Circadian rhythm of swimming activity in juvenile pink salmon (Oncorhynchus gorbuscha)

The circadian rhythm of swimming activity and the role of the daily illumination cycle in the synchronization of this rhythm were studied in individual juvenile pink salmon. Sixty eight percent of all fish examined (n=38) were day-active when exposed to a 12 h L:12 h D cycle; the remaining fish were nocturnally active. One half of the fish tested under laboratory conditions of continuous, constant light intensity (LL) and constant temperature showed unambiguously endogenous activity rhythms with circadian periods for up to 10 d. The remaining fish were arrhythmic. Mean period length of the free-running activity rhythms for diurnal fish in LL shortened with constant light intensity increasing from 6 to 600 lx, as predicted by the circadian rule. In contrast, mean free-running period for nocturnal fish did not vary significantly with similarly increasing constant light intensity. Mean swimming speed (activity level) of both diurnal and nocturnal fish increased significantly with increasing light intensity. This is suggestive of a positive photokinetic response. When subjected to a phase-delayed LD cycle, the fish resynchronized their daily rhythms of activity with this new LD cycle after only one transient cycle in most instances. Hence, the timing of the daily activity rhythms appeared to occur through the direct masking action of the illumination cycle on activity, rather than through entrainment of an endogenous circadian system.     

Temporal patterns of foraging activity in some wood warblers in relation to the availability of insect prey

Summary I monitored the temporal pattern of diurnal feeding activity in several wood warbler (Parulidae) species and concomitantly recorded the numbers of active (flying) insects in 2 willow habitats in the western United States. At one site the temporal relationship between the density of active and inactive (nonflying) insects was investigated. The diurnal patterns of insect and bird activity were inversely related and each pattern was significantly nonuniform throughout the day; the wood warblers were largely inactive during the middle of the day when insects were most active.As foliage-gleaning birds, wood warblers depend primarily on the availability of inactive (nonflying) insects that they pick from the foliage, and they appear to be limited in their foraging activity by the unavailability of such insects during midday. Interestingly, the duration of midday inactivity for a given bird species varied inversely with the proportion of time that species spent flycatching. Thus, food availability may play an important role in determining the temporal patterns of feeding activity in these insectivorous bird species.     

Feeding over the 24-h cycle: dietary flexibility of cathemeral collared lemurs (<Emphasis Type="Italic">Eulemur collaris</Emphasis>)

Animals show specific morphological, physiological and behavioural adaptations to diurnal or nocturnal activity. Cathemeral species, i.e. animals with activities distributed over the 24-h period, have to compromise between these specific adaptations. The driving evolutionary forces and the proximate costs and benefits of cathemerality are still poorly understood. Our goal was to evaluate the role of predator avoidance, food availability and diet quality in shaping cathemeral activity of arboreal mammals using a lemur species as an example. For this, two groups of collared lemurs, Eulemur collaris, were studied for 14 months in the littoral forest of southeastern Madagascar. Data on feeding behaviour were collected during all-day and all-night follows by direct observation. A phenological transect containing 78 plant species was established and monitored every 2 weeks to evaluate food availability during the study period. Characteristics of food items and animal nutritional intake were determined via biochemical analyses. The ratio of diurnal to nocturnal feeding was used as response variable in the analyses. The effects of abiotic environmental variables were removed statistically before the analyses of the biotic variables. We found that diurnal feeding lasted longer during the hot–wet season (December–February), whereas nocturnal feeding peaked during the hot–dry and cool–wet seasons (March–August). Although the lemurs foraged mostly in lower forest strata during daylight and used emergent trees preferably at night, the variables which measured animal exposure to birds of prey failed to predict the variation of the ratio of diurnal/nocturnal feeding. Ripe fruit availability and fiber intake are the two variables which best predicted the annual variation of the lemur diurnality. The data indicate that feeding over the whole 24-h cycle is advantageous during lean periods when animals have a fibre-rich, low-quality diet.     

Endogenous swimming rhythms underlying the spawning migration of the blue crab, Callinectes sapidus: ontogeny and variation with ambient tidal regime

Spawning female blue crabs, Callinectes sapidus, use ebb-tide transport (ETT) to migrate seaward. In estuaries with semi-diurnal tides, ETT in ovigerous blue crabs is driven by a circatidal rhythm in vertical swimming in which crabs ascend into the water column during ebb tide. The ontogeny of this rhythm was examined by monitoring swimming behavior of females before the pubertal molt, females that had recently undergone the molt but had not yet produced a clutch of eggs, and ovigerous females from an estuary with strong semi-diurnal tides. To assess variation in swimming rhythms with ambient tidal regime, swimming rhythms of ovigerous females from semi-diurnal (Beaufort, North Carolina), diurnal (St. Andrew Bay, Florida), and non-tidal (South River, North Carolina) estuaries were compared. Experiments were conducted during the summers of 2006–2008. Female crabs prior to oviposition had variable endogenous swimming rhythms (circadian, circatidal, or circalunidian). Ovigerous females from estuaries with semi-diurnal and diurnal tides had pronounced circatidal or circalunidian rhythms with swimming during the time of ambient ebb tide. Swimming rhythms of several ovigerous crabs switched between circatidal and circalunidian during the ~5-day observation period. Ovigerous crabs from a non-tidal estuary had a circadian rhythm with vertical swimming around the time of sunset. These results suggest that, while endogenous swimming rhythms are present in some female blue crabs prior to oviposition, rapid seaward movement via ETT in tidal estuaries begins following oviposition of the first clutch of eggs.     

Optimal daily routines of singing and foraging in a bird singing to attract a mate

Summary In this paper we develop a dynamic programming model to explore the optimal organization of daily routines of singing and foraging in a small bird. While singing the bird may attract a mate but uses up energy. Most of the patterns of daily variation in singing generated have basic features very characteristic of typical passerine song output. The predictions are remarkably robust to changes in a wide range of parameters, showing which parameters are important. A peak of singing at dawn can result from variability in overnight energy expenditure in the absence of any circadian patterns in the environment.     

Influences of moonlight,ambient temperature,and food availability on the diurnal and nocturnal activity of owl monkeys (<Emphasis Type="Italic">Aotus azarai</Emphasis>)

The study of activity rhythms, their potential zeitgebers and masking factors among free-ranging primates has received relatively little attention in the past. Most primates are diurnal, a few of them nocturnal, and even fewer are cathemeral. Owl monkeys (Aotus azarai azarai) regularly show diurnal, as well as nocturnal, activity in the Argentinean and Paraguayan Chaco. The goal of this study was to examine how changes in activity patterns in owl monkeys of Formosa, Argentina are related to daily, monthly, and seasonal changes in temperature, light and food availability . During 1 year, I collected activity data from five groups followed continuously from dawn to dusk, dusk to dawn or uninterruptedly during 24 or 36 h for approximately 1,500 h. I kept hourly and daily records of temperature and light conditions, and I gathered monthly information on the density, distribution and abundance of food resources available to the monkeys. I found that the area of study is highly seasonal, and characterized by significant fluctuations in rainfall, temperature, photoperiod, and food availability. Owl monkeys had on average 5 h of activity during the day and 4 h during the night. The amount of diurnal activity remained fairly constant through the year despite seasonal changes in exogenous factors. Owl monkeys did not show changes in their activity patterns that could be attributed to changes in food availability. Nocturnal activity increased as the amount of moonlight increased, whereas diurnal activity decreased following a full-moon night. Ambient temperature was a good predictor of activity only when the moon was full. These results argue convincingly for an interaction between ambient temperature and moonlight in determining the observed activity pattern. It is then highly advisable that any evaluation of diurnal activity in cathemeral animals be analyzed controlling for the possible effects of moonlight during the previous night.Communicated by P. Kappeler     

Social dominance modifies behavioral rhythm in a queenless ant

Social insects provide an intriguing model system in chronobiology. Typically, an egg-laying queen exhibits arrhythmicity in activity while foraging worker has clear rhythmicity. In the queenless ant, Diacamma sp., from Japan, colony members lack morphological caste, and reproductive differentiation occurs as a consequence of dominance hierarchy formation. Their specialized dominance interaction “gemmae mutilation”, provide us a fascinating model system to investigate the effect of social dominance on rhythmic ontogeny. Measurement of individual rhythms revealed that they have clear circadian rhythm at eclosion but it is diminished by social mutilation of gemmae. Moreover, unlike highly eusocial species, mated egg-layer (i.e., gamergate) possessed a circadian rhythm even after mating in Diacamma. Measurement of colony-level rhythms revealed that gemmae mutilations are performed in the limited time of the day, but foraging occurs around-the-clock. The above finding is a novel form of temporal organization in social insects, providing a new insight in morphologically casteless species. We discuss the causes and consequences of rhythmic variability in social organization.     

Food abundance affects energy intake and reproduction in frugivorous female Assamese macaques

In most mammals, female fertility and reproduction are strongly influenced by nutritional status and, therefore, by foraging conditions. Here, we investigate the relationship between food resources, feeding competition, energy intake and reproduction in a group of wild female Assamese macaques (Macaca assamensis) in northeastern Thailand. Over 2,100 h of data on feeding behaviour, energy intake and activity budgets were combined with data on resource characteristics, female reproduction and physical condition. We found that an increase in food availability had a positive effect on female energy intake and conception rates. In addition, it appeared that females incurred energetic costs during lactation and that females with a better physical condition during the mating season were more likely to conceive. The annual birth season occurred a few months before the annual peak in food availability, causing peak lactation to coincide with a period of high food availability. This suggests that females use the increased food abundance to compensate for the energetic costs of lactation. Neither energy intake rates nor activity budgets were influenced by female dominance rank, even during periods when the levels of contest competition were predicted to be high. In line with this, we found no evidence for rank-related differences in reproduction. The apparently limited influence of feeding competition in female Assamese macaques adds to the debate on the extent to which patterns in feeding competition and fitness can reliably be predicted based on ecological conditions. We suggest that this may partially be resolved by including potential competition-reducing mechanisms into the predictive framework.     

Pattern-oriented modeling of bird foraging and pest control in coffee farms

We develop a model of how land use and habitat diversity affect migratory bird populations and their ability to suppress an insect pest on Jamaican coffee farms. Bird foraging—choosing which habitat patch and prey to use as prey abundance changes over space and time—is the key process driving this system. Following the “pattern-oriented” modeling strategy, we identified nine observed patterns that characterize the real system's dynamics. The model was designed so that these patterns could potentially emerge from it. The resulting model is individual-based, has fine spatial and temporal resolutions, represents very simply the supply of the pest insect and other arthropod food in six habitat types, and includes foraging habitat selection as the only adaptive behavior of birds. Although there is an extensive heritage of bird foraging theory in ecology, most of it addresses only the individual level and is too simple for our context. We used pattern-oriented modeling to develop and test foraging theory for this across-scale problem: rules for individual bird foraging that cause the model to reproduce a variety of patterns observed at the system level. Four alternative foraging theories were contrasted by how well they caused the model to reproduce the nine characteristic patterns. Four of these patterns were clearly reproduced with the “null” theory that birds select habitat randomly. A version of classical theory in which birds stay in a patch until food is depleted to some threshold caused the model to reproduce five patterns; this theory caused lower, not higher, use of habitat experiencing an outbreak of prey insects. Assuming that birds select the nearby patch providing highest intake rate caused the model to reproduce all but one pattern, whereas assuming birds select the highest-intake patch over a large radius produced an unrealistic distribution of movement distances. The pattern reproduced under none of the theories, a negative relation between bird density and distance to trees, appears to result from a process not in the model: birds return to trees at night to roost. We conclude that a foraging model for small insectivorous birds in diverse habitat should assume birds can sense higher food supply but over short, not long, distances.     

Locomotor activity patterns and feeding habits in the prawn Palaemon xiphias (Crustacea: Decapoda: Palaemonidae) in Alfacs Bay,Ebro Delta (northwest Mediterranean)

Diel activity rhythms of the prawn Palaemon xiphias Risso, 1818 from Alfacs Bay, Ebro Delta, were studied by time-lapse videorecordings. Activity displayed an endogenous circadian rhythm, with maximum activity at night. Feeding habits were studied by frequency of occurrence and by the points method. The food of P. xiphias consisted mainly of crustaceans; the remains of amphipods, isopods, mysids, copepods, decapods and ostracods were identified. The remaining items consisted of molluscs, polychaetes, ophiuroids, plant material, sand, and unidentified organic debris. The results indicate that P. xiphias is a predator of benthic invertebrates rather than a scavenger or detritus feeder. Diet composition changes with increasing size of the prawn.     

Self-organization of circadian rhythms in groups of honeybees (Apis mellifera L.)

Workers in social groups of honeybees (Apis mellifera L.) synchronize their individual free-running circadian rhythms to an overall group rhythm. By monitoring the activity of bees by recording the oxygen consumption and intragroup temperature, it is shown that the rhythm coordination is in part achieved by temperature fluctuations as an intragroup Zeitgeber. Trophallaxis was shown to have only a minor (if any) effect on circadian rhythm synchronization. A model incorporating a feed back loop between temperature and activity can plausibly explain the observed synchronization of individual rhythms in social groups as a self-organization phenomenon. Correspondence to: R.F.A. Moritz     

Endogenous circatidal rhythm in the Manila clam Ruditapes philippinarum (Bivalvia: Veneridae)

Manila clams, Ruditapes philippinarum, removed from their natural environment and maintained for 9 weeks in continuously immersed conditions exhibited a clear endogenous circatidal rhythm in oxygen consumption. The clams exhibited a semidiurnal rhythmicity in oxygen consumption after showing a diurnal pattern in the first few days (5 to 7 d) of the experiment. The results of the present study indicate that activity rhythms of clams are controlled not only by exogenous factors, but also by an endogenous circatidal periodicity. Received: 8 April 1998 / Accepted: 1 February 1999     

Environmental influence on the locomotor activity rhythm of Talitrus saltator (Crustacea: Amphipoda)

The persistence and precision of the endogenous, nocturnal, circadian locomotor rhythm of the sand-beach amphipod Talitrus saltator is characterised, and the influence of substrate availability, photoperiod, temperature and relative humidity as potential environmental synchronizers of the rhythm is assessed. Individual, cyclic light-dark and temperature regimes synchronize and modify the laboratory activity pattern, but substrate availability and relative humidity have no significant long-term effect. Under simulateneous experimental regimes simulating field conditions of photoperiod, temperature and relative humidity the rhythm is entrained solely by photoperiod. The implications are discussed in relation to field behaviour.     

Arrived in the light: diel and seasonal activity patterns in wild Verreaux’s sifakas (<Emphasis Type="Italic">Propithecus v. verreauxi</Emphasis>; Primates: Indriidae)

Madagascar is characterised by pronounced annual climatic and ecological seasonality and harbours a radiation of closely related sympatric primates (Lemuriformes) that exhibit diurnal, nocturnal and cathemeral activity patterns. We collected continuous activity data over a complete annual cycle from wild diurnal Verreauxs sifakas (Propithecus v. verreauxi) to contribute detailed and comparative data: (1) to characterise their diel and seasonal activity rhythms, (2) to identify factors shaping variation in activity rhythms, and (3) to help reconstruct the evolutionary transition from nocturnal to diurnal activity. We fitted eight adult sifakas from seven different groups living in Kirindy forest with an accelerometer/data logger device and recorded their activity in 5-min bins for a total of 12 months. We characterise P. verreauxi as a strictly diurnal species with a pronounced bimodal activity pattern that briefly changed to a more unimodal pattern during their annual mating season (January to March). We documented significant annual variation in total daily activity, activity time, and activity level, as well as in most parameters characterising their bimodal activity pattern. Despite a significant positive correlation of the animals activity time with day length, pronounced annual variation in the phase position of onset and end of activity in relation to sunrise and sunset times could also be discerned. Minor enhancing effects of moonlight on nocturnal activity were only found for the first 3 h of the inactivity period. Bimodality of the activity pattern and the additional reduction of activity time during the cold and dry winter months associated with reduced food availability can be interpreted as flexible behavioural adaptations to reduce energy expenditure. We therefore propose that energetic and thermoregulatory benefits are important factors shaping these primates activity pattern.Communicated by C. Nunn     

Diel vertical migration of the marine copepod<Emphasis Type="Italic"> Calanopia americana</Emphasis>. II. Proximate role of exogenous light cues and endogenous rhythms

The marine copepod Calanopia americana Dahl undergoes twilight diel vertical migration (DVM) in the Newport River estuary, North Carolina, USA, in synchrony with the light:dark cycle. Copepods ascend to the surface at sunset, descend to the bottom around midnight, and make a second ascent and descent before sunrise. Behavioral assays with C. americana in the laboratory during fall 2002/2003 and summer 2004 investigated aspects of three hypotheses for the proximate role of light in DVM: (1) preferendum hypothesis (absolute irradiance), (2) rate of change hypothesis (relative rates of irradiance change), and (3) endogenous rhythm hypothesis. Results suggest that C. americana responds to exogenous light cues consistent with its DVM pattern; changes in absolute irradiance evoked swimming responses that would result in an ascent at sunset and descent at sunrise, while relative rates of irradiance decrease at sunset (–0.0046 s^–1) evoked an ascent response, and relative rates of irradiance increase at sunrise (0.0042 s^–1) evoked a descent response. Furthermore, C. americana expressed an endogenous rhythm in vertical migration that was positively correlated with field observations of twilight DVM. Collectively, these results indicate that both exogenous light cues and endogenous rhythms play a proximate role in twilight DVM of C. americana, providing redundancy in the causes of its vertical migration.Communicated by J.P. Grassle, New Brunswick     

Timekeeping in the honey bee colony: integration of circadian rhythms and division of labor

The daily patterns of task performance in honey bee colonies during behavioral development were studied to determine the role of circadian rhythmicity in age-related division of labor. Although it is well known that foragers exhibit robust circadian patterns of activity in both field and laboratory settings, we report that many in-hive tasks are not allocated according to a daily rhythm but rather are performed 24 h per day. Around-the-clock activity at the colony level is accomplished through the performance of some tasks by individual workers randomly with respect to time of day. Bees are initially arrhythmic with respect to task performance but develop diel rhythmicity, by increasing the occurrence of inactivity at night, prior to becoming foragers. There are genotypic differences for age at onset of rhythmicity and our results suggest that these differences are correlated with genotypic variation in rate of behavioral development: genotypes of bees that progressed through the age polyethism schedule faster also acquired behavioral rhythmicity at an earlier age. The ontogeny of circadian rhythmicity in honey bee workers ensures that essential in-hive behaviors are performed around the clock but also allows the circadian clock to be engaged before the onset of foraging. Received: 6 October 1997 / Accepted after revision: 28 March 1998     

The role of the queen in circadian rhythms of honeybees (Apis mellifera L.)

Summary Colonies and smaller social groups of honeybees (Apis mellifera carnica L.) show distinct free-running circadian rhythms similar to that of individual organisms. The workers of a colony synchronize their individual rhythms to one overall group rhythm. Caste plays an important role in this synchronization process. Queens were introduced into worker groups which were entrained to a phase-shifted light/dark cycle. The introduction of the queen caused a shift in the free-running phase under constant dark conditions. Single introduced workers had no effect on the free-running rhythms. This indicates that the queen plays an important role in the synchronization of circadian rhythms of honeybee colonies.     

Feeding activity rhythm of<Emphasis Type="Italic"> Nephrops norvegicus</Emphasis> of the western Mediterranean shelf and slope grounds

Over a 24-h time scale, benthic decapods show complex patterns of behavioural activity associated with feeding. Information is scarce on the feeding rhythms of the burrowing decapod Nephrops norvegicus (L.), especially in those deep, benthic environments typical of western Mediterranean slopes, where the fishery of this species is highly developed. In the present study, the feeding of this species was determined by means of stomach sampling, from animals captured during four continuous 24-h cycles of trawl hauls at 100–110 m and 400–430 m, in October (the autumn equinox) and June (the spring solstice). A total of 3348 stomachs were analysed for fullness and classified by means of a visual, qualitative scale. A high proportion of the stomachs collected was empty (47.2%). Feeding rhythmicity was analysed by computing a percentage value of empty stomachs for each catch. Over 4 days, the percentage values presented marked fluctuations. Therefore, periodogram and form estimate analyses were performed on time series of percentages of empty stomachs, to determine the phase and periodicity of feeding rhythms. In October, animals ingested food during the daytime, presenting nocturnal peaks of empty stomachs at both depths considered in this study. In June, nocturnal peaks of empty stomachs were recorded for the slope, while a marked feeding arrhythmia was present on the shelf. A hypothesis linking feeding activity to endogenous rhythms in locomotion, cardiac and respiratory activities is presented to account for the animals diurnal ingestion, irrespective of the depth of distribution.Communicated by S.A. Poulet, Roscoff     

Ontogenetic differences in the circadian locomotor activity rhythm of the talitrid amphipod crustacean Orchestoidea tuberculata

 Adult talitrid amphipods often display a nocturnal activity pattern, which has been shown in a number of cases to be partly controlled by an endogenous rhythm of circadian period. Juvenile talitrids have been little studied in the past, but evidence from field studies indicates that they may display a different diel pattern of locomotor activity from that of adults. This study presents evidence from the field in south–central Chile that juvenile Orchestoidea tuberculata Nicolet show crepuscular activity peaks, contrasting with the nocturnal peaks shown by adult conspecifics. Under constant conditions in the laboratory, both juveniles and adults exhibit endogenous rhythms of circadian periodicity. However, the phasing of activity differs in juveniles and adults, corresponding to the differences observed in the field. It is suggested that differing phasing of peak locomotor activity may be a mechanism to reduce contact between juvenile and adult talitrids. Experimental evidence indicates a significant negative influence on survival of juveniles in the presence of adult conspecifics by intraspecific predation. Other factors influencing the timing of surface activity are also discussed. Received: 3 August 1999 / Accepted: 26 May 2000     

Looking for the clock mechanism responsible for circatidal behavior in the oyster Crassostrea gigas

Valve activity rhythm of the oyster Crassostrea gigas is mainly driven by tides in the field, but in the laboratory, only a circadian clock mechanism has been demonstrated. In an attempt to reconcile these results, the mechanisms underlying the circatidal rhythm were studied in the laboratory under different entrainment or free-running regimes and in the field at Arcachon (44°39′N/1°09′W) in February–April 2011). Results confirm the existence of a circadian clock in C. gigas. Under entrainment regimes (12-h dark/12-h light photoperiod and tidal cycles simulated by a reversing current flow), oysters exhibited both circadian and circatidal cycles. Under free-running conditions (e.g., continuous darkness), the endogenous rhythm appeared to be circadian. There was no experimental evidence for an endogenous circatidal rhythm, even in oysters just transferred from the field, where a clear tidal cycle was expressed. There are two possible mechanisms to explain tidal behavior in C. gigas: an exogenous tidal cue that drives tidal activity and masks the circadian rhythm and an endogenous circatidal clock that is sensitive to tidal zeitgebers and runs at tidal frequency.