Daily and seasonal rhythms of locomotor activity and adrenal function in male hedgehogs(Erinaceus europaeus L.)

Abstract

Male hedgehogs were maintained in outdoor parks, under natural climatic conditions. The general locomotor activity was recorded over one year in five hedgehogs, using an infra‐red producer‐receptor system registering entering or leaving the nest box in relation to time. Simultaneously, thecortico‐adrenal activity was studied monthly by sampling without disturbing, at 4 h intervals over a period of 24 h, in groups of 6 animals with a catheter inserted in the left carotid artery. The corticosteroids were assayed using a competitive protein binding technique.

In our study, hedgehogs living in Western Central France showed:

(1) from February‐March to August‐September a daily nocturnal locomotor activity well synchronized with sunset and sunrise,

(2) between August‐September and February‐March an alternation of resting and activity periods which constitute the hibernating period. The activity periods occur still during the night, but there are no relationships with sunset or sunrise,

(3) from February to October, a nycthemeral rhythm of plasma corticosteroids wi th an evening maximum and a morning minimum, as in laboratory animals with nocturnal locomotor activity,

(4) after October, during hibernation, significant variations of plasma corticos teroids with an irregular.pattern (shifting of the maximum or biphasic profile),

(5) a clearly annual cycle of the daily plasma corticosteroids levels, with a considerable increase in autumn (maximum in December), a significant decrease in January and a minimum from January to May.

These daily and seasonal variations of locomotor activity and cortico‐adrenal activity are related to the seasonal variations of the environment.     

The Arctic charr story: development of subarctic freshwater fish farming in Sweden

We present an overview of 27 years of experience of domestication and farming of Arctic charr (Salvelinus alpinus) in Sweden. The domestication process included an evaluation of suitable strains for farming, a breeding programme and the study of the biological and behavioural characteristics of the species. Traits of three different Arctic charr populations differing in ecology and appearance were compared during initial 2-year trials under farming conditions. The best-performing population with respect to the growth rate and the lowest frequency of early sexual maturation was a piscivore form and this became the foundation for a breeding programme intended to select for an Arctic charr strain suitable for farming. After 23 years and 7 generations, our selective breeding has resulted in a fast-growing, late-maturing strain much appreciated by farmers. The biological and behavioural characteristics studied included annual and diel locomotor activity, feeding, social and thermal behaviour. Applying our findings in these areas has greatly improved both profits and conditions for the fish. Other investigations have focused on the application and further evaluation of the results from research in practical farming trials, such as evaluation of growth at different farms with different temperature conditions, optimal time and stocking density for start-feeding and evaluation of different feeding schedules. In Sweden, Arctic charr is mainly farmed in net pens situated in nutritionally depleted and extremely unproductive water reservoirs formed by damming rivers to create electric power. Judicious farming of Arctic charr in such reservoirs can restore their nutritional and productivity state to that which existed before regulation. Site selection criteria for Arctic charr farming in such waters have been developed. The development of intensive farming of Arctic charr in Sweden is discussed together with current limitations and future possibilities.     

From autochthonous to allochthonous resources: seasonal shifts in food use by stream‐dwelling YOY Arctic charr Salvelinus alpinus through the ice‐free season

Substantial seasonal changes in resource use associated with enhanced water‐column use were revealed in stream‐living YOY Arctic charr Salvelinus alpinus during the ice‐free season. In July, YOY individuals showed a diet dominated by aquatic invertebrates (mainly Chironomidae larvae), but despite the small size of the fish, the abundance of terrestrial insects in their diet increased markedly from July to September (from 1·9 to 62·8%). Similarly, the frequency of surface drifting foragers, i.e. individuals feeding on allochthonous resources, increased from July to September (from 20·6 to 80%); allochthonous resources thus constituting an important energy subsidy for YOY S. alpinus during the late sub‐Arctic summer.     

Temporal and Spatial Habitats of Anadromous Brook Charr in the Laval River and its Estuary

Anadromy in brook charr occurs across the species’ range, but few studies have examined contiguous seasonal movements and habitat utilization of freshwater and estuarine environments. We used acoustic telemetry to track movements of anadromous brook charr for 1 year in the Laval River, Quebec. Fish entered the marine environment in May and June, inhabited shallow (<1.7 m), near-shore areas (<500 m from shore), and rarely ventured beyond the headlands of the bay. They were found where salinities were 1–34 ppt and temperatures 5–18°C. There was a strong tidal periodicity to observed movements with fish running up into bays as they became inundated at high tide and then returning to deep river channels in the bays at low tide. Between late July and early September, the charr returned to freshwater and spawned in the middle and upper sections of the river in October and November. Winter habitats were dispersed over 22 km of the river including two lakes. These detailed observations are an important contribution to our understanding of the evolutionary significance of anadromy in this species. The salinity and temperature co-tolerance provides insights into the species’ post-glacial dispersal, present-day distribution, and potential habitat maps for conservation, restoration, and enhancement programmes.     

Diet, gastric evacuation rates and food consumption in a stunted population of Arctic charr, Salvelinus alpinus L., in Takvatn, northern Norway

Diet and food consumption of a stunted population of Arctic charr in Takvatn, northern Norway, was studied throughout the ice-free season. Chironomid pupae dominated the diet in June, July and August, while zooplankton and three-spined sticklebacks dominated in September and October. Estimates of gastric evacuation rates were obtained from laboratory experiments with wild-caught, acclimated charr, and these estimates were used for in situ estimation of food consumption from weights of stomach contents. The daily food consumption was greatest in July, after which it decreased markedly towards autumn. The estimated food intake rates were low, especially in September and October. The food supply appears to be restricted, which is probably the main reason why the charr in Takvaln exhibit stunted growth.     

Behavioral rhythms of the Japanese newts, Cynops pyrrhogaster, under a semi-natural condition

 Locomotor activity rhythms of the Japanese newt, Cynops pyrrhogaster, were recorded under a semi-natural condition using phototransistor systems. The daily activity rhythm showed a seasonal change: the locomotor activity was mainly diurnal (active during the daytime) from spring to early summer; mainly nocturnal (active during the night-time) from summer to autumn; and showed either a diurnal or nocturnal pattern, depending on the ambient temperature, in winter. To analyze the daily activity in detail, we observed the behavior of a group of newts (three males, three females) throughout 24 h. Four types of behavior (respiration, feeding, mating, and resting on the land) were observed. Each behavior had daily rhythms and showed a seasonal change. The behavior on land showed mainly a nocturnal or bimodal pattern (activity rhythms with two peaks) throughout the year and was more frequently observed in summer. Mating behavior also showed a seasonal change: high activity in spring, with peaks in the early morning and evening, but no activity in summer. Except in winter, feeding and respiratory behavior showed no seasonal changes in either activity period or frequency. Coupling between behavior and the clock seems to be weak in the Japanese newt because of indistinct daily rhythms and frequent phase changes of locomotor activity in water. Physical factors such as humidity and temperature seem to affect strongly the daily activity of the newts. Received: 21 April 1997 / Accepted: 1 September 1997     

Seasonal and diel behaviour of radio-tagged Clarias gariepinus in Lake Ngezi, Zimbabwe (Pisces: Clariidae)

Seasonal and diel behaviour of six sharptooth catfish, Clarius gariepinus (Burchell), was monitored in Lake Ngezi, Zimbabwe, from May 1986 to February 1987. All specimens were equipped with surgically-implanted radiotransmi tters, 48–49 mHz, after they were anaesthetized in 6 mg l^-1 quinaldine. This represented the first known use of this methodology in subSaharan Africa.
It was determined that C. gariepinus exhibited three movement patterns on a day-to-day basis: long-distance movements ( 2200 m), moderate movements (c. 40–200 m) and local movements (<40 m). Major movements were not restricted to night-time conditions, nor were inshore/offshore movements related to time of day. Seasonal activity peaks occurred in July/August 1986 and again in January 1987, with rate of swimming exceeding 470 m/h on some occasions. Home range of some individuals encompassed the entire lake (> 5 km²).     

Seasonal changes in sea-water tolerance of Arctic charr (Salvelinus alpinus)

Summary Groups of Arctic charr,Salvelinus alpinus, which had been acclimated to water with a salinity of 7 g·l^–1 and natural temperature and photoperiod, were exposed to water with different salinities and temperatures in June, September and February. At a salinity of 15 g·l^–1, plasma osmolality, plasma Na⁺, Cl^–, Mg²⁺ concentrations and the activity of gill Na-K-ATPase were stable, irrespective of temperature and season. In June, the charr were able to regulate blood plasma ionic levels within narrow limits when exposed to a salinity of 34 g·l^–1 (sea water) and a temperature of 8°C. The hypo-osmoregulatory capacity was less, but sufficient if the temperature was only 1°C during the seawater exposure. At the start of the experiment, the gill Na-K-ATPase activity was significantly higher in June than corresponding enzyme activities in September and February. Furthermore, an increase in gill Na-K-ATPase activity during the seawater exposure (8°C) was seen in June. Irrespective of ambient temperature and salinity, no fish died during the June experiments. In September and February, exposure to sea water produced marked increases in plasma osmolality and plasma ion concentrations. There were no changes in gill Na-K-ATPase activity. Consequently, the fish became dehydrated and were moribund after a short period of seawater exposure. Highest mortality was recorded when charr were exposed to winter sea conditions (34 g·l^–1 and 1°C) in February. The results indicate that an increase in daylength induce a hypo-osmoregulatory capacity in the Arctic charr during summer. In fall and winter, however, reduced daylength are accompanied by poor hypo-osmoregulatory capacity. This leads to high mortality as a result of increased electrolyte levels and dehydration.     

Polymorphic segregation in Arctic charr Salvelinus alpinus (L.) from Vatnshlidarvatn, a shallow Icelandic lake

We studied the salmonid fish Arctic charr, Salvelinus alpinus , in a small and shallow landlocked lake in NW Iceland. The lake is productive but die only fish present is Arctic charr. Despite the apparent absence of discrete benthic and limnetic habitats for fish, two forms of Arctic charr are found in the lake. They show subde differences in morphology related to swimming performance and manoeuvrability, but differences in life history such as growth, and age and size at sexual maturation are more pronounced. Both forms have benthic feeding habits with one form consuming greater number of species than the other. We suggest that the segregation of these forms is based on the evolution of a specialist from a local generalist and that this has been made possible by the absence of a common fish competitor in similar lakes, the threespined stickleback Gasterosteous aculeatus.     

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

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

Melatonin implantation during spring and summer does not affect the seasonal rhythm of feeding in anadromous Arctic charr (Salvelinus alpinus)

Plasma melatonin levels in the high-latitude teleost Arctic charr (Salvelinus alpinus) are constantly low during summer when feeding activity is high, and high during the dark winter when they eat little and loose weight. The question arises if melatonin is involved in the phase-setting of annual rhythms of feeding and growth and if low summer melatonin production is permissive for high summer growth in this species. The present study was therefore set out to compare the seasonal appetite and growth rhythms in Arctic charr with constantly high plasma melatonin levels from February throughout the Arctic summer (melatonin implanted, average mid-day plasma melatonin levels 1,106 ± 147 pg/ml) with those of fish with natural plasma melatonin levels (vehicle implanted and untreated fish with average mid-day plasma melatonin levels of 94 ± 13 and 58 ± 6 pg/ml, respectively). Feed intake, body mass or body length, as well as the timing of the seasonal growth rhythm, were not affected by the high summer plasma melatonin level. Further, Arctic charr fasted for 3 months had a 24 h plasma profile of melatonin which was consistently higher throughout the scotophase compared to fed charr. Although the daily melatonin production seems to be affected by the energy status of the fish, melatonin does not seem to be directly involved in regulation of the seasonal feeding and growth rhythm in the high-latitude, anadromous Arctic charr.     

Individual variation in distribution, activity and growth rate of Arctic charr kept in a three-tank system

The movements and distribution of groups of Arctic charr Salvelinus alpinus were examined in a rearing system that offered a choice between two different feeding tanks separated by a larger non-feeding tank. The passages of individual fish were monitored continuously during a period of 3 weeks using the PIT (passive integrated transponder)-tag technique. The primary aim was to examine if only some charr were occupying the feeding tanks, thereby excluding other individuals, and whether differences in visit activity explained within-group variation in individual growth. On average, about 35 of the 40 charr in each group shoaled in the large non-feeding tank leaving only five individuals in the feeding tanks. Charr that spent a long total time in one of the feeding tanks made frequent excursions to the other tanks resulting in a continuous exchange of individuals. Individual growth rates were correlated positively with visit activity rather than with the total time spent in the feeding tanks. Thus, individuals with low growth rate spent as much time in feeding tanks as charr with high growth rate. However, less successful fish tended to visit the feeding tanks at night when the feeders were switched off. Based on behavioural and growth results obtained in this experiment, the use of multitank systems in the cultivation of Arctic charr is discussed.     

Adaptations to Stochastic Environmental Variations: The Effects of Seasonal Temperatures on the Migratory Window of Svalbard Arctic Charr

The circumpolar Arctic charr, Salvelinus alpinus, is ideal for studying how environmental factors affect life history in fishes. Charr populations demonstrate a tremendous ecological plasticity and adaptations to harsh environments. Arctic charr is the only freshwater fish on Svalbard, including anadromous, resident, and landlocked stocks. Freshwater lake systems on Svalbard are characterized by very low water temperatures, long-term or even permanent ice cover, and low levels of nutrients. Food is thus limited and may lead to growth stagnation and early maturity in Arctic charr. The individual growth pattern may alternatively follow a sigmoid-shaped curve, caused by a shift to either cannibalistic or anadromous (migration to sea) behaviour. In lake systems that include migratory charr, the population may consist of a mixture of parr, postsmolt, and adult migratory individuals, as well as small-sized resident, large-sized resident (cannibals), and large formerly resident individuals transformed to anadromy. Our study in the Lake Dieset watercourse (79°N), Svalbard, demonstrates that the annual water flow in the outlet river is strongly correlated to air temperatures and provides a passage to the sea, allowing the charr access to the nutrient-rich seawater environment, during at most two months each year. During one of the years studied, the youngest and small-sized part of the sea-going stock was prevented from ascending the river and probably suffered mortality during winter. The migratory window of the Arctic charr in Lake Dieset is therefore highly variable among years and thus unpredictable. We hypothesize that in worst case scenarios (cold years, low water discharge), climatic variations may occasionally prevent charr from migrating upstream in Svalbard lake systems in late autumn, resulting in high mortality in the population.     

Seasonal changes in hypoosmoregulatory ability in landlocked and anadromous populations of Arctic charr,Salvelinus alpinus,and Atlantic salmon,Salmo salar

Synopsis Seasonal changes in hypoosmoregulatory ability were compared in landlocked and anadromous strains of Arctic charr and Atlantic salmon. Seawater adaptability was assessed using periodic 48 h seawater challenge tests with 25. seawater. The landlocked strains of Arctic charr, two from northern Sweden and one from Southern Norway, displayed similar seasonal changes in seawater adaptability as the anadromous strain. Seawater tolerance increased during spring and remained high until the end of July — early August after which it declined. The two strains of Atlantic salmon displayed different seasonal patterns in hypoosmoregulatory ability. The anadromous strain showed a pronounced seasonal pattern with maximal seawater adaptability in early June. In contrast, seawater tolerance in the landlocked strain improved steadily during spring and remained high until late autumn. During the period of enhanced seawater tolerance, hypoosmoregulatory ability increased significantly with body size in both Arctic charr and anadromous Atlantic salmon. The minimum size at which fish were able to regulate plasma sodium following seawater transfer at a level comparable to freshwater levels (<170 mmol I^–1) differed significantly between anadromous Atlantic salmon (ca. 14 cm) and Arctic charr (ca. 22 cm). The results show that seasonal changes in hypoosmoregulatory ability are present in both Atlantic salmon and Arctic charr, and that these physiological traits are retained in the corresponding landlocked strains. However, the seasonal pattern of seawater adaptability as well as the minimum size at which seawater tolerance occurs differs between the two species.     

Sex steroids, growth and condition of Arctic charr broodstock during an annual cycle

Changes in plasma concentrations of sex steroids, growth rate and condition of repeat spawning (3+) male and female Arctic charr were studied throughout an annual reproductive cycle. Individually marked fish (mean weight approx. 500 g) were held under conditions of liberal food supply, constant temperature (4° C) and simulated natural photoperiod (Tromsø, 70° N). Once each month fish were weighed, measured and blood samples taken for steroid analysis. Plasma concentrations of testosterone (T), 11-ketotestosterone (11-KT) and oestradiol-17β (E₂) were determined using radioimmunoassay (RIA). Both male and female fish displayed distinct seasonal changes in plasma concentrations of sex steroids, growth rate and condition. From February (minimal concentrations) to March all sex steroids increased slightly and these elevated concentrations were maintained until May. Thereafter, there was a second, and far more pronounced, increase in plasma steroid concentrations which culminated in peak steroid concentrations in September–October. There was then a rapid decline during the spawning period. In winter, growth rate and condition were generally low, then increased during the spring, reached a peak during the summer, and then declined with the onset of autumn. During spring (March–May), the frequency distributions of plasma testosterone concentrations in both male and female fish were bimodal. The fish of the upper modal group of the distribution had significantly higher growth rates and condition than those in the lower modal group. In summer and early autumn (June–September) the association between T and growth rate changed. Significant negative correlations between T and growth rates were observed in females. There was an increase in endocrine activity, indicated by elevated plasma sex steroid concentrations in March, 7–8 months prior to maturation. It is suggested that this may be one factor influencing the onset of spring growth and energy deposition among maturing charr.     

Migratory patterns of anadromous white-spotted charr Salvelinus leucomaenis in Eastern Hokkaido, Japan: The solution to a mystery?

We evaluated the migratory patterns of white-spotted charr in eastern Hokkaido, Japan using: (1) field observation of fish, (2) salinity-recording archival tags, and (3) analysis of otolith Sr and Ca concentrations. Field observations suggest that the majority of anadromous white-spotted charr outmigrate to the sea in spring (April to June) and ascend the river between late summer and autumn (August to November). Salinity records from archival tags revealed a fluctuating pattern consisting of both low and high values within a month, indicating frequent short-term movements between freshwater (including non-natal rivers) and brackish/salt water habitats. Otolith Sr:Ca ratios exhibited a sharp rise at about 40–50% of the otolith radius, which was followed by several spikes consisting of both low and high values, indicating that fish migrated between freshwater and marine environments multiple times during their life history. However, the peaks in Sr:Ca ratios were generally associated with annulus zones on the otolith, indicating that the anadromous migration was generally an annual event.     

Life History of Ilybius fenestratus (Fabricius) (Coleoptera,Dytiscidae) in a Central Norwegian Lake

The life history of Ilybius fenestratus was studied in Målsj?en, a lake in S?r-Tr?ndelag, Central Norway (63°14'N, 10°26'E), during 1971–1972. Adults and larvae were sampled in activity traps every week during March–October and every 2–3 weeks during November–February. Newly-emerged adults were generally found from mid-July to September, and after overwintering in torpidity, probably in the water, they again appeared in the second half of June to August/September. Laying of eggs took place from early/mid July to early August. Larvae occurred in the traps from late July to early May. I. fenestratus was thus found to be a semivoltine summer breeder, with overwintering larvae the first winter and overwintering adults the next. Both adults and larvae lived in large areas of the lake's littoral zone.     

Empirical analysis of the influence of swimming pattern on the net energetic cost of swimming in fishes

We tested the hypothesis that the energetics of swimming in a flume accurately represent the costs of various spontaneous movements using empirical relationships between fish swimming costs, weight, and speed for three swimming patterns: (1) 'forced swimming' corresponded to movements adopted by fish forced to swim against a unidirectional current of constant velocity; (2) 'directed swimming' was defined as quasi-rectilinear movements executed at relatively constant speeds in a stationary body of water and (3) 'routine swimming' was characterized by marked changes in swimming direction and speed. Weight and speed explained between 76% (routine swimming) and 80% (forced swimming) of net swimming cost variability. Net costs associated with different swimming patterns were compared using ratios of model predictions (swimming cost ratio; SCR) for various weight and speed combinations. Routine swimming was the most expensive swimming pattern (SCR for routine and forced swimming =6.4 to 14.0) followed by directed (SCR for directed and forced swimming =0.9 to 2.8), and forced swimming. The magnitude of the difference between the net costs of forced and spontaneous swimming increases with movement complexity and decreases as fish weight increases.     

哀牢山中山湿性常绿阔叶林土壤呼吸季节和昼夜变化特征及影响因子比较

 由于受到多种生物和非生物因素的影响,土壤呼吸在不同时间尺度上的动态变化可能不一致。对不同时间尺度的土壤呼吸动态变化的研究有助 于深入了解土壤呼吸变化的机理,也有利于精确推算土壤碳的排放。采用红外CO₂分析法测定哀牢山中山湿性常绿阔叶林季节间(2004年4月～ 2005年3月)和昼夜间 (2004年7、9和11月及2005年1、3和5月共6次)的土壤呼吸。哀牢山中山湿性常绿阔叶林中土壤呼吸的季节变化显著,其中 湿季(5～10月)的土壤呼吸高于干季(11月～翌年4月),全年土壤呼吸的平均值为0.442 g CO₂&;#8226;m^－2&;#8226;h^－1。6 次测定的土壤呼吸日变化模式并不 相同,7和9月、翌年1和3月夜间土壤呼吸大于昼间土壤呼吸,11月和翌年5月则相反;5、7和9月昼夜间的土壤呼吸最大值与最小值的差异比11 月、翌年1和3月的测定结果大。季节间土壤呼吸与土壤温度(p=0.000)和土壤含水量(p=0. 007) 均有显著的指数相关,土壤温度可以解释土壤 呼吸变化的56.1％,土壤含水量可以解释土壤呼吸变化的11.1％。不同季节测定的土壤呼吸日变化与土壤温度、气温和土壤含水量则没有显著 的指数相关。由土壤呼吸与土壤温度拟合的指数方程计算Q₁₀值,在温度为 5.9～16.6 ℃内,全年土壤呼吸的Q₁₀值为4.53,在温度为5.9～ 11.0 ℃内,干季土壤呼吸的Q₁₀值为7.17,在温度为10.3～16.6 ℃内,湿季土壤呼吸的Q₁₀值为2.34。在不同时间尺度上,生物和非生物因素 对哀牢山中山湿性常绿阔叶林的土壤呼吸表现出不同的影响。土壤呼吸的季节变化主要受非生物因子温度和水分变化的调控,而土壤呼吸的昼 夜变化则可能主要受植物的生理活动周期性等生物因素的影响。通过温度的指数函数关系,用土壤呼吸的瞬时值来推算土壤呼吸的日通量和年 通量时,需要考虑温度和水分外的其它生物因子的影响。