Circadian and circatidal rhythms of oxygen consumption in the sandy-beach isopod Tylos granulatus Krauss

The sandy-beach isopod Tylos granulatus Krauss burrows at the high tide mark, and has rhythms of nocturnal emergence coincident with the low tide period. Measurements of the respiration rate show that a low rate of oxygen consumption is maintained throughout the day but there is a circatidal rhythmic increase of between 300 and 700% during the nocturnal low tide. The height of this peak and the percentage of animals displaying a peak both increase from spring tide to neap tide suggesting a semi-lunar rhythm. The respiratory peaks have a 24.8 h periodicity, being later each night until low tide falls in the dawn, when there is a ‘switch-back’ so that peaks are then during the subsequent evening low tide. These respiratory rhythms are persistent under constant light and coincide with previously described activity rhythms. The respiratory rhythms considerably reduce metabolic energy losses, particularly as the activity rhythms ensure avoidance of high diurnal temperatures and activity during the cooler nocturnal period.     

Comparative studies of tidal rhythms. VIII. A translocation experiment involving Circalunidian rhythms

One of the earliest experiments that provided support for the exogenous clock hypothesis was a translocation experiment involving fiddler crabs. The activity rhythm of a sample of crabs placed in constant conditions, abandoned (in a week's time) the phase of the tidal cycle to which the crabs had been exposed in nature, and the peaks attuned themselves to the approximate times of lunar nadir and zenith. The study reported in the present paper was an attempt to repeat this interesting and important finding.

The effort at replication was unsuccessful. But other more recent findings of tide‐associated rhythms, such as splitting, uncoupling, and temporary arrhythmicity, were confirmed. And an “ultradian”; rhythm was discovered to exist in conjunction with a typical circalunidian frequency.     

Diurnal Variation in Temperature,Mental and Physical Performance,and Tasks Specifically Related to Football (Soccer)

Football (soccer) training and matches are scheduled at different times throughout the day. Association football involves a variety of fitness components as well as psychomotor and game‐related cognitive skills. The purpose of the present research, consisting of two separate studies, was to determine whether game‐related skills varied with time of day in phase with global markers of both performance and the body clock. In the first study, eight diurnally active male association football players (19.1±1.9 yrs of age; mean±SD) with 10.8±2.1 yrs playing experience participated. Measurements were made on different days at 08:00, 12:00, 16:00, and 20:00 h in a counterbalanced manner. Time‐of‐day changes in intra‐aural temperature (used as a marker of the body clock), grip strength, reaction times, flexibility (markers of aspects of performance), juggling and dribbling tasks, and wall‐volley test (football‐specific skills) were compared. Significant (repeated measures analysis of variance, ANOVA) diurnal variations were found for body temperature (p<0.0005), choice reaction time (p<0.05), self‐rated alertness (p<0.0005), fatigue (p<0.05), forward (sit‐and‐reach) flexibility (p<0.02), and right-hand grip strength (p<0.02), but not left-hand grip strength (p=0.40) nor whole‐body (stand‐and‐reach) flexibility (p=0.07). Alertness was highest and fatigue lowest at 20:00 h. Football‐specific skills of juggling performance showed significant diurnal variation (p<0.05, peak at 16:00 h), whereas performance on the wall‐volley test tended to peak at 20:00 h and dribbling showed no time‐of‐day effect (p=0.55). In a second study, eight diurnally active subjects (23.0±0.7 yrs of age) completed five test sessions, at the same times as in the first study but with a second session at 08:00 h. Test‐re‐test comparisons at 08:00 h for all components indicated good reliability. Intra‐aural temperature showed a significant time‐of‐day effect (p<0.001) with mean temperature at 16:00 h (36.4°C) higher than at 08:00 h (35.4°C). There was no significant effect of chronotype on the temperature acrophase (peak time) (p>0.05). Diurnal variation was found for performance tests, including sit‐and‐reach flexibility (p<0.01) and spinal hyper‐extension (p<0.05). Peaks occurred between 16:00 and 20:00 h and the daytime changes paralleled the temperature rhythm. Diurnal variation was also found for football‐specific tests, including dribbling time (p<0.001, peak at 20:00 h) and chip test performance (p<0.01), being more accurate at 16:00 h (mean error=0.75 m) than at 08:00 h (mean error=1.01 m). Results indicate football players perform at an optimum between 16:00 and 20:00 h when not only football‐specific skills but also measures of physical performance are at their peak. Body temperature peaked at a similar time, but positive mood states seemed to peak slightly earlier. While causal links cannot be established in these experiments, the results indicate that the diurnal variation of some aspects of football performance is affected by factor(s) other than body temperature alone.     

24‐H entrainment and ultradian fluctuations in the activity of the lizard Gallotia galloti galloti (sauria‐lacertidae)

Abstract

Activity data from two experimental lizard groups were analysed in order to search for 24 h‐entrained and ultradian periodicities. The data of a first group were obtained through motion sensitive platforms situated under the animals’ cage and continuously for up to 12 consecutive days; those from a second group were collected by manual recording of the behaviour patterns of individual animals for 2 h each day over a 10‐day period. Lizards from both groups were situated in cages inside isolated chambers in which a light‐dark cycle (12:12), a temperature of 28°C ±1° and a relative humidity of 50–60% were maintained. Periodogram analysis showed the existence of a significant period peak (p < 0.01) at 24 h. Autocorrelation functions and spectral analysis of different data‐segment lengths showed that ultradian periodicities were present in the daily motor activity, appearing as noisy though frequency‐band limited. Differences in the frequency band‐limited fluctuations were found between morning and afternoon activities: during the morning ultradian activity appears distributed in two bands (4.5–36 c/day and 63–94 c/day), the power being mainly concentrated in the second one, while during the afternoon it was in the 4.5–36 c/day band. Results from the second lizard group showed only one band (24–60 c/day) overlapped with the first one from the latter group. Although activity recording methods and some experimental conditions are discussed as possible sources of these differences, the possibility of endogenous ultradian variation within the individuals is also suggested.     

Juvenile Chinese mitten crabs (<Emphasis Type="Italic">Eriocheir sinensis</Emphasis>) in the Thames estuary: distribution,movement and possible interactions with the native crab <Emphasis Type="Italic">Carcinus maenas</Emphasis>

The Chinese mitten crab, Eriocheir sinensis is a successful invader. Whilst non-breeding adult E. sinensis have been associated with the destruction of riverbanks, little is known about the ecology of this species in its invaded areas. This is especially true of the juveniles which are a key migratory stage. Intertidal surveys along the Thames estuary indicated an increase in the population since the 1990s. Juvenile E. sinensis were abundant in the sampled upper tidal region of the Thames, refuging under boulders in the intertidal at low tide. Seasonal differences in sampled populations were observed, with a significantly lower abundance of crabs found during winter compared to summer. Mark-recaptures indicated movement in the intertidal occurring during high tide, with an influx of new crabs evident after a single tidal cycle. Endogenous rhythms were also apparent, with peaks in activity occurring corresponding with night-time high tide conditions. E. sinensis juveniles can successfully exclude similar sized native Carcinus maenas from shelters in the laboratory, regardless of which species originally inhabited the shelter. This may have implications for native estuarine Carcinus populations, which can rely on such intertidal shelters for refuges.     

Review of diel rhythmic activities in physiological functions of scorpions

In this study I review the subject of biological rhythms in scorpions. This will include only some of the diel rhythmic endogenous physiological functions and not the locomotory rhythmicity, which will be dealt separately. Most studies reported here were conducted on 13 scorpion species that were studied in 39 different studies. Most of these (66.7%) were studies on a single species (Heterometrus fulvipes). Being a scorpionid, it sits and waits near its burrow not being very active, especially the females. The fact that experimenting was carried out irrespective of species diversity, gender, ecological or physiological conditions, and was usually done on animals kept in captivity for some time before the experimenting had started, is a major drawback to this kind of study. Although the main conclusion appears to be that enzymes reached their peak activity at 20.00 h, there are some exceptions showing otherwise that need further study in order to explain them.     

IMPAIRED EXPRESSION OF THE mPer2 CIRCADIAN CLOCK GENE IN THE SUPRACHIASMATIC NUCLEI OF AGING MICE

The expression of circadian clock genes was investigated inthe suprachiasmatic nuclei (SCN) of young adult and old laboratory mice. Sampleswere taken at two time points, which corresponded to the expected maximum(circadian time 7 [CT7]) or minimum (CT21) of mPermRNA expression. Whereas the young mice had a stable and well-synchronizedcircadian activity/rest cycle, the rhythms of old animals were less stableand were phase advanced. The expression of mPer1mRNA and mPer2 mRNA was rhythmic in bothgroups, with peak values at CT7. The levels of mClockand mCry1 mRNA were not different dependingon the time of day and did not vary with age. In contrast, an age-dependentdifference was found in the case of mPer2(but not mPer1) mRNA expression, with themaximum at CT7 significantly lower in old mice. The decreased expression of mPer2 may be relevant for the observed differencesin the overt activity rhythm of aged mice. (ChronobiologyInternational, 18(3), 559–565, 2001)     

Making up for lost time: Biophysical constraints on the temporal abundance of two fiddler crabs in wet–dry tropical mangroves

Biophysical models are used to predict the spatial distributions of organisms. Nevertheless, understanding factors influencing the temporal distributions of animals may often be additionally required. It is expected that intertidal macrofauna of the wet–dry tropics face a multitude of temporal challenges because there is not only seasonal drying but also variation in surface moisture over the circatidal cycle. Activities of fiddler crabs (Uca spp.) depend on adequate surface moisture being available for feeding and respiration. A recent study monitored crab abundance during spring tides and found that one Uca species in the mangroves of Darwin Harbour, Australia, U. flammula, is most abundant in the wet season, while another, U. elegans, is most abundant in the dry season. We hypothesized here that these seemingly contradictory abundance patterns are driven by temporal variation in the availability of soil moisture within each species habitat. We thus monitored crab abundance and measured soil moisture content across four types of habitat (low gap centres, low gap edges, mid‐height gap centres and high gap centres) seasonally and across the circatidal cycle. We found that crab abundance and soil moisture both varied over time among habitat types. We used a log‐linear model to show that habitat type influenced soil moisture and this in turn influenced crab abundance. Sampling across the circatidal cycle showed that U. flammula was more abundant in the wet season, as reported previously, while the abundance of U. elegans did not vary between seasons. Our model suggested that U. elegans ‘makes up for lost time’ in the dry season by undertaking all activities during spring tide low water as only at this time is the substratum moist enough for feeding and respiration. We highlight the importance of measuring multiple variables across habitats over small and large scales when assessing temporal abundance patterns of intertidal tropical organisms.     

DIEL RHYTHMS OF CALLING BEHAVIOR AND SEX PHEROMONE PRODUCTION OF BEET ARMYWORM,SPODOPTERA EXIGUA (LEPIDOPTERA:NOCTUIDAE)*

Abstract Beet armyworm (BAW), Spodoptera exigua, is becoming one of more and more serious pests in China in recent years. As a part of research program of sex pheromone and its application of BAW in China, the hourly and daily variation of calling behavior and pheromone production of BAW females were investigated. Both calling behavior and titers of 4 sex pheromone components showed distinct diel rhythms, and the two peak periods were synchronous. In comparison, the calling activity lasted shorter period of time with a longer peak time, whereas the production of the sex pheromone lasted throughout the whole scotophase and part of the photophase with a very short peak time. The calling behavior began at the middle scotophase, reached the maximum at the middle‐later scotophase, and continued the maximal calling activity until the end of the scotophase. When the light was on, the calling percentage reduced sharply, and all females stopped calling 1 hour later. The variation patterns of the 4 pheromone components in the glands of the 3 day old moths were similar from one to another. From 0.5 h before to 4.5 h into scotophase, the titers increased slightly, but at 6.5 h they showed a significant increase up to the peak values of the scotophase. Soon at 8.5 h into the scotophase, they decreased significantly and thereafter gradually to undetectable level at 4.5 h into the next photophase. The daily change experiment showed that BAW females began calling at 0 scotophase, became fully active in calling at 3rd scotophase, and maintained the calling activity to 7th scotophase. There was no significant difference in pheromone titers among different day‐old moths.     

Tidal,Daily, and Lunar‐Day Activity Cycles in the Marine Polychaete Nereis virens

The burrow emergence activity of the wild caught ragworm Nereis virens Sars associated with food prospecting was investigated under various photoperiodic (LD) and simulated tidal cycles (STC) using a laboratory based actograph. Just over half (57%) of the animals under LD with STC displayed significant tidal (～12.4 h) and/or lunar‐day (～24.8 h) activity patterns. Under constant light (LL) plus a STC, 25% of all animals were tidal, while one animal responded with a circadian (24.2 h) activity rhythm suggestive of cross‐modal entrainment where the environmental stimulus of one period entrains rhythmic behavior of a different period. All peaks of activity under a STC, apart from that of the individual cross‐modal entrainment case, coincided with the period of tank flooding. Under only LD without a STC, 49% of the animals showed nocturnal (～24 h) activity. When animals were maintained under free‐running LL conditions, 15% displayed significant rhythmicity with circatidal and circadian/circalunidian periodicities. Although activity cycles in N. virens at the population level are robust, at the individual level they are particularly labile, suggesting complex biological clock‐control with multiple clock output pathways.     

FORAGING AND SPAWNING RHYTHMS OF THE PULMONATE LIMPET SIPHONARIA SIRIUS (PILSBRY): SWITCHING OF ACTIVITY PERIOD BY A DIURNAL FORAGER

Foraging and spawning rhythms of the intertidal pulmonate limpetSiphonaria sirius were studied at Shirahama, Wakayama Prefecture,Japan. Continuous 24 h observations were made five times, andtwo series of daytimeobservations were conducted for over 10days almost successively. This species inhabited the lower intertidalzones and had a diurnal habit in spring to autumn. At springtide most marked individuals showed foraging activity when awashand submerged both in ebb and flood tides before and after thetime of lowest water. When the tide progressed towards neaps,the activity peaks became ca. 30 min later per day and smaller.A new activity peak appeared early in the morning at neap tidein waxinghalf moon, and in summer, many adult limpets copulatedduring the morning activity. Simultaneous spawning by such limpetsoccurred one or two days after the peak of copulation. S. siriusswitched the activity period from late evening back to earlymorning at neap tide. The adaptive significances of such activitypatterns are discussed focusing on the responses against daytimedesiccation and nocturnal aquatic predators. (Received 5 September 1994; accepted 30 October 1994)     

The effects of different light–dark cycles on the metabolism of the diazotrophic,unicellular cyanobacteria Cyanothece sp. ATCC 51142, and Cyanothecesp. PCC 7822

The diazotrophic unicellular cyanobacterium Cyanothece sp. ATCC 51142 demonstrates circadian patterns in nitrogenase activity, H₂ production and glycogen storage when grown under nitrogen‐fixing, 12:12 light:dark (L:D) conditions. In this study, we grew Cyanothece sp. ATCC 51142, and another strain in this genus, Cyanothece sp. PCC 7822, under long‐day (16:8 L:D) and short‐day (8:16 L:D) nitrogen‐fixing conditions to determine if they continued to display circadian rhythms. Both strains demonstrated similar circadian patterns for all three metabolic parameters when grown under long‐day conditions. However, the strains responded differently to short‐day growth conditions. Cyanothece sp. ATCC 51142 retained reasonable circadian patterns under 8:16 L:D conditions, whereas Cyanothece sp. PCC 7822 had quite damped patterns without a clear circadian pattern. In particular, glycogen storage changed very little throughout the day and we ascribe this to the difference in the type of glycogen granules in Cyanothece sp. PCC 7822 which has small β‐granules, compared to the large, starch‐like granules in Cyanothece sp. ATCC 51142. The results suggested that both mechanistic and regulatory processes play a role in establishing the basis for these metabolic oscillations.     

Rhythmic cortisol secretion in the equine: Analysis and physiological mechanisms

Abstract

Four Thoroughbred mares (no. 1–4) were maintained under constant temperature (24°C) and controlled light (L/D:12/12 with lights on at 06.00 hr) conditions. They were fed and watered ad libitum with fresh feed and water given at 09.00 hr. After a 45‐day pre‐conditioning period, blood samples were obtained by veinipuncture at 4‐hr intervals for 14 days to determine circadian and day‐to‐day variation. The horses exhibited a circadian rhythm with maximum values attained at about 12.00 hr, however, there are periods of days in which no rhythm is distinguishable. Ultradian rhythms with mean periods of 105 to 128 and 24 to 31 min are superimposed upon the circadian rhythm. The individual rhythms are quite variable from horse to horse and within the same horse. During periods of decline in plasma cortisol with metabolic half‐lives of approximately 70 min, secretion of cortisol was very low or had ceased. During periods of increasing plasma concentration, secretion was occurring at a faster rate than degradation. Rapid decreases in plasma concentration (metabolic half‐life of approximately 30 min) was accompanied by a rise in specific activity indicating cortisol with a high specific activity was entering the plasma pool from other storage pools.     

Social stimulation as a circadian synchronizer in Periplaneta americana

Abstract

Social stimulation is not usually considered to be a circadian synchronizer of activity in insects. We investigated whether it could have any effect on synchronization and resetting capacity in insects with cauterized eyes. Direct continuous observations were carried out in groups of Periplaneta americana composed of two operated animals and eight controls. When night fell at the same time as before surgery, sexual and locomotor activity of operated animals remained synchronized with the activity of control animals. But when nightfall was postponed by 6 h, operated animals were unable to lengthen the activity peak, as long as the activity threshold stayed at its minimum level. Social stimulation resulted in modifications of and interactions with the temporal activity distribution within the threshold limits, but was unable to reset the circadian clock; so it may be questioned whether it should be regarded as a circadian synchronizer in a strict sense. Circadian rhythm of food intake disappeared in operated animals; this unstable character which has already been mentioned by other authors, even in intact animals, is discussed.     

Variation of the Locomotor Activity Rhythms in Three Species of Talitrid Amphipods,Talitrus saltator,Orchestia montagui,and O. gammarellus,from Various Habitats

The expression of biological rhythms was investigated in five populations of three different species of talitrid amphipods from various habitats in the Maremma Regional Park, Grosseto, Italy: Talitrus saltator (from a sandy beach and a canal), Orchestia montagui (form a Posidonia banquette), and O. gammarellus (from a cave entrance and a river bank). Locomotor activity rhythms were recorded in individual animals over 21 days in constant dark at a temperature of 18°±1°C. A high variability in rhythm expression was evident, not only among species and populations but also within populations and the activity pattern of individuals. Activity rhythms of T. saltator and O. montagui were similar, with a good definition and precise circadian periodicity, whereas O. gammarellus showed a high variability and low definition of the circadian rhythm. Significant differences were also observed between two populations of O. gammarellus and T. saltator from different habitats. Within the O. gammarellus species, a significantly higher percentage of active animals (p<0.001) was observed in the cave than the river‐bank population; within T. saltator, a significantly lower percentage of active animals (p<0.01) and higher percentage of periodic animals (p<0.05) was found in the canal than the sandy beach population. With reference to environmental stability and variability, the differences observed are explained as a need for plasticity to adapt to environmental changes.     

Phenology and flight periodicity of Sirex noctilio (Hymenoptera: Siricidae) in central New York,U.S.A.

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Daily Torpor Alters Multiple Gene Expression in the Suprachiasmatic Nucleus and Pineal Gland of the Djungarian Hamster (Phodopus sungorus)

Circadian rhythms are still expressed in animals that display daily torpor, implying a temperature compensation of the pacemaker. Nevertheless, it remains unclear how the clock works in hypothermic states and whether torpor itself, as a temperature pulse, affects the circadian system. To reveal changes in the clockwork during torpor, we compared clock gene and neuropeptide expression by in situ hybridization in the suprachiasmatic nucleus (SCN) and pineal gland of normothermic and torpid Djungarian hamsters (Phodopus sungorus). Animals from light‐dark (LD) 8∶16 were sacrificed at 8 time points throughout 24 h. To investigate the effect of a previous torpor episode on the clock, we sacrificed a group of normothermic hamsters 1 day after torpor. In normothermic animals, Per1 peaked at zeitgeber time (ZT)4; whereas, Bmal1 reached maximal expression between ZT16 and ZT19. AVP mRNA in the SCN showed highest levels at ZT7. On the day of torpor, the levels of all mRNAs investigated, except for AVP mRNA, were increased during the torpor bout. Moreover, the Bmal1 rhythm was advanced. On the day after the hypothermia, Bmal1 and AVP rhythms showed severely depressed amplitude. Those distinct amplitude changes of Bmal1 and AVP on the day after a torpor episode expression suggests that torpor affects the circadian system, probably by altered translational processes that might lead to a modified protein feedback on gene expression. In the pineal gland, an important clock output, Aanat expression, peaked between ZT16 and ZT22 in normothermic animals. Aanat levels were significantly advanced on the day of hypothermia, an effect which was still visible 1 day afterward. In summary, this study showed that daily torpor affects the phase and amplitude of rhythmic clock gene and clock‐controlled gene expression in the SCN. Furthermore, the rhythmic gene expression in a peripheral oscillator, the pineal gland, is also affected.     

ASSOCIATION OF THE ANTIDIABETIC EFFECTS OF BROMOCRIPTINE WITH A SHIFT IN THE DAILY RHYTHM OF MONOAMINE METABOLISM WITHIN THE SUPRACHIASMATIC NUCLEI OF THE SYRIAN HAMSTER

Bromocriptine, a dopamine D₂ agonist, inhibits seasonal fattening and improves seasonal insulin resistance in Syrian hamsters. Alterations in daily rhythms of neuroendocrine activities are involved in the regulation of seasonal metabolic changes. Changes in circadian neuroendocrine activities that regulate metabolism are believed to be modulated by central circadian oscillators within the hypothalamic suprachiasmatic nuclei (SCN) of seasonal animals. We examined the association of metabolic responses to bromocriptine with its effects on the daily rhythms of metabolic hormones and daily monoamine profiles within the SCN, a primary circadian pacemaker known to regulate metabolism, in Syrian hamsters. Obese glucose-intolerant male Syrian hamsters (body weight [BW] 185 ± 10 g) held on 14h daily photoperiods were treated at light onset with bromocriptine (800 μg/animal/day, ip) or vehicle for 2 weeks. Animals were then subjected to a glucose tolerance test (GTT) (3 g/kg BW, ip). Different subsets of animals (n = 6) from each treatment group were sacrificed at 0h/24h, 5h, 10h, 15h, or 20h after light onset for analyses of SCN monoamines, plasma insulin, prolactin, cortisol, thyroxin (T₄), triiodothyronine (T₃), glucose, and free fatty acids (FFAs). Compared with control values, bromocriptine treatment significantly reduced weight gain (14.9 vs. ?2.9 g, p <. 01) and the areas under the GTT glucose and insulin curves by 29% and 48%, respectively (p <. 05). Basal plasma insulin concentration was markedly reduced throughout the day in bromocriptine-treated animals without influencing plasma glucose levels. Bromocriptine reduced the daily peak in FFA by 26% during the late light span(p <. 05). Bromocriptine significantly shifted the daily plasma cortisol peak from the early dark to the light period of the day, reduced the plasma prolactin (mean 1.8 vs. 39.4 ng/dL) and T₄ throughout the day (mean 1.6 vs. 3.8 μg/dL), and selectively reduced T₃ during the dark period of the day (p <. 01). Concurrently, bromocriptine treatment significantly reduced SCN dopamine turnover during the light period and shifted daily peaks of SCN serotonin and 5-hydroxy-indoleacetic acid (5-HIAA) content by 12h from the light to the dark period of the day (p <. 05). This was confirmed by a further in vivo microdialysis study in which bromocriptine increased SCN extracellular 5-HIAA of glucose-intolerant hamsters during the dark phase (47% increase, p <. 05) toward levels observed in normal glucose-tolerant hamsters. Thus, bromocriptine-induced resetting of daily patterns of SCN neurotransmitter metabolism is associated with the effects of bromocriptine on attenuation of the obese insulin-resistant and glucose-intolerant condition. A large body of corroborating evidence suggests that such bromocriptine-induced changes in SCN monoamine metabolism may be functional in its effects on metabolism. (Chronobiology International, 17(2), 155–172, 2000)     

Activity rhythms and cyclical changes of microhabitat preferences in the intertidal pipefish Nerophis lumbriciformis (Pisces: Syngnathidae)

As an inhabitant of the intertidal zone, Nerophis lumbriciformis predictably should show a clear rhythmic activity, with its maximum closely correlated with the high tide period, as observed in several other rocky intertidal fish species. We investigated the rhythmic activity patterns of N. lumbriciformis and analysed to what extent specific substratum preferences may be linked to water level and, ultimately, to the particular activity rhythm patterns of the fish. To determine N. lumbriciformis substratum preferences, two different experiments were conducted. The first evaluated substratum preferences from among a group of the four most common substrates found in the sample area. The second experiment evaluated the fish's maintenance of that substratum preference, after alteration of the water level in a simulated ebb tide. From the available substrata, N. lumbriciformis displayed a clear preference for algae until the water level began to decrease. Then, the preference radically shifted from algae to boulders. N. lumbriciformis showed a clear rhythmic activity, affected by both tidal and circadian periodicity, producing asymmetrical activity peaks. Unlike the rhythms of other rocky intertidal fish species, the maximum activity peaks were not centred at the expected high tide period. The adaptive explanation for the apparently peculiar activity peaks appears to be related to the particular substratum preferences of N. lumbriciformis. The detected circatidal period seems, consequently, closely related to small migrations between substrata, whilst the observed increase in diurnal activity, independent of the tidal cycle, may be related to predatory activity, since N. lumbriciformis is a visual feeder. Electronic Publication     

Two Oscillators Might Control the Locomotor Activity Rhythm of the High‐Altitude Himalayan Strain of Drosophila Helvetica

The properties of the pacemaker controlling the adult locomotor activity rhythm of the high‐altitude Himalayan (haH) strain (Hemkund Sahib, 4121 m above sea level) of Drosophila helvetica are strikingly different from those of the low‐altitude Himalayan (laH) strain (Birahi, 1132 m above sea level) of the same species. The haH strain has a unimodal activity pattern with a delayed peak occurring about 4.5 h after lights‐on of the entraining light‐dark (LD) cycle, while the laH strain has a bimodal activity pattern with the morning and evening peaks. It is rather unusual for a wild type strain of any Drosophila species to have a unimodal activity pattern during entrainment as observed in the haH strain. The single activity peak of the haH strain is regarded as a consequence of delayed morning peak merging with the evening one. Three experiments were performed to test this hypothesis. The first experiment examined whether the single activity peak could be dissociated into two components by LD cycles in which photoperiods varied from 10 to 16 h per 24 h. The haH strain again exhibited a unimodal activity pattern with a delayed peak in 10, 12, and 14 h photoperiods but a bimodal activity pattern in 16 h photoperiod. The laH strain had bimodality in 10 and 12 h photoperiods, unimodality in a 14 h photoperiod, but complete arrhythmicity in a 16 h photoperiod.

In the second experiment, the haH flies were transferred from LD 16∶8 to LL at 5 lux to confirm whether the bimodality of this strain in LD 16∶8 cycles was not the result of masking by the long photoperiod of 16 h. Bimodality of the haH strain persisted in LL too; moreover, the morning component free‐ran with period (τ) <24 h, while the evening component free‐ran with τ>24 h. The third experiment examined the LL‐induced splitting of activity peak of the haH strain. Flies were transferred from LD 12∶12 cycles to LL at 0, 1, 5, and 15 lux. The haH strain was rhythmic in LL at 0 and 1 lux with a unimodal activity pattern. It was also rhythmic in LL at 5 lux, but the single activity peak was split into two discrete components; the morning component free‐ran with τ<24 h, while the evening component free‐ran with τ>24 h. This strain, however, was completely arrhythmic in LL at 15 lux. The laH strain was uniformly arrhythmic in LL at all levels of light intensity. These results suggest that the single but late activity component of the haH strain during entrainment appears to be the consequence of merging the delayed morning peak with the evening one as an adaptation to the environmental conditions at the altitude of origin of this strain, where these flies begin activity in the forenoon owing to non‐permissible low temperature in the morning.