动物内源褪黑素调控生物节律的研究进展

内源褪黑素对人类和其他哺乳动物的节律行为具有调控功能。生物节律是自然进化赋予生命的基本特征之一,生物体的生命活动受到生物节律的控制与影响。在哺乳动物中,节律调控中心是松果体,其主要功能是合成和分泌褪黑素。褪黑素广泛参与生物体节律行为的调节,本文从褪黑素的产生和作用机制,分别阐述褪黑素对昼夜节律行为和多种年节律行为的调控作用,同时明确褪黑素与生物钟及神经内分泌系统的直接作用和反馈互动的复杂集合,进一步揭示褪黑素调控生物节律的重要作用,以期为褪黑素的基础研究以及未来探究生物体的生物钟内源性发生机制提供参考。     

Endogenous rhythms of locomotion in the American horseshoe crab, Limulus polyphemus

American horseshoe crabs (Limulus polyphemus) exhibit clear circadian rhythms of visual sensitivity in the laboratory and in the field they exhibit seasonal patterns of mating behavior that are closely associated with the tides. Recent reports suggest that Limulus locomotor activity may be controlled by endogenous circadian and/or circatidal clocks and that light:dark (LD) cycles may affect the rhythmic output of both of these clocks. In this study, we examined locomotor behavior in the laboratory to determine the extent of this endogenous activity and to examine the influence of LD cycles on these rhythms. Thirty-three L. polyphemus were captured during the breeding season and their activity was monitored with activity boxes and “running wheels” in seawater kept at constant temperature and salinity. Activity patterns were analyzed using visual inspection of actograms and Chi-square and Lomb-Scargle periodograms. Overall, 36% of the animals was significantly more active during L, while only 12% was more active during D (52% showed no preference). Circatidal rhythms were observed in LD in 67% of the horseshoe crabs. Surprisingly, LD cycles appeared to synchronize these rhythms at times. In DD, the majority of animals tested (63%) exhibited circatidal rhythms that persisted for at least seven days. Overall, the results demonstrate that an endogenously controlled tidal rhythm of locomotion operates during, and significantly after, the breeding season in this species. In addition, the present results are consistent with the presence of circalunidian oscillators controlling these rhythms.     

Epigenetic Maternal Effects on Endogenous Rhythms in Precocial Birds

Development involves interactions between genetic and environmental influences. Vertebrate mothers are generally the first individuals to encounter and interact with young animals. Thus, their role is primordial during ontogeny. The present study evaluated non‐genomic effects of mothers on the development of rhythms of precocial Japanese quail (Coturnix c. japonica). First, we investigated the influence of mothering on the ontogeny of endogenous rhythms of young. We compared circadian and ultradian rhythms of feeding activity of quail reared with or without adoptive mothers. More brooded than non‐brooded quail presented a circadian and/or an ultradian rhythm. Thus, the presence of the mother during the normal brooding period favors, in the long term, expression of rhythms in the young. Second, we investigated the influence of rhythmic phenotype of the mother on the development of endogenous rhythms of young by comparing quail brooded by circadian‐rhythmic adoptive mothers (R) to quail brooded by circadian‐arrhythmic adoptive mothers (A). More R‐brooded than A‐brooded quail expressed circadian rhythmicity, and circadian rhythm clarities were greater in R‐brooded than A‐brooded quail. Ultradian rhythmicity did not differ between R‐ and A‐brooded quail, nor between R and A adoptive mothers. Thus, the rhythmic phenotypes of quail mothers influence the rhythmic phenotypes of their young. Our results demonstrate that mothers of precocial birds influence epigenetically the ontogeny of endogenous rhythms of the young they raise.     

The circadian rhythm controls telomeres and telomerase activity

Circadian clocks are fundamental machinery in organisms ranging from archaea to humans. Disruption of the circadian system is associated with premature aging in mice, but the molecular basis underlying this phenomenon is still unclear. In this study, we found that telomerase activity exhibits endogenous circadian rhythmicity in humans and mice. Human and mouse TERT mRNA expression oscillates with circadian rhythms and are under the control of CLOCK–BMAL1 heterodimers. CLOCK deficiency in mice causes loss of rhythmic telomerase activities, TERT mRNA oscillation, and shortened telomere length. Physicians with regular work schedules have circadian oscillation of telomerase activity while emergency physicians working in shifts lose the circadian rhythms of telomerase activity. These findings identify the circadian rhythm as a mechanism underlying telomere and telomerase activity control that serve as interconnections between circadian systems and aging.     

Demography of the sessile rotifers, <Emphasis Type="Italic">Limnias ceratophylli</Emphasis> and <Emphasis Type="Italic">Limnias melicerta</Emphasis> (Rotifera: Gnesiotrocha), in relation to food (<Emphasis Type="Italic">Chlorella vulgaris</Emphasis> Beijerinck, 1890) density

Circadian rhythms occur widely amongst living organisms, often in response to diel changes in environmental conditions. In aquatic animals, circadian activity is often synchronised with diel changes in the depths individuals occupy and may be related to predator–prey interactions, where the circadian rhythm is determined by ambient light levels, or have a thermoregulatory purpose, where the circadian rhythm is governed by temperature. Here, these two hypotheses are examined using animal-attached accelerometers in juvenile freshwater sawfish occupying a riverine environment displaying seasonal changes in thermal stratification. Across seasons, diel patterns of depth use (shallow at night and deep in the day) tended to occur only in the late dry seasons when the water was stratified, whereas individuals were primarily shallow in the early dry season which featured no thermal stratification. Activity was elevated during crepuscular and nocturnal periods compared to daytime, regardless of the thermal environment. Our observation of resting at cooler depths is consistent with behavioural thermoregulation to reduce energy expenditure, whereas activity appears linked to ambient light levels and predator–prey interactions. This suggests that circadian rhythms in activity and vertical migrations are decoupled in this species and respond to independent environmental drivers.     

邦亮东黑冠长臂猿日食性与活动节律的季节性变化

日食性变化和活动节律是动物行为的重要方面,通过对日食性变化与活动节律的研究,可以了解食物、温度和光照时间等环境因素的变化对动物行为的影响,以及动物在行为上的应对策略。2009 年1 -12 月,在广西靖西县邦亮东部黑冠长臂猿自治区级自然保护区内,采用瞬时扫描取样法对2 群东黑冠长臂猿觅食行为和活动节律进行了研究。结果显示,东黑冠长臂猿的活动和食性在旱季和雨季都具有明显的节律。鸣叫只发生在12:00之前,但旱季的鸣叫时间偏晚。在日节律上,东黑冠长臂猿在离开过夜树后和进入过夜树前各有一个取食高峰,07:00 喜欢取食果实和无花果,16:00 也选择更多的果实,但无花果较少,13:00 - 15:00 取食更多的叶和芽。在10:00 - 12:00,其用于休息和社会活动的时间增加。在雨季,东黑冠长臂猿以果实和无花果为主要食物,并且花大量时间觅食无脊椎动物;但在旱季,其食物以叶和芽为主。与之相适应,东黑冠长臂猿在旱季通过减少移动增加休息来节约能量开支和应对低温,同时它们增加取食的时间以获得等价的能量和营养。雨季时社会行为的比例明显高于旱季,而且其高峰出现在08:00 - 10:00,但旱季社会行为主要出现在11:00 - 14:00。在雨季东黑冠长臂猿没有出现休息高峰,相反在旱季10:00 左右出现一个明显的休息高峰。上述行为节律体现了东黑冠长臂猿对环境季节性变化的良好适应,使其能够在寒冷并且退化的喀斯特森林中生存繁衍。     

Effects of Pregnancy and Parturition on Free-Running Circadian Activity Rhythms in the Rat

Alterations in circadian rhythms have previously been associated with estrous and seasonal changes in reproductive state. In the present study we explored the effects of the reproductive events of pregnancy and parturition on free-running circadian activity rhythms in the rat. Free-running rhythms were monitored before mating, during pregnancy, and following parturition and removal of pups. Systematic and long-lasting alterations of the period of the free-running activity rhythm were seen following parturition. The effects of estrous, seasonal, and gestational reproductive states on circadian rhythms may be mediated by the endocrine events which accompany these states.     

Effects of Pregnancy and Parturition on Free-Running Circadian Activity Rhythms in the Rat

Alterations in circadian rhythms have previously been associated with estrous and seasonal changes in reproductive state. In the present study we explored the effects of the reproductive events of pregnancy and parturition on free-running circadian activity rhythms in the rat. Free-running rhythms were monitored before mating, during pregnancy, and following parturition and removal of pups. Systematic and long-lasting alterations of the period of the free-running activity rhythm were seen following parturition. The effects of estrous, seasonal, and gestational reproductive states on circadian rhythms may be mediated by the endocrine events which accompany these states.     

Activity budgets and activity rhythms in red ruffed lemurs (Varecia rubra) on the Masoala Peninsula, Madagascar: seasonality and reproductive energetics

The activity budgets and daily activity rhythms of Varecia rubra were examined over an annual cycle according to season and reproductive stage. Given the relatively high reproductive costs and patchy food resources of this species, I predicted that V. rubra would 1) travel less and feed more during seasonal resource scarcity in an attempt to maintain energy balance, and 2) show sex differences in activity budgets due to differing reproductive investment. Contrary to the first prediction, V. rubra does not increase feeding time during seasonal food scarcity; rather, females feed for a consistent amount of time in every season, whereas males feed most during the resource-rich, hot dry season. The results are consistent with other predictions: V. rubra travels less in the resource-scarce cold rainy season, and there are some pronounced sex differences, with females feeding more and resting less than males in every season and in every reproductive stage except gestation. However, there are also some provocative similarities between the sexes when activity budgets are examined by reproductive stage. During gestation, female and male activity budgets do not differ and appear geared toward energy accumulation: both sexes feed and rest extensively and travel least during this stage. During lactation, activity budgets are geared toward high energy expenditure: both sexes travel most and in equal measure, and rest least, although it remains the case that females feed more and rest less than males. These similarities between female and male activity budgets appear related to cooperative infant care. The high energetic costs of reproduction in V. rubra females may require that they allot more time to feeding year round, and that their overall activity budget be more directly responsive to seasonal climate change, seasonal food distribution, and reproductive schedules.     

Melatonin and sleep

Sleep-wake cycle is the predominant example of circadian rhythms. Melatonin is commonly used to treat insomnia and in additional neurodevelopmental disorders in which sleep disturbance is frequent. In mammals, melatonin receptors are present in the membrane and cell nucleus of many tissues and systems where it exhibits various actions, including the regulation of circadian rhythms. The rhythmic pattern of melatonin secretion is imperative since it endows with vital information to the organism concerning time, which permits for alterations of a number of physiological functions consistent with daily and seasonal variations. Melatonin as well has sleep promoting effects demonstrated in changes in brain activation patterns and tiredness generation. The SCN’s (suprachiasmatic nuclei) function and melatonin production capability turns down with age consequently depriving the brain from an important time cue and sleep regulator.     

Circadian rhythmicity and photoperiodism in the pitcher-plant mosquito: can the seasonal timer evolve independently of the circadian clock?

The two major rhythms of the biosphere are daily and seasonal; the two major adaptations to these rhythms are the circadian clock, mediating daily activities, and the photoperiodic timer, mediating seasonal activities. The mechanistic connection between the circadian clock and the photoperiodic timer remains unresolved. Herein, we show that the rhythmic developmental response to exotic light:dark cycles, usually used to infer a causal connection between the circadian clock and the photoperiodic timer, has evolved independently of the photoperiodic timer in the pitcher-plant mosquito Wyeomyia smithii across the climatic gradient of eastern North America from Florida to Canada and from the coastal plain to the mountains. We conclude that the photoperiodic timing of seasonal events can evolve independently of the daily circadian clock.     

Reproductive phenology, life-forms, and habitats of the Venezuelan Central Plain

Reproductive phenology of 171 plant species belonging to 57 families of angiosperms was studied according to life-forms in four habitat types in a savanna-forest mosaic on the Venezuelan Central Plain. Flowering, unripe fruit, and mature fruit patterns were affected significantly according to life-forms and habitats respectively. Production of flowers, unripe fruits, and mature fruits showed marked seasonality for all habitats except for the forest. Flowering peaked during the rainy season, and fruiting peaked toward the end of the rainy season. The savanna and the disturbed area had similar proportions of species that flowered over the year. The percentage of species with unripe fruits produced throughout the year was more seasonal for the disturbed area than for the other habitats. Mature fruit patterns showed an increase during the late rainy season for the ecotone and savanna. A large number of herbaceous (annual and perennial) and liana species flowered during the wet season, and a smaller fraction flowered during the dry season; and trees, shrubs, and epiphytes increased flowering activity during the dry season. Unripe fruit patterns were similar to those of flowering for all life-forms, however, tree species were less seasonal. Mature fruit production by shrubs peaked in the period of maximum rainfall, while the peak for perennial herbs was in the late rainy season and the peak for annual herbs was during the transition between the rainy season and the dry season. The largest proportion of tree and liana species with ripe fruits occurred during the dry season. Differences among phenological patterns in habitats were caused mainly by life-forms and promote a wider distribution of reproductive events in habitats and overall community in the Venezuelan Central Plain.     

韭菜迟眼蕈蚊成虫行为活动昼夜节律

【目的】本研究旨在明确韭菜迟眼蕈蚊Bradysia cellarum成虫行为活动的昼夜节律,为其室内饲养、行为学研究和研发精准、高效的监测与防控新技术提供依据。【方法】室内系统观察记录了韭菜迟眼蕈蚊成虫羽化盛期求偶、交配、产卵、移动与静息等行为的昼夜节律,分析了其特定行为的时间分配特征。【结果】韭菜迟眼蕈蚊成虫的运动行为和静息行为白天和黑夜均有发生,夜间的繁殖行为(求偶、交配和产卵)频次高于白天的,各时间点的移动、静息和繁殖行为频次占比间均具有显著性差异。韭菜迟眼蕈蚊雌雄成虫的求偶节律相同,一天中求偶有两次高峰,分别出现在3:00和7:00。韭菜迟眼蕈蚊雌雄成虫的交配节律与雌成虫的产卵节律相似,雄成虫移动行为频次占比的峰值时刻均出现在3:00时,雌成虫移动行为频次占比的峰值时刻则出现在19:00时。雌雄成虫静息行为的频次占比均无明显昼夜节律高峰。在日时间分配上,雌、雄成虫以静息行为的时间占比最多,分别为89.53%和89.90%;求偶行为的时间占比最少,分别为0.56%和0.89%。雌雄成虫各行为的日时间占比差异不显著。【结论】韭菜迟眼蕈蚊成虫的求偶、交配、产卵、移动和静息行为具有明显...     

Drosophila cryb mutation reveals two circadian clocks that drive locomotor rhythm and have different responsiveness to light

Cryptochrome (CRY) is a blue-light-absorbing protein involved in the photic entrainment of the circadian clock in Drosophila melanogaster. We have investigated the locomotor activity rhythms of flies carrying cryb mutant and revealed that they have two separate circadian oscillators with different responsiveness to light. When kept in constant light conditions, wild-type flies became arrhythmic, while cryb mutant flies exhibited free-running rhythms with two rhythmic components, one with a shorter and the other with a longer free-running period. The rhythm dissociation was dependent on the light intensities: the higher the light intensities, the greater the proportion of animals exhibiting the two oscillations. External photoreceptors including the compound eyes and the ocelli are the likely photoreceptors for the rhythm dissociation, since rhythm dissociation was prevented in so1;cryb and norpAP41;cryb double mutant flies. Immunohistochemical analysis demonstrated that the PERIOD expression rhythms in ventrally located lateral neurons (LNvs) occurred synchronously with the shorter period component, while those in the dorsally located per-expressing neurons showed PER expression most likely related to the longer period component, in addition to that synchronized to the LNvs. These results suggest that the Drosophila locomotor rhythms are driven by two separate per-dependent clocks, responding differentially to constant light.     

Circadian rhythms of adult emergence and activity but not eclosion in males of the parasitic wasp Nasonia vitripennis

An endogenous circadian system is responsible for the rhythms observed in many physiological and behavioural traits in most organisms. In insects, the circadian system controls the periodicity of eclosion, egg-laying, locomotor and mating activity. The parasitoid wasp Nasonia vitripennis has been extensively used to study the role of the circadian system in photoperiodism. In this study, behavioural activities expected to be under the control of the endogenous circadian system were characterized in Nasonia. Male emergence from the host puparium is rhythmic under light-darkness conditions while eclosion from the own pupal integument is not rhythmic but continuous. Following entrainment in light-dark conditions, males show robust free-running circadian activity rhythms with a period (τ, tau) of approximately 25.6 h in constant darkness. While the endogenous circadian system is enough to trigger male emergence in Nasonia, light seems to have a modulatory effect: when present it induces more males to emerge. Our results add to the understanding of chronobiological phenotypes in insects and provide a basis towards the molecular characterization of the endogenous circadian system in Nasonia.     

Constitutive expression of CIR1 (RVE2) affects several circadian-regulated processes and seed germination in Arabidopsis

Circadian clocks are endogenous auto-regulatory mechanisms that allow organisms, from bacteria to humans, to advantageously time a wide range of activities within 24 h environmental cycles. Here we report the identification and characterization of an MYB-related gene, designated Circadian 1 (CIR1), that is involved in circadian regulation in Arabidopsis. Expression of CIR1 is transiently induced by light and oscillates with a circadian rhythm. The rhythmic expression of CIR1 is controlled by the central oscillator. Constitutive expression of CIR1 resulted in a shorter period length for the rhythms of four central oscillator components, and much lower amplitude for the rhythms of central oscillator components CCA1 and LHY. Furthermore, CIR1 over-expression severely affected the circadian rhythms of its own RNA and those of the slave oscillator EPR1 and effector genes Lhcb and CAT3. Plants that constitutively expressed CIR1 displayed delayed flowering, longer hypocotyls and reduced seed germination in the dark. These results suggest that CIR1 is possibly part of a regulatory feedback loop that controls a subset of the circadian outputs and modulates the central oscillator.     

Age-related effects on the biological clock and its behavioral output in a primate

In humans, activity rhythms become fragmented and attenuated in the elderly. This suggests an alteration of the circadian system per se that could in turn affect the expression of biological rhythms. In primates, very few studies have analyzed the effect of aging on the circadian system. The mouse lemur provides a unique model of aging in non-human primates. To assess the effect of aging on the circadian system of this primate, we recorded the circadian and daily rhythms of locomotor activity of mouse lemurs of various ages. We also examined age-related changes in the daily rhythm of immunoreactivities for vasoactive intestinal polypeptide (VIP) and arginine-vasopressin (AVP) in suprachiasmatic nucleus neurons (SCN), two major peptides of the biological clock. Compared to adult animals, aged mouse lemurs showed a significant increase in daytime activity and an advanced activity onset. Moreover, when maintained in constant dim red light, aged animals exhibited a shortening of the free-running period compared to adult animals. In adults, AVP immunoreactivity (ir) peaked during the second part of the day, and VIP ir peaked during the night. In aged mouse lemurs, the peaks of AVP ir and VIP ir were significantly shifted with no change in amplitude. AVP ir was most intense at the beginning of the night; whereas, VIP ir peaked at the beginning of the daytime. A weakened oscillator could account for the rhythmic disorders often observed in the elderly. Changes in the daily rhythms of AVP ir and VIP ir may affect the ability of the SCN to transmit rhythmic information to other neural target sites, and thereby modify the expression of some biological rhythms.     

Annual reproductive activity of Caiman crocodilus fuscus in captivity

We describe the gonadal and abdominal fat storage cycles for a population of Caiman crocodilus fuscus kept in captivity at a breeding farm. The reproductive cycles of males and females are clearly seasonal. Vitellogenic follicles were observed in females year‐round; however, the follicular growth that produces the preovulatory follicles for the breeding season occurs in November–May (most of the dry season to the beginning of the rainy season). These months correspond with the presence of physiologically reproductive males. Oviposition takes place during the wet season (April–August) and the births occur at the end of this season (September–November). Fat storage was not related to reproductive activity, and did not vary by month or between sexes. Although environmental and hormonal conditions have been improved for these captive animals, they maintain a strongly seasonal reproductive cycle, historically fixed to oviposit in the most favorable period for nest construction and hatching during the rainy season. Zoo Biol 22:121–133, 2003. © 2003 Wiley‐Liss, Inc.     

Non-stationary time series and the robustness of circadian rhythms

Circadian rhythms are regular oscillations in the value of behavioral and physiological variables of organisms that recur on a daily basis. The purpose of this study was to evaluate the extent of non-stationarity of circadian rhythms over several days, to determine how damaging is the violation of the assumption of stationarity in the analysis of circadian rhythms, and to formalize the concept of "rhythm robustness" as an index of oscillatory ("weak") stationarity. Simulated (computer-generated) and experimental data sets (rhythms of body temperature and running-wheel activity in several rodent species) were analysed. Tests of stationarity based on the variance of the daily means and the variance of the daily variances revealed that most experimental data sets are not stationary. Analysis of linear trends indicated that significant trends are rare in experimental data sets. Although the non-stationarity of the experimental data sets reduced the spectral energy of the Enright periodogram used to assess rhythmicity, detection of circadian rhythmicity was not prevented in any of the rhythmic data sets. The results of the various analyses allow the inference that, after high-frequency noise is filtered out, the value of the periodogram's Q(P) statistic reflects the extent of stationarity of the time series. Thus, the "robustness" of a circadian rhythm (i.e. the magnitude of the empirical Q(P) value as compared to the Q(P) value associated with a perfectly rhythmic time series) can serve as an index of the stationarity of the rhythm.     

Interseasonal variation in the circadian rhythms of locomotor activity and temperature selection in sleepy lizards, <Emphasis Type="Italic">Tiliqua rugosa</Emphasis>

Few studies in non-mammalian vertebrates have examined how various effectors of the circadian system interact. To determine if the daily locomotor and behavioural thermoregulatory rhythms of Tiliqua rugosa are both controlled by the circadian system in different seasons, lizards were tested in laboratory thermal gradients in four seasons and in constant darkness. Circadian rhythmicity for both rhythms was present in each season, being most pronounced in spring and summer and least evident in autumn. Most lizards displayed a unimodal locomotor activity pattern across all seasons. However, some individuals presented a bimodal locomotor activity pattern in spring and summer. Seasonal variations in the phase relationships of both rhythms to the light:dark (LD) cycle were demonstrated. No seasonal differences in the free-running period lengths of either rhythm were detected, raising the possibility that a single circadian pacemaker drives both rhythms in this species. Our present results demonstrate that both rhythms are similarly controlled by the circadian system in each season. Although seasonal variations in the thermal preferences of reptiles both in the field and laboratory have previously been well documented, this study is the first to demonstrate circadian rhythms of temperature selection in a reptile species in each season.