Zeitgebers for animals in the continuous daylight of high arctic summer

Summary To find out which of daily oscillating environmental factors in the continuous daylight of arctic summer are responsable for synchronization of animal activity period with the rotation of the earth, in Spitsbergen (78° N) a whole summer continuous measurements of temperature (°C), light intensity (lux) and spectral composition of the light (colour temperature in °K) were done. With a daily amplitude of 4–5° C in the air, resp. 10° C at the ground surface, temperature can only act as a Zeitgeber for heterothermic animals. Measurements of light intensity (with a photocell directed vertical upward to the zenith) showed relatively large daily oscillations. But since light intensity depends on the cosine of the angle of incidence of the light (Lambert's cosine law) and in high latitudes the angle of incidence of the sun is always very great, in fact measurements have to be done with the photocell directed to the sun or, as the latter meets with mechanical problems, all dates got with the photocell vertical upward must be converted after Lambert's cosine law. Converted dates of light intensity oscillate not nearly as intense as measured dates. This small daily amplitude of light intensity, which in addition is strongly influenced by sudden changes of the wheather, can hardly come into consideration as a Zeitgeber of animal activity period. Colour temperature however showed large and regular daily oscillations and therefore could be a Zeitgeber of animal activity period in high arctic summer, particularly because in the laboratory the synchroniing effect of an artificial daily alternation of colour temperature is already proved. Nest (feeding) activity of local snowbuntings and perch hopping activity of greenfinches, brought from Germany, were registrated with light traps. Activity period of both bird species was clearly synchronized with the rotation of the earth. In a narrow valley the activity pattern of greenfinches was identical as in open country, in spite of temporarily total other conditions of light intensity (shading by mountains). Apparently even the activity period of non arctic animals can be synchronized by weak Zeitgebers like colour temperature.     

The importance of illumination level in the daily onset of flight activity in nocturnal moths

ABSTRACT. The daily onset of flight activity in nocturnal moths from Denmark and Florida was investigated in natural and experimental reductions of illumination. The daily cycle of illumination is described, especially the variation of the rate of change during the evening. Different species commence activity ('onset') at different mean light intensities. These means occur around the termination of twilight, at between 0 and –2.50 log lux. In a given species the illumination at mean onset is constant, and the dispersion of onsets in the population is constant with respect to the illumination. As a consequence, the dispersion with respect to time increases, the longer the duration of twilight. The temporal dispersion, therefore, depends upon season and latitude. A model describing the number of onsets in relation to the decrease of illumination is presented. At the physiological level it is proposed that a preparatory process precedes the actual onset of activity. This is observed as a latency between the light change and the activity onset. It is presumed to represent a change, from an inhibiting to a disinhibiting output from a controlling CNS centre, brought about by the reduction in illumination increasing the sensitivity of the centre to a 'sensitizer'. This might represent the Zeitgeber effect of illumination reduction. The results indicate that during a gradual decrease of illumination, the latency process is initiated by a certain low illumination and that it proceeds further, the lower the illumination. The faster the rate of decrease, the faster is the rate of the process, so that it is completed at about the same time as a threshold illumination for overt activity is reached, irrespective of the rate of decrease. However, at very fast reductions of illumination, the physiological process cannot keep pace with the reduction, and activity onset occurs at a lower illumination than normal.     

Daily biphasic behaviour in keratin-feeding desert trogid beetles in relation to climate

Abstract. 1. Patterns of daily activity of two ectothermic species of the trogid genus Omorgus were studied in the Kalahari of southern Africa in summer. The relationship between time and type of activity, and environmental temperature, relative humidity and light intensity, was investigated.
2. Beetle activity was biphasic, with peaks in activity at sunset and sunrise in both species.
3. Afternoon activity coincided with high temperatures and low humidity and morning activity with minimum daily temperature and maximum humidity.
4. Afternoon activity, which peaked at about 34°C and 25% r.h., consisted predominantly of complex breeding behaviour and dispersal. High temperatures are necessary for these activities.
5. In the morning feeding predominated. At this time temperature was about 27°C and r.h. above 60%. High r.h. is important because it restricts respiratory loss of water, improves food (moist hair and skin keratin) quality and may compensate for faecal water loss.
6. No evidence of competition between the two species was found.     

Weather-induced changes in moth activity bias measurement of long-term population dynamics from light trap samples

Interpretation of light trap catches of moths is complicated by daily variation in weather that alters flight activity and numbers caught. Light trap efficiency is also modified by wind and fog, and daily weather may effect absolute abundance (numbers actually present). However, actograph experiments and other sampling methods suggest that changes in daily activity are large by comparison to changes in absolute abundance. Daily variation in weather (other than wind and fog) is therefore a form of sampling error in absolute abundance estimates. We investigated the extent of this sampling bias in 26 years of population dynamics from 133 moth species. In a subset of 20 noctuid and geometrid species, daily numbers caught were positively correlated with temperature in 14 species, and negatively correlated with rainfall in 11 species. The strength of correlations varied between species, making it difficult to standardize catches to constant conditions. We overcame this by establishing how weather variation changed with time and duration of the flight period. Species flying later in the summer and for shorter periods experienced more variable temperatures, making sampling error greater for these species. Of the 133 moth species, those with shorter flight periods had greater population variability and more showed significant temporal density dependence. However, these effects were weak, which is encouraging because it suggests that population analyses of light trap data largely reflect factors other than sampling error.     

Daily rhythms of oxygen consumption, body temperature, activity and melatonin in the Norwegian lemming Lemmus lemmus under northern summer photoperiod

Rodents inhabiting high latitudes, close to the Arctic Circle or above it, are exposed to near 24 h daylight during the summer season. An example to such rodent species is the Norwegian lemming Lemmus lemmus, which is distributed in northern Fennoscandia. We measured daily rhythms of heat production (VO₂), body temperature (T_b), motor activity and melatonin secretion (measured from its metabolite 6-sulfatoximelatonin 6-SMT) in individuals exposed to natural day light, Oulu Finland, in August 1997 and at a controlled ambient temperature of 22 °C. Our results show a daily rhythm of VO₂ with an acrophase at 04:00 h and minimal values measured at 16:00 h, with a significant difference (p<0.001) between mean day and night values. 6-SMT also shows a daily rhythm with maximal secretion during the dark phase 24:00 and 06:00 h in which total 6-SMT values were 2.92±1.1 and 3.87±1.2 ng, respectively. The lowest values were recorded at 12:00 h; 0.86±0.63 ng. These results suggest that lemmings show a VO₂ and melatonin daily rhythms, which seem to correlate with each other and it appears that melatonin secretion increases heat production.     

Usefulness and limitation of a tiny light-temperature logger to monitor daily activity levels of arboreal squirrels in temperate areas

Owing to recent advances in technology, biologging has revealed in-depth animal behavior at a level of detail that previously could not be obtained. However, previous studies were limited to relatively large species due to constraints caused by the size of data loggers. Some recent studies, however, applied tiny light loggers (geolocator) to monitor daily activity levels of small rodents in the southern arctic and far north Holarctic. In this study, we tested whether a similar method can be applied to arboreal squirrels in temperate regions, by combining light intensity and temperature data. Inside the squirrel’s nest was dark enough during daylight hours to capture out of nest activity with light intensity, clearly showing daily and seasonal changes in diel activity patterns. The new temperature function on the device showed consistently high temperature within the nests (35–37 °C) and suggested additional activity before sunrise; the squirrels were primarily diurnal, but occasionally left their nest for short periods (an average of 13 min) during the night. We also found that estimates of daily activity time changed depending on the threshold cut-off values of light intensity and provide a recommendation of 5 to 25 lx for future studies. In addition, temperature loggers were not always effective during summer when warmer ambient temperature approached the squirrel’s body temperature. While the inexpensive and tiny light-temperature logger has potential to monitor activity levels of small mammals, care should be taken when analyzing the output.     

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.     

Locomotor activity and body temperature rhythms in the Mahali mole-rat (C. h. mahali): The effect of light and ambient temperature variations

African mole-rats (family: Bathyergidae) are strictly subterranean mammals that reside in extensive networks of underground tunnels. They are rarely, if ever, exposed to light and experience muted temperature ranges. Despite these constant conditions, the presence of a functional circadian clock capable of entraining to external light cues has been reported for a number of species. In this study, we examine a social mole-rat species, Cryptomys hottentotus mahali, to determine if it possesses a functional circadian clock that is capable of perceiving light and ambient temperature cycles, and can integrate these cues into circadian rhythms of locomotor activity and core body temperature. Eight male and eight female, non-reproductive individuals were subjected to six cycles of varying light and temperature regimes. The majority of the individuals displayed daily rhythms of locomotor activity and body temperature that are synchronised to the external light and temperature cycles. Furthermore, endogenous rhythms of both locomotor activity and core body temperature were displayed under constant conditions. Thus, we can conclude that C. h. mahali possesses a functional circadian clock that can integrate external light and temperature cues into circadian rhythms of locomotor activity and core-body temperature.     

Circadian mating activity and effect of pheromone pre-exposure on pheromone response rhythms in the moth Spodoptera littoralis

Mating in moths is generally mediated by female-produced sex pheromones. Mating activity, female pheromone production/release and male pheromone responsiveness all show diurnal variations in many species. We found that the response of the male Egyptian cotton leafworm, Spodoptera littoralis, to sex pheromone gland extracts showed a diel rhythm in olfactometer tests, and the variation was persistent for at least 1 day in constant darkness. High male response to sex pheromone was correlated in time with high mating and locomotor activity. Male S. littoralis, maintained in constant darkness and exposed to pheromone gland extracts on a daily basis, showed an induced temporal variation in response after several days, in contrast to unexposed males. This suggests that in the absence of other external zeitgebers, exposure to sex pheromone may function to synchronise circadian behavioural rhythms in male moths. The daily rhythm in mating activity in S. littoralis is also shown to be persistent for at least 2 days in constant darkness. Pairs mated significantly less when either the male or female had been raised in a light:dark cycle 10 h out of phase, indicating that the proposed circadian rhythm in mating activity is composed of rhythmic mating preference/ability in both sexes.     

Interplay of temperature and light influences wild guppy (Poecilia reticulata) daily reproductive activity

In this study we investigated the environmental regulation of daily reproductive activity of guppies (Poecilia reticulata). We observed male guppy reproductive behaviour for 17 days over three time periods (dawn, noon, and dusk) while recording natural variation in water temperature and light level. Water temperatures recorded during the experiments were highly variable (up to 7 °C per day), and there was a recurring pattern in daily variation for both light and temperature. Levels of activity were highest at dawn and similarly low during noon and dusk, but reproductive behaviour was recorded throughout the day. Mixed‐effects models indicate that light and temperature affect reproductive behaviour differently at different times of the day, and can also have opposing effects. We suggest that the environmental heterogeneity of streams in Trinidad has led to a broad thermal tolerance, and has contributed to the high level of phenotypic plasticity in the guppy and its success as an invasive species. Furthermore, our results show that daily variation in temperature and its interaction with light should be considered in future studies of guppy reproductive behaviour. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 511–520.     

On the occurence of a short period of delayed implantation in Schreibers'long-fingered bat (Miniopterus schreibersii) from a tropical latitude in Zimbabwe

Mating in Schrcibers'long-fingered bat ( Miniopterus schreibersii ) from central Zimbabwe (18°S) occurred between mid-April and mid-May, after a five-month period of spermatogenesis. Implantation was delayed until early July and parturition occurred between late October and mid-November, about four months after implantation and six to seven months after fertilization. The timing and length of gestation at 18°S are similar to that described for M. schreibersii from Zaire (11σS) and when these date are compared with those for the same species from higher latitudes in Africa, it is apparent that there is a small, but distinct, increase in the total length of pregnancy and the length of delayed implantation with increasing latitude. The reproductive bilogy of Miniopterus schreibersii from Africa and France differs fundamentally from that of the same species from Australia and Japan, and it is suggested that studies of the relationship between latitude and duration of delayed implantation should be based on comparisons within single species from different latitudes on the same continent, and that comparisons between species and between continents should be avoided.     

Respiratory Q10 varies between populations of two species of Myrmica ants according to the latitude of their sites

Metabolic respiration by groups of resting Myrmica ruginodis and M. scabrinodis worker ants from five sites representing a range of latitudes, have been compared by measuring rates of CO(2) production-standardised by fat-free weight-at 5 and 25 degrees C. M. ruginodis which lives in cooler habitats than M. scabrinodis consistently produced more CO(2). At 5 degrees C ants of both species from southern latitudes were metabolically more active than those from more northerly latitudes, whereas at 25 degrees C the situation was reversed. Estimates of Q10 were positively correlated with latitude indicating that the respiratory metabolism of northern populations increases relatively more in response to rising temperatures than southern populations. Values of Q10 at different latitudes were the same for both species. The results are discussed in terms of seasonal fluctuations of temperature at different latitudes.     

Effects of age, ambient temperature and reproductive status on wing beat frequency of the rice leafroller Cnaphalocrocis medinalis (Guenée) (Lepidoptera: Crambidae)

The rice leafroller Cnaphalocrocis medinalis (Guenée) is a serious insect pest of rice with migratory ability. In this work, we studied the effects of age, temperature and reproductive status on the wing beat frequency (WBF). The longevity of this moth reached 30.6 days at 16 °C, which was significantly longer than that at 26 and 32 °C (10.7 and 8.8 days, respectively). The WBF of the 1-day-old moths did not differ significantly among the three temperature treatments. However, the WBF of the 2- to 7-day-old moths differed significantly among those temperatures, with the highest WBF at 26 °C and the lowest WBF at 32 °C. Both age and temperature significantly affected the WBF. No significant differences in WBF were detected either between mated and unmated or between parous and nulliparous female moths. There were no significant differences on WBF among field-collected females on different stages of ovary development. Significant correlations were found between the WBF and the tethered flight duration in field-collected moths as well as in laboratory-reared moths. These results indicated that the WBF was intensively influenced by age and temperature, and it may be a potential index to evaluate the flight ability of this species.     

Evaluation of pheromone-baited traps for winter moth and Bruce spanworm (Lepidoptera: Geometridae)

We tested different pheromone-baited traps for surveying winter moth, Operophtera brumata (L.) (Lepidoptera: Geometridae), populations in eastern North America. We compared male catch at Pherocon 1C sticky traps with various large capacity traps and showed that Universal Moth traps with white bottoms caught more winter moths than any other trap type. We ran the experiment on Cape Cod, MA, where we caught only winter moth, and in western Massachusetts, where we caught only Bruce spanworm, Operophtera bruceata (Hulst) (Lepidoptera: Geometridae), a congener of winter moth native to North America that uses the same pheromone compound [(Z,Z,Z)-1,3,6,9-nonadecatetraene] and is difficult to distinguish from adult male winter moths. With Bruce spanworm, the Pherocon 1C sticky traps caught by far the most moths. We tested an isomer of the pheromone [(E,Z,Z)-1,3,6,9-nonadecatetraene] that previous work had suggested would inhibit captures of Bruce spanworm but not winter moths. We found that the different doses and placements of the isomer suppressed captures of both species to a similar degree. We are thus doubtful that we can use the isomer to trap winter moths without also catching Bruce spanworm. Pheromone-baited survey traps will catch both species.     

Influence of environmental factors on daily flight activity of the filbertworm, Melissopus latiferreanus (Lepidoptera, Olethreutidae)

ABSTRACT.

• 1 Flight of Melissopus latiferreanus (Walsinghzm), the filbertworm, was influenced by a number of environmental factors including temperature, wind, and rainfall.
• 2 Few M. latiferreanus moths were trapped in light or suction traps or found in sweep nets samples when air temperatures were above 31°C or below 15°C. Moth captures were optimum when prevailing temperature was between 21 and 26°C.
• 3 Typically moth flights began at about sunset and continued throughout the night with a peak at 22.00 hours, about an hour after sunset.
• 4 Very few moths were trapped under showery and gusty conditions when wind velocity was over 16km h^?1.
• 5 More moths were captured in light traps during dark nights than on full moon nights. The pattern of captures indicated that females flew earlier than males.

     

Seed dormancy in Rumex species in response to environmental factors

Abstract. Many Rumex species show similar seed dormancy characteristics but there is more information concerning R. crispus and R. obtusifolius than other species. These species respond positively to red or white light. Far-red light applied for short periods may promote or inhibit germination depending on the timing of the irradiation in relation to temperature change; but long periods of far-red inhibit germination. Seeds may also be stimulated to germinate in the dark by low-temperature stratification at 15°C or less providing the temperature of the seeds is subsequently raised to a minimum of about 15°C. Seeds can, however, germinate at lower temperatures providing they have received other appropriate stimulatory treatment. Seeds also respond to alternating temperatures. In a diurnal cycle the minimum upper temperature required is about 15°C and the maximum lower temperature is about 25°C. The optimum period spent at the upper temperature is about 8 h when it is 15–25°C but the optimum period decreases as the upper temperature is increased above this range so that at 45°C, for example, it is only about 30 min. The period spent at the lower temperature in a diurnal cycle is not critical. Providing these criteria are met, the percentage germination increases with the number and amplitude of the cycles. The warming part of the cycle is necessary for the response but so far there is no convincing evidence that cooling itself is important. Secondary dormancy is induced at constant temperatures at a rate dependent on temperature, but apparently only in the presence of oxygen. This feature affects the optimum timing of a temperature change or exposure to light. Strong positive interactions are shown between stimulatory temperature treatments and white or red light. Unlike many other weed species the seeds respond only slightly to nitrate ions. The implications of these responses are discussed in relation to field behaviour.     

Cormorants dive through the Polar night

Most seabirds are visual hunters and are thus strongly affected by light levels. Dependence on vision should be problematic for species wintering at high latitudes, as they face very low light levels for extended periods during the Polar night. We examined the foraging rhythms of male great cormorants (Phalacrocorax carbo) wintering north of the Polar circle in West Greenland, conducting the first year-round recordings of the diving activity in a seabird wintering at high latitudes. Dive depth data revealed that birds dived every day during the Arctic winter and did not adjust their foraging rhythms to varying day length. Therefore, a significant proportion of the dive bouts were conducted in the dark (less than 1 lux) during the Polar night. Our study underlines the stunning adaptability of great cormorants and raises questions about the capacity of diving birds to use non-visual cues to target fish.     

The effect of temperature on the activity of three species of Helicid snail (Mollusca: Gastropoda)

The activity of the land snails Cepaea nemoralis (L.), C. hortensis (Müll.) and Arianta arbustorum (L.) was measured over 24 hours at the constant temperatures of 0, 3, 8, 17, 22 and 26°C at 100% relative humidity and with 16 hours light and 8 hours dark. The activity of all three species varies significantly with temperature and light, and there are differences in the activities of the species at each temperature. A. arbustorum is relatively more active than Cepaea at low rather than high temperatures, as is C. hortensis with respect to C. nemoralis , though to a lesser extent. There is little diurnal rhythm in activity at 0°C, but all species become more nocturnal as temperature increases; C. nemoralis is most, and A. arbustorum least nocturnal. The results obtained assist in explaining the observed geographical and local distributions of the species concerned.