies show for the first time that the high amplitude circadian rhythm in ethanol metabolism persists under constant conditions of illumination (DD and LL), demonstrating that it may well be a truly internal circadian rhythm and not a response to exogenous cues of the light/dark cycle. During both LD and DL, maximal and minimal ethanol clearance rates fell near the end of the dark and light phases, respectively, and followed circadian peak and trough control temperatures by approximately 6 hr. A fixed internal phase relationship between the core body temperature and the circadian rhythm in ethanol metabolism is demonstrated, thus establishing the rhythm in body temperature as a suitable and convenient internal marker rhythm for studies of the metabolism of low-to-moderate ethanol doses. These studies demonstrate that the phase relationships of blood ethanol clearance rate and body temperature can be manipulated by the illumination regimen selected, an observation of both basic and practical importance.     

Circadian Locomotor Activity Rhythm in the Freshwater Crab Pseudothelphusa americana (De Saussure, 1857): Effect of Eyestalk Ablation

The circadian rhythm of locomotor activity in the freshwater crab, Pseudothelphusa americana , was studied in aquaria using infrared crossing sensors. Individuals with ablated eyestalks were compared with intact individuals in constant darkness (DD) and in light-dark cycles (LD). Our results showed that intact animals in DD displayed bimodal rhythms. In LD conditions the two peaks were associated with lights on and lights off, respectively. A significant difference in the free running periods before and after LD was observed in all intact animals. After eyestalk ablation (ES-X), the circadian rhythm of locomotor activity disappeared immediately, but reappeared several days later. Diurnal activity was seen in some ES-X animals when exposed to LD. Our results indicate that locomotor activity rhythm in P. americana is driven primarily by oscillators located outside the eyestalks, and that extraretinal photoreceptors mediate either entrainment or masking effects.     

Circadian Locomotor Activity Rhythm in the Freshwater Crab Pseudothelphusa americana (De Saussure, 1857): Effect of Eyestalk Ablation

The circadian rhythm of locomotor activity in the freshwater crab, Pseudothelphusa americana, was studied in aquaria using infrared crossing sensors. Individuals with ablated eyestalks were compared with intact individuals in constant darkness (DD) and in light-dark cycles (LD). Our results showed that intact animals in DD displayed bimodal rhythms. In LD conditions the two peaks were associated with lights on and lights off, respectively. A significant difference in the free running periods before and after LD was observed in all intact animals. After eyestalk ablation (ES-X), the circadian rhythm of locomotor activity disappeared immediately, but reappeared several days later. Diurnal activity was seen in some ES-X animals when exposed to LD. Our results indicate that locomotor activity rhythm in P. americana is driven primarily by oscillators located outside the eyestalks, and that extraretinal photoreceptors mediate either entrainment or masking effects.     

Entrainment Properties of the Locomotor Activity Rhythm of Drosophila melanogaster Under Different Photoperiodic Regimens

In this paper we report the results of an experiment to assess how closely repeated brief light pulses (LPs) mimic the effects of 12:12 h light/dark (LD) cycles (PPc). The locomotor activity rhythm of individual fruit flies from a laboratory population of Drosophila melanogaster was monitored under four different photoperiodic regimens, created using 12 h of light and 12 h of darkness or brief light pulses (LPs). The phase relationship (Ψ) and the stability (precision) of the locomotor activity rhythm during entrainment were estimated in order to compare the state of the circadian clocks under the four different photoperiodic regimens. The flies (n = 72) were subjected to four different LD cycles: (i) 12 h of light and 12 h of darkness (complete photoperiod, PPc); (ii) a single brief LP of 15 min duration presented close to the onset of activity (SLP-1); (iii) a single brief LP of 15 min duration administered close to the offset of activity (SLP-2); and (iv) two brief LPs administered 12 h apart (skeleton photoperiod, PPs). The locomotor activity rhythm of the flies was first monitored under constant darkness (DD) for about 10 days and then under the four different photoperiodic regimens for about 10 days, and finally in DD for the remainder of the experiment. The Ψ of the locomotor activity rhythm and its precision under PPc and PPs did not differ significantly, but they were significantly different from the SLP-1 and SLP-2 conditions. The results provide interesting insights into photoentrainment mechanisms of circadian clocks in D. melanogaster, and suggest that skeleton photoperiods, but not single brief LPs, mimic the actions of complete photoperiods.