Integrative Action and Temporal Order - A Review of Complex Annual Rhythms

Complex rhythms during the interval of a year (those rhythms with more than one peak and one trough) are reviewed for two species, cats and human beings. The quantitative variation during the year of grooming reflexes in the cat represents a systematic change in the expression of integrative action, a concept first formulated by Sherrington. The evaluation of these changes as systematic rather than random is addressed by a simple simulation and by the application of a periodic regression analysis. The synchrony among individuals within a group and the synchrony among groups studied during different years establish that environmental factors control and regulate the complex rhythms. Two hypotheses are presented: 1. The complex rhythms are driven by an environmental variable with the same complex pattern; and 2. the complex rhythms are generated by a complex photoperiodic response curve. These hypotheses are described and examples of the evidence supporting them are presented. The review of the literature establishes that complex multi-modal rhythms exist during the interval of a year; that these may be considered adaptive in providing more than one “window” for producing offspring; that a basic multi-modal variation in the physiological substrate is a common and ubiquitous aspect of temporal order, and may be the rhythmic source of periodic diseases.     

Circannual clocks in avian reproduction and migration

Many behavioural and physiological functions of organisms are adjusted to the periodic changes in their environment, particularly to those related to the natural day and year. This adjustment is often achieved through the action of endogenous daily (circadian) and annual (circannual) clocks. Studies of the control of avian moult, migration and reproduction have played a major role in understanding how biological clocks function and interact with rhythms in the environment. Investigations on tropical birds such as the East African subspecies of the Stonechat ( Saxicola torquata axillaris ). and long-distance migrants like the Garden Warbler ( Sylvia borin ). have provided the longest records of circannual rhythms, some of them running for more than 12 years, with periods ranging from about 9 to 13 months. Avian circannual rhythms are organized in a characteristic way for a particular species or population, and cross-breeding experiments have shown that some of the differences found among them are genetically determined. In African Stonechats circannual rhythms guarantee that seasonal events occur at the appropriate times of the year and in the characteristic sequence. They also control a "reproductive window" that provides the temporal framework for breeding. The width of this window is rather inflexible but the performance of a bird within this framework (e.g. whether it breeds once or twice per season) is subject to modification by environmental conditions. In migratory birds circannual programs are involved in determining the time course, distance and direction of migration. Circannual rhythms are synchronized with and modified by environmental factors in a complex way, but the endogenous mechanisms usually respond to environmental cues such that an optimal adjustment to season and latitude is guaranteed.     

Multiple environmental determinants of regional species richness and effects of geographic range size

Understanding patterns of species richness at broad geographic extents remains one of the most challenging yet necessary scientific goals of our time. Many hypotheses have been proposed to account for spatial variation in species richness; among them, environmental determinants have played a central role. In this study, we use data on regional bat species richness in the New World to study redundancy and complementarity of three environmental hypotheses: energy, heterogeneity and seasonality. We accomplish this by partitioning variation in species richness among components associated with unique and combined effects of variables from each hypotheses, and by partitioning the overall richness gradient into gradients of species with varying breadths of geographic distribution. These three environmental hypotheses explain most variation in the species richness gradient of all bats, but do not account for all positive spatial autocorrelation at short distances. Although environmental predictors are highly redundant, energy and seasonality explain different and complementary fractions of variation in species richness of all bats. On the other hand, heterogeneity variables contribute little to explain this gradient. However, results change dramatically when richness is estimated for groups of species with different sizes of geographic distribution. First, the amount of variation explained by environment decreases with a decrease in range size; this suggests that richness gradients of small‐ranged species can not be explained as easily as those of broadly distributed species, as has been implied by analyses that do not consider differences in range size among species. Second, the relative contribution of environmental predictors to explained variation also changes with change in range size. Seasonality and energy are good predictors of species with broad distributions, but they loose almost all explanatory power for richness of species with small ranges. In contrast, heterogeneity, which is a relatively poor predictor of richness of species with large ranges, becomes the main predictor of richness gradients of species with restricted distributions. This suggests that range size is a different dimension on which heterogeneity and other environmental characteristics are complementary to each other. Our results suggest that determinants of species richness gradients might be complex, or at least more complex than many studies have previously suggested.     

Circannual clocks in avian reproduction and migration

Many behavioural and physiological functions of organisms are adjusted to the periodic changes in their environment, particularly to those related to the natural day and year. This adjustment is often achieved through the action of endogenous daily (circadian) and annual (circannual) clocks. Studies of the control of avian moult, migration and reproduction have played a major role in understanding how biological clocks function and interact with rhythms in the environment. Investigations on tropical birds such as the East African subspecies of the Stonechat ( Saxicola torquata axillaris ) and long-distance migrants like the Garden Warbler ( Sylvia boriri ) have provided the longest records of circannual rhythms, some of them running for more than 12 years, with periods ranging from about 9 to 13 months. Avian circannual rhythms are organized in a characteristic way for a particular species or population, and cross-breeding experiments have shown that some of the differences found among them are genetically determined. In African Stonechats circannual rhythms guarantee that seasonal events occur at the appropriate times of the year and in the characteristic sequence. They also control a "reproductive window" that provides the temporal framework for breeding. The width of this window is rather inflexible but the performance of a bird within this framework (e.g. whether it breeds once or twice per season) is subject to modification by environmental conditions. In migratory birds circannual programs are involved in determining the time course, distance and direction of migration. Circannual rhythms are synchronized with and modified by environmental factors in a complex way, but the endogenous mechanisms usually respond to environmental cues such that an optimal adjustment to season and latitude is guaranteed.     

Regression Models for the Estimation of Orcadian Rhythms in the Presence of Effects Due to Masking

The estimation of human circadian rhythms from experimental data is complicated by the presence of “masking” effects associated with the sleep-wake cycle. The observed rhythm may include a component due to masking, as well as the endogenous component linked to a circadian pacemaker. In situations where the relationship between the sleep-wake cycle and the circadian rhythm is not constant, it may be possible to obtain individual estimates of these two components, but methods commonly used for the estimation of circadian rhythms, such as the cosinor analysis, spectral analysis, average waveforms and complex demodulation, have not generally been adapted to identify the modulations that arise from masking. The estimates relate to the observed rhythms, and the amplitudes and acrophases do not necessarily refer to the endogenous rhythm.

In this paper methods are discussed for the separation of circadian and masking effects using regression models that incorporate a sinusoidal circadian variation together with functions of time since sleep and time during sleep. The basic model can be extended to include a time-varying circadian rhythm and estimates are available for the amplitude and phase at a given time, together with their joint confidence intervals and tests for changes in amplitude and acrophase between any two selected times. Modifications of these procedures are discussed to allow for non-sinusoidal circadian rhythms, non-additivity of the circadian and time-since-sleep effects and the breakdown of the usual assumptions concerning the residual errors.

This approach enables systematic masking effects associated with the sleep-wake cycle to be separated from the circadian rhythm, and it has applications to the analysis of data from experiments where the sleep-wake cycle is not synchronized with the circadian rhythm, for example after time-zone transitions or during irregular schedules of work and rest.     

Ultradian and circadian rhythms of sleep-wake and food-intake behavior during early infancy

The early development of sleep-wake and food-intake rhythms in human infants is reviewed. The development of a 24h day-night rhythm contains two observable developmental processes: the alterations in the periodic structure of behavior (decreased ultradian, increased circadian components) and the process of synchronization to external time (entrainment). The authors present the results of their studies involving 26 German children and compare them with previous investigations. In their research, it became evident that, during the first weeks of life, the time pattern of sleep-wake and food-intake behavior is characterized by different ultradian periodicities, ranging from 2h to 8h. In the course of further ontogenesis, the share of ultradian rhythms in the sleep-wake behavior decreases, while it remains dominant for food-intake behavior. The circadian component is established as early as the first weeks of life and increases in the months that follow. Besides, the authors' study supports the notion of broad interindividual variation in ultradian rhythms and in the development of a day-night rhythm. Examples of free-running rhythms of sleep-wake and food-intake behavior by various authors are strong indicators of the endogenous nature of the circadian rhythms in infants and show that the internal clock is already functioning at birth. It is still uncertain when the process of synchronization to external and social time cues begins and how differences in the maturation of perceptive organs affect the importance of time cues for the entrainment. Prepartally, the physiological maternal entrainment factors and mother-fetus interactions may be most important; during the first weeks of life, the social time cues gain importance, while light acts as a dominant “zeitgeber” at a later time only.     

Causes and consequences of variation in competitive ability in plant communities

Abstract. Several factors may define the cause and pattern of variation in competitive ability among individuals within a plant community. Variation may be a consequence of genetic or environmental variability. These two sources of variation may vary in their relative magnitudes. The relevant scale of genetic variation may occur at the individual genotype level or at the species level. The relevant scale of environmental variation may occur at the individual plant level or at the neighbourhood (or community) level. Relative competitive abilities may be effected by genotype-environment interaction or by genotype-genotype (or species-species) interaction. The complex relationship among these factors reveals the mechanistic basis for establishing a clear distinction among five specific hypotheses for species coexistence and diversity that are all variations of the general hypothesis that competitive abilities do not differ sufficiently among coexisting species to cause any competitive exclusion at the community level. These hypotheses are compared in terms of the degree to which they are restricted by assumptions and supported by existing data, and in the extent to which they involve evolutionary consequences of competition.     

Seasonal variation of activity patterns in roe deer in a temperate forested area

We investigated the activity patterns of a European roe deer (Capreolus capreolus) population living in a forested Apennine area in central Italy, in order to shed light on the environmental and biological factors that were expected to account for the observed activity patterns on daily and yearly bases. Daily and seasonal activity patterns of 31 radio-collared roe deer were assessed through sessions of radio tracking for a total period of 18 consecutive months. Roe deer showed bimodal activity patterns throughout the year, with the two highest peaks of activity recorded at dawn and dusk. Activity patterns of males and females differed during the territorial period (from early spring to late summer), whereas they did not during the nonterritorial period. Most likely, behavioral thermoregulation can be held responsible for variation of daily activity patterns in different seasons. In winter, for instance, activity during the dawn period was significantly higher than in other seasons and daylight activity was significantly higher than at night. Nocturnal activity was highest in summer and lowest in winter. During the hunting season, moreover, roe deer showed lower activity levels than during the rest of the year. The prediction that roe deer would show lower activity levels during full moon nights, when the predation risk was assumed to be higher, was not confirmed by our data. Activity rhythms in roe deer were thus subjected to both endogenous and environmental factors, the latter working as exogenous synchronization cues. Accordingly, in changing environmental and ecological conditions, a circadian cycle of activity could be seen as the result of complex interactions among daily behavioral rhythm, digestive physiology, and external modifying factors.     

Geographic variation of wing morphology of great fruit‐eating bats (Artibeus lituratus): environmental,genetic and spatial correlates of phenotypic differences

Responses of species to environmental gradients are important and frequent determinants of geographic phenotypic variation that can drive adaptive processes. Nonetheless, random genetic processes such as drift can also result in geographic variation in phenotypes, and should be evaluated before implicating selection as the process driving phenotypic change. We examined geographic variation in wing morphology of Artibeus lituratus among 18 different sites distributed across interior Atlantic Forest of Paraguay and Argentina. Moreover, we contrasted geographic variation with environmental, spatial, and genetic variation to test hypotheses related to selection and drift and their impacts on wing morphology. For A. lituratus distributed across interior Atlantic Forest, significant differences among sites characterized variation in wing morphology. Geographic variation was significantly related to climatic variables but not spatial or genetic distances. Such a pattern suggests that phenotypic variation is related to selection for particular environmental regimes, and not genetic drift. Four significant dimensions of phenotypic variation were determined. Three dimensions were related to variation among individuals in terms of wing tips, whereas one was related to overall body size. Wing tips are important for manoeuverability during flight and differences among sites likely reflect differences in forest and vegetation structure that must be managed during foraging. Although climate provides good surrogates for environmental variation, it is probably only an indirect cue of selection regimes that determine variation in wing morphology. Future studies should evaluate more direct environmental measures such as vegetation structure when attempting to interpret geographical variation in wing morphology.     

Individual Variation in Sleep-Wake Rhythms in Free-Living Birds

Ultradian rhythms, such as sleep-wake periodicities, during the night might represent basic rest-activity cycles of organisms that are fundamental to the temporal organization and synchronization of behavior throughout the day. However, in contrast to circadian rhythms, little is known about the underlying oscillators and molecular mechanisms of higher-frequency rhythms. A fundamental step for the understanding of the mechanisms of these latter periodicities is the analysis of variation in sleep-wake cycles in free-living animals, which can help in estimating the relative importance of genetic and environmental influence on the rhythmicity. We analyzed variation in the level of rhythmicity and period length (τ) of behaviorally defined sleep-wake cycles in a natural population of blue tits Cyanistes caeruleus. Our results indicate that the expression of periodicity in sleep-wake patterns, but not τ, has a strong individual-specific basis. The within-individual repeatability estimate of the expression of periodicity was .45 (95% confidence interval: .35–.55) when data from males and females were combined. In addition, periodicity was influenced by specific environmental factors, such as night temperature, seasonal date, and age of the individual. Most strikingly, low nighttime temperature negatively affected periodicity of sleep-wake patterns, potentially via a hypothermic response of the birds. Our results further suggest that τ is influenced by photoperiod. Blue tits showed longer sleep-wake rhythms when the nights were longer. These observations suggest a genetic basis for the incidence of rhythmic sleep-wake behavior in addition to environmental modifications of their specific expression. (Author correspondence: mueller@orn.mpg.de)     

Extrinsic and intrinsic factors influencing the emergence and return of the Asian particolored bat Vespertilio sinensis to the summer roost

Circadian rhythms play a crucial role in the health and survival of organisms. However, little is known concerning how intrinsic and extrinsic factors affect animal daily rhythms in the field, especially in nocturnal animals. Here, we investigated the first emergence, mid‐emergence, and return times of Vespertilio sinensis, and also integrated environmental conditions (temperature, humidity, and light intensity) and biotic factors (reproductive status and predation risk) to determine causes of variation in the activity rhythms of the bats. We found that variation in the first emergence time, the mid‐emergence time, and the final return time were distinct. The results demonstrated that the emergence and return times of bats were affected by light intensity, reproductive status, and predation risk in a relatively complex pattern. Light intensity had the greatest contribution to activity rhythms. Moreover, we first investigated the effects of actual predators on the activity rhythms of bats; the results showed that the mid‐emergence time of bats was earlier as predators were hunting, but the final return time was later when predators were present. Finally, our results also highlighted the importance of higher energy demands during the lactation in bats to variation in activity rhythms. These results improve our understanding of the patterns and causes of variation in activity rhythms in bats and other nocturnal animals.     

Environmental determinants of geographic butterfly richness pattern in eastern China

A long-standing task for ecologists and biogeographers is to reveal the underlying mechanisms accounting for the geographic pattern of species diversity. The number of hypotheses to explain geographic variation in species diversity has increased dramatically during the past half century. The oldest and the most popular one is environmental determination. However, seasonality, the intra-annual variability in climate variables has been rarely related to species richness. In this study, we assessed the relative importance of three environmental hypotheses: energy, seasonality and heterogeneity in explaining species richness pattern of butterflies in Eastern China. In addition, we also examined how environmental variables affect the relationship between species richness of butterflies and seed plants at geographic scale. All the environmental factors significantly affected butterfly richness, except sampling area and coefficient of variation of mean monthly precipitation. Energy and seasonality hypotheses explained comparable variation in butterfly richness (42.3 vs. 39.3 %), higher than that of heterogeneity hypothesis (25.9 %). Variation partitioning indicated that the independent effect of seasonality was much lower (0.0 %) than that of energy (5.5 %) and heterogeneity (6.3 %). However, seasonality performed better in explaining butterfly richness in topographically complex areas, reducing spatial autocorrelation in butterfly richness, and more strongly affect the association between butterflies and seed plants. The positive relationship between seed plant richness and butterfly richness was most likely the result of environmental variables (especially seasonality) influencing them in parallel. Insufficient sampling may partly explain the low explanatory power of environmental model (52.1 %) for geographic butterfly richness pattern. Our results have important implications for predicting the response of butterfly diversity to climate change.     

Circadian Pattern of Cercarial Emergence in Schistosoma Mansoni(Platyhelminthes: Digenea) from Isolated Biomphalaria Glabrata

The present work aimed to compare the acrophases (peak hours) of emergence of Schistosoma mansoni cercariae among isolated individuals of the snail Biomphalaria glabrata. Laboratory stocks of melanic B. glabrata from the same biotope as the S. mansoni strain (Belo Horizonte, Minas Ger-ais) were used. Twenty-two snails individually exposed to five miracidia were tested. Chronobiological trials were performed outdoors after an acclimation period of at least a week. Three groups of snails were tested between November 1989 and April 1991. Cercarial emergence from individual isolated snails was quantified every 3 h for 3 consecutive days. In all trials, most cercariae were found to emerge during daytime (94.9%). Time series and chronograms showed recurrent peaks during the daytime. The periodogram suggested that 24 h was the period that best fitted cercarial emergence data in 90.9% of the snails. The single cosinor analysis confirmed 24-h rhythms in 95.5% of the snails. Acrophases of cercarial emergence among individual snails occurred between 14:15 and 17:02. They did not differ significantly. The population cosinor analysis indicated greater homogeneity in the 24-h rhythms of cercarial emergence than in the snail groups of each chronobiological trial. Acrophases of cercarial emergence occurred between 14:53 and 15:27 and did not differ significantly among all trials. Data from the three trials were pooled and analyzed using the population cosinor. This statistical method indicated a homogeneity in the 24-h rhythms of cercarial emergence from all snails, with acrophase occurring around 15:00. Results showed that the acrophases of cercarial emergence of S. mansoni are similar among isolated B. glubrura specimens. Data support the hypothesis of a “gate” rhythm in the dynamics of cercarial production and emergence. It is suggested that the adaptive importance of the “gate” mechanism is associated with the concentration of cercariae in the water at times when the vertebrate is present, optimizing the contact between the parasite and the host. The emergence of some cercariae at night (5.1% of the total number of emerged cercariae) suggests a flexible “gate” that could be associated with a residual light effect or with experimental procedures in the laboratory.     

Environmental change drives long‐term recruitment and growth variation in an estuarine fish

How individuals respond to environmental change determines the strength and direction of biological processes like recruitment and growth that underpin population productivity. Ascertaining the relative importance of environmental factors can, however, be difficult given the numerous mechanisms through which they affect individuals. This is especially true in dynamic and complex estuarine environments. Here, we develop long‐term otolith‐based indices of recruitment and growth for estuary perch Percalates colonorum (Bemm River, Australia), to explore the importance of intrinsic (individual, demographic) and extrinsic (hydrologic, climatic, density‐dependent) factors in driving estuarine fish productivity. Analyses involved a novel zero‐inflated specification of catch curve regression and mixed effects modelling. The 39 years of recruitment and 46 years of growth data, spanning a period of environmental change including severe drought, displayed considerable inter‐annual variation. Recruitment success was strongly related to high freshwater inflows during the spawning season, suggesting that these conditions act as spawning cues for adults and potentially provide favourable conditions for larvae. Individuals displayed age‐dependent growth, with highest rates observed at younger ages in years characterized by warm temperatures, and to a lesser degree, greater magnitude base inflow conditions. We detected systematic among‐year‐class growth differences, but these were not attributable to year class strength, suggesting that environmental conditions experienced by individuals as juveniles can have long‐lasting effects of greater importance to population productivity than density‐dependent growth responses. The primacy of temperature in driving growth variation highlights that under‐appreciated climatic variation can affect estuarine fish productivity through direct physiological and indirect food web mechanisms. We predict that climatic warming will promote individual growth in southerly populations of P. colonorum but concurrently limit recruitment due to forecast reductions in spawning season river discharge. Disparate trait responses are likely in other fishes as they respond to multiple and changing environmental drivers, making predictions of future population productivity challenging.     

Effects of Depressed Affect on Diurnal and Ultradian Variations in Mood in a Healthy Sample

The study investigated diurnal and ultradian variations in cheerful and depressed mood in a healthy sample, and examined whether these variations were related to severity of depressed affect. Thirty employed volunteers used a pocket computer to complete self-rating scales of depression and cheerfulness every 2 h for 14 days except during sleep periods. They also completed the Beck Depression Inventory (BDI) and the General Health Questionnaire (GHQ). Analysis of variance of depressed mood showed a significant interaction between time of day and BDI/GHQ, in which only the mood of the high BDI/GHQ groups got worse during the day. The low BDI group showed a diurnal variation in cheerful mood in which mood was at its best in the early evening. Cosinor and “binfit” analysis of raw and residual time-series showed that more than half of the participants had significant fitting ultradian rhythms in mood in the range 3-9 h. However, only the number of significant fits for residual cheerful mood was substantially above the level of chance. The amplitudes of the ultradian rhythms in raw depressed mood were greater in participants scoring higher on the BDI/GHQ scales, but this seemed to reflect an increase in serial dependency between depressed mood states. The results are discussed in relation to circadian rhythms in mood and depression.     

Ultradian and asymmetric rhythms of hemispheric processing speed

Daily changes in cognitive performance have been documented, both in time of day/effect paradigm studies and in time-isolation studies. However, in both types of studies, phenomena such as the “post-lunch dip” have been found that were difficult to explain in terms of theoretical backgrounds. These phenomena may suggest ultradian rhythms in cognitive performance. A number of studies have also shown ultradian and asymmetric rhythms in activity indices of the brain hemispheres. The aim of this study was to test three hypotheses: the first two assumed that there is a significant ultradian frequency in a component of the endogenous rhythm of processing speed, and the third assumed that the ultradian endogenous rhythms of the processing speed (encoding and recognition) of stimuli addressed to the left brain hemisphere differ in period length from those addressed to right hemisphere. During a 24 h constant-routine experiment, the memory performance of 30 participants was measured eight times (every 2.5-3 h), starting at 06:30 h. Parallel sets of words and pictures were shown to subjects in a random order in either the left or the right visual field on a computer screen. The participants pressed one of two buttons in response to the picture or word, or when answering a question concerning the meaning of a presented stimulus. Cosinor analysis was applied to individual time series data. Two significant ultradian components were found in a majority of the time series. Dominant periods were analyzed using three factor ANOVA. The results showed an asymmetry between both hemispheres in the frequency of ultradian rhythms in encoding speed.     

Ultradian, circahoral and circadian structures in endothermic vertebrates and humans

1. For more than 30 years many studies have been carried out concerning rhythms with periods approaching 24 hr (circadian rhythms). 2. The latter have been demonstrated as resulting from environmental 24 hr synchronizers (zeitgebers), but they usually persist in the absence of a 24 hr synchronization, which proves their endogenous nature. 3. Biological rhythms with periods less than 20 hr (ultradian rhythms) and particularly those approaching 1 hr (circahoral rhythms) have been determined: for motility, rest-activity, sleep phases, endocrine secretions and other physiological functions. 4. These ultradian and circahoral rhythms have been found in rodents, birds, monkeys and humans. 5. Existing at all stages of ontogeny, they have been proved to be endogenous and species and strain specific. 6. As these ultradian rhythms can be influenced by environmental factors and sometimes by circadian rhythms they are not truly periodic, so therefore cannot be computed by the usual processes of mathematical time analysis.     

Daily temporal organization of laying in Japanese quail: variability and heritability

In birds, many behavioral and physiological processes that occur during reproduction show daily rhythms in response to environmental temporal constraints. In this study, the individual daily organization of laying and its genetic determinant in Japanese quail (Coturnix coturnix japonica) were analyzed. For this purpose, the oviposition time of 102 randomly chosen females, maintained in long-day photoperiodic conditions (LD 14h:10h) for 1 mon was observed and extreme phenotypes selected. Laying is characterized by two parameters: oviposition interval and laying hour.

The birds showed a specific time of laying during 24 h. All eggs were laid in the afternoon between 6.5 and 14 h after lights on (HALO). Two laying profiles were determined: 20% of females with an oviposition interval greater than 24 h (24.7±0.2 h) (the “delayed” profile) laid progressively later each day until a pause day. The remaining 80% of the females laid at the same time each day, with few pause days and an oviposition interval close to 24 h (24.0±0.2 h) (the “stable” profile). Among the females, showing this last profile, an intra-individual stability and an inter-individual variability of laying hour was established. Two extreme laying phenotypes were then determined: the “early” phenotype (E) for females laying on average between 7.5 and 9.5 HALO and the “late” phenotype (L) for females laying between 12.5 and 14 HALO.

In order to study the genetic basis of the laying hour, three females of each extreme phenotype were selected and crossed with two different males. The E and L females produced 57 F1E and 42 F1L daughters, respectively. F1 females displayed both laying profiles. However, the proportion of females displaying a “delayed” profile was higher in the L line (50%) than in the E line (29.8%). For the “stable” daughters, artificial selection induced an advance in laying hour of 4.7% for the E line and a delay of 4.7% for the L line. Realized heritability was estimated at 0.5. Moreover, the laying hour of the daughters was correlated positively to that of the mothers (N=61;r=0.45). These results support the notion of heritability of oviposition time in Japanese quail.     

Temporal pattern of LH secretion: regulation by multiple ultradian oscillators versus a single circadian oscillator

The possibility that the 24h rhythm output is the composite expression of ultradian oscillators of varying periodicities was examined by assessing the effect of external continuously or pulsed (20-minute) Gonadotropinreleasing hormone (GnRH) infusions on in vitro luteinizing hormone (LH) release patterns from female mouse pituitaries during 38h study spans. Applying stepwise analyses (spectral, cosine fit, best-fit curve, and peak detection analyses) revealed the waveform shape of LH release output patterns over time is composed of several ultradian oscillations of different periods. The results further substantiated previous observations indicating the pituitary functions as an autonomous clock. The GnRH oscillator functions as a pulse generator and amplitude regulator, but it is not the oscillator that drives the ultradian LH release rhythms. At different stages of the estrus cycle, the effect of GnRH on the expression of ultradian periodicities varies, resulting in the modification of their amplitudes but not their periods. The functional output from the system of ultradian oscillators may superimpose a “circadian or infradian phenotype” on the observed secretion pattern. An “amplitude control” hypothesis is proposed: The temporal pattern of LH release is governed by several oscillators that function in conjunction with one another and are regulated by an amplitude-controlled mechanism. Simulated models show that such a mechanism results in better adaptive response to environmental requirements than does a single circadian oscillator. (Chronobiology International, 18(3), 399-412, 2001)     

The daily pattern of nitrogen uptake by phytoplankton in dynamic mixed layer environments

The transport and assimilation of the various forms of biologically available nitrogen by phytoplankton, and the subsequent biosynthesis of N-containing macromolecules, have the potential to respond in different ways during the daily growth cycle. This review examines five types of effect that may influence the daily pattern of nitrogen uptake and metabolism: light versus dark (the day/night cycle); changes in irradiance during the day (including the diurnal rise and fall in photon fluence rates); circadian rhythms (endogenous patterns of variation which may continue in the absence of external environmental forcing); periodic variations in exogenous nitrogen supply; and the 24-hour dynamics of stratification and mixing. The hydrodynamic effects operate through a variety of direct and indirect controls, and can substantially modify the diel rhythmicity of phytoplankton growth.