Range Of Cycle Periods Research Articles

Noise resistant synchronization and collective rhythm switching in a model of animal group locomotion.

Biology is suffused with rhythmic behaviour, and interacting biological oscillators often synchronize their rhythms with one another. Colonies of some ant species are able to synchronize their activity to fall into coherent bursts, but models of this phenomenon have neglected the potential effects of intrinsic noise and interspecific differences in individual-level behaviour. We investigated the individual and collective activity patterns of two Leptothorax ant species. We show that in one species (Leptothorax sp. W), ants converge onto rhythmic cycles of synchronized collective activity with a period of about 20 min. A second species (Leptothorax crassipilis) exhibits more complex collective dynamics, where dominant collective cycle periods range from 16 min to 2.8 h. Recordings that last 35 h reveal that, in both species, the same colony can exhibit multiple oscillation frequencies. We observe that workers of both species can be stimulated by nest-mates to become active after a refractory resting period, but the durations of refractory periods differ between the species and can be highly variable. We model the emergence of synchronized rhythms using an agent-based model informed by our empirical data. This simple model successfully generates synchronized group oscillations despite the addition of noise to ants' refractory periods. We also find that adding noise reduces the likelihood that the model will spontaneously switch between distinct collective cycle frequencies.

Brainstem Steering of Locomotor Activity in the Newborn Rat.

Control of locomotion relies on motor loops conveying modulatory signals between brainstem and spinal motor circuits. We investigated the steering control of the brainstem reticular formation over the spinal locomotor networks using isolated brainstem-spinal cord preparations of male and female neonatal rats. First, we performed patch-clamp recordings of identified reticulospinal cells during episodes of fictive locomotion. This revealed that a spinal ascending phasic modulation of reticulospinal cell activity is already present at birth. Half of the cells exhibited tonic firing during locomotion, while the other half emitted phasic discharges of action potentials phase locked to ongoing activity. We next showed that mimicking the phasic activity of reticulospinal neurons by applying patterned electrical stimulation bilaterally at the ventral caudal medulla level triggered fictive locomotion efficiently. Moreover, the brainstem stimuli-induced locomotor rhythm was entrained in a one-to-one coupling over a range of cycle periods (2-6 s). Additionally, we induced turning like motor outputs by either increasing or decreasing the relative duration of the stimulation trains on one side of the brainstem compared to the other. The ability of the patterned descending command to control the locomotor output depended on the functional integrity of ventral reticulospinal pathways and the involvement of local spinal central pattern generator circuitry. Altogether, this study provides a mechanism by which brainstem reticulospinal neurons relay steering and speed commands to the spinal locomotor networks.SIGNIFICANCE STATEMENT Locomotor function allows the survival of most animal species while sustaining the expression of fundamental behaviors. Locomotor activities adapt from moment to moment to behavioral and environmental changes. We show that the brainstem can control the spinal locomotor network outputs through phasic descending commands that alternate bilaterally. Manipulating the periodicity and/or the relative durations of the left and right descending commands at the brainstem level is efficient to set the locomotor speed and sustain directional changes.

Magnetic cycles and rotation periods of late-type stars from photometric time series

We investigate the photometric modulation induced by magnetic activity cycles and study the relationship between rotation period and activity cycle(s) in late-type (FGKM) stars. We analyse light-curves spanning up to 9 years of 125 nearby stars provided by the ASAS survey. The sample is mainly conformed by low-activity main sequence late A to mid M-type stars. A search is performed for short (days) and long-term (years) periodic variations in the photometry. We modelled with combinations of sinusoids the light-curves to measure the properties of these periodic signals. To provide a better statistical interpretation of our results we complement them with the results from previous similar works. We have been able to measure long-term photometric cycles of 47 stars. Rotational modulation was also detected and rotational periods measured in 36 stars. For 28 stars we have simultaneous measurements of both, activity cycles and rotational periods, being 17 of them M-type stars. From sinusoidal fits we measured both photometric amplitudes and periods. The measured cycle periods range from 2 up to 14 yr with photometric amplitudes in the range of 5-20 mmag. We have found that the distribution of cycle lengths for the different spectral types is similar. On the other hand the distribution of rotation periods is completely different, trending to longer periods for later type stars. The amplitudes induced by magnetic cycles and rotation show a clear correlation. A trend of photometric amplitudes with rotation period is also outlined in the data. For a given activity index the amplitudes of the photometric variability induced by activity cycles of main sequence GK stars are lower than those of early and mid-M dwarfs. Using spectroscopic data we also provide an update in the empirical relationship between the level of chromospheric activity as given by log(Rhk) and the rotation periods.

DISCOVERY OF A 1.6 YEAR MAGNETIC ACTIVITY CYCLE IN THE EXOPLANET HOST STAR ι HOROLOGII

The Mount Wilson Ca HK survey revealed magnetic activity variations in a large sample of solar-type stars with timescales ranging from 2.5 to 25 years. This broad range of cycle periods is thought to reflect differences in the rotational properties and the depths of the surface convection zones for stars with various masses and ages. In 2007 we initiated a long-term monitoring campaign of Ca II H and K emission for a sample of 57 southern solar-type stars to measure their magnetic activity cycles and their rotational properties when possible. We report the discovery of a 1.6-year magnetic activity cycle in the exoplanet host star iota Horologii, and we obtain an estimate of the rotation period that is consistent with Hyades membership. This is the shortest activity cycle so far measured for a solar-type star, and may be related to the short-timescale magnetic variations recently identified in the Sun and HD49933 from helio- and asteroseismic measurements. Future asteroseismic observations can be compared to those obtained near the magnetic minimum in 2006 to search for cycle-induced shifts in the oscillation frequencies. If such short activity cycles are common in F stars, then NASA's Kepler mission should observe their effects in many of its long-term asteroseismic targets.

Coregulation of ionic currents maintaining the duty cycle of bursting

Central pattern generators (CPGs) are the oscillatory neuronal networks which control rhythmic movements of animals. Some CPGs keep the phase relationships between the neurons’ oscillatory activities over a wide range of the cycle periods. Maintenance of the duty cycle of the bursting activity could be a key feature for a variety of dynamical mechanisms supporting phase constancy in oscillatory neuronal networks. It is a form of cellular homeostasis of neuronal activity. Among other currents, hyperpolarization-activated currents and potassium currents have been shown to be a ubiquitous target for modulation and homeostasis [1,2]. Here we present a novel mechanism of coregulation of currents which preserves duty cycle of bursting activity over a range of cycle periods. We develop a generic lowdimensional Hodgkin-Huxley type model stemming from a model of the leech heart interneuron under certain pharmacological conditions [3]. Application of Co 2+ and 4-AP blocks Ca 2+ currents, the synaptic currents and most of the K + currents. The model contains the slow potassium current (IK2), the fast sodium current (INa). Our new model also includes the hyperpolarization activated current (Ih). Bifurcation theory allows us to make predictions concerning the temporal characteristics of the dynamics of bursting nearby the critical transitions between activities. Shilnikov & Cymblayuk sho wed that the transition from bursting into tonic spiking (blue sky catastrophe) determines the dependence of the burst duration on the voltage of half-activation of IK2 (θ K2 )a s one over square root of the parameter value [4]. Here we show that the half-activation potential of Ih (θ h) controls the interburst interval as one over square root of the parameter value. We investigate the activity of the model to identify mechanisms of coregulation of IK2 and Ih maintaining the duty cycle. Bifurcation analysis of the model was performed using θ K2 and θ h, as controlling parameters. We investigated the temporal characteristics of bursting activity. We identified a saddle node bifurcation for periodic orbits determining the blue sky catastrophe [4] and a saddle node bifurcation for stationary states (SNIC) [5]. We showed the temporal characteristics of bursting depend on the location of the bifurcation curves. By coordinating the two parameters, we were able to increase the period such that the burst duration and interburst interval maintained constant proportion. The coregulation consists of a negative correlation of θ K2 and θ h, which is steeper for the higher duty cycles.

Navigating parameter space between bifurcations to preserve the duty cycle of bursting activity

Event Abstract Back to Event Navigating parameter space between bifurcations to preserve the duty cycle of bursting activity William Barnett1, Martin Anquez2 and Gennady Cymbalyuk2, 3* 1 Georgia State University, Faculty of Electrical Engineering and Computing, United States 2 Georgia State University, Department of Physics and Astronomy, United States 3 Georgia State University, Neuroscience Institute, United States We study the leech heart interneuron, and our model simulates activity in a pharmacologically reduced scenario. Central pattern generators control rhythmic activity in many animals, and neurons in these circuits often conserve functional characteristics, such as duty cycle, over a range of cycle periods. We present a mechanism through which coregulation of the half activation voltage of two conductances in a model of the leech heart interneuron maintains constant duty cycle over a wide range of cycle periods. These currents are a non-inactivating potassium current (IK) and a hyperpolarization activated current (Ih). The proportionality is conserved if the location of two bifurcations is taken into account in coregulation: a saddle-node for periodic orbits and a saddle-node for stationary states. If the shift of the half activation of IK brings the system close to the saddle-node for periodic orbits, the burst duration grows without bound. If the shift of the half activation of Ih brings the system close to the saddle-node for stationary states, the interburst interval grows without bound. In both cases, the cycle period grows proportionally to one over the square root of the control parameter. By varying the two parameters together, the system approaches both bifurcations and burst duration and interburst interval grow proportionally to one another. This mechanism of coregulation maintains constant duty cycle over a threefold range of cycle periods. Supported by NSF PHY-0750456. Conference: 2010 South East Nerve Net (SENN) and Georgia/South Carolina Neuroscience Consortium (GASCNC) conferences, Atlanta , United States, 5 Mar - 7 Mar, 2010. Presentation Type: Oral Presentation Topic: Talks Citation: Barnett W, Anquez M and Cymbalyuk G (2010). Navigating parameter space between bifurcations to preserve the duty cycle of bursting activity. Front. Neurosci. Conference Abstract: 2010 South East Nerve Net (SENN) and Georgia/South Carolina Neuroscience Consortium (GASCNC) conferences. doi: 10.3389/conf.fnins.2010.04.00004 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 15 Mar 2010; Published Online: 15 Mar 2010. * Correspondence: Gennady Cymbalyuk, Georgia State University, Department of Physics and Astronomy, Atlanta, United States, wbarnett2@gsu.edu Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers William Barnett Martin Anquez Gennady Cymbalyuk Google William Barnett Martin Anquez Gennady Cymbalyuk Google Scholar William Barnett Martin Anquez Gennady Cymbalyuk PubMed William Barnett Martin Anquez Gennady Cymbalyuk Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Genetic Ablation of V2a Ipsilateral Interneurons Disrupts Left-Right Locomotor Coordination in Mammalian Spinal Cord

The initiation and coordination of activity in limb muscles are the main functions of neural circuits that control locomotion. Commissural neurons connect locomotor circuits on the two sides of the spinal cord, and represent the known neural substrate for left-right coordination. Here we demonstrate that a group of ipsilateral interneurons, V2a interneurons, plays an essential role in the control of left-right alternation. In the absence of V2a interneurons, the spinal cord fails to exhibit consistent left-right alternation. Locomotor burst activity shows increased variability, but flexor-extensor coordination is unaffected. Anatomical tracing studies reveal a direct excitatory input of V2a interneurons onto commissural interneurons, including a set of molecularly defined V0 neurons that drive left-right alternation. Our findings imply that the neural substrate for left-right coordination consists of at least two components; commissural neurons and a class of ipsilateral interneurons that activate commissural pathways.

Modeling the mammalian locomotor CPG: insights from mistakes and perturbations

A computational model of the mammalian spinal cord circuitry incorporating a two-level central pattern generator (CPG) with separate half-center rhythm generator (RG) and pattern formation (PF) networks is reviewed. The model consists of interacting populations of interneurons and motoneurons described in the Hodgkin-Huxley style. Locomotor rhythm generation is based on a combination of intrinsic (persistent sodium current dependent) properties of excitatory RG neurons and reciprocal inhibition between the two half-centers comprising the RG. The two-level architecture of the CPG was suggested from an analysis of deletions (spontaneous omissions of activity) and the effects of afferent stimulation on the locomotor pattern and rhythm observed during fictive locomotion in the cat. The RG controls the activity of the PF network that in turn defines the rhythmic pattern of motoneuron activity. The model produces realistic firing patterns of two antagonist motoneuron populations and generates locomotor oscillations encompassing the range of cycle periods and phase durations observed during cat locomotion. A number of features of the real CPG operation can be reproduced with separate RG and PF networks, which would be difficult if not impossible to demonstrate with a classical single-level CPG. The two-level architecture allows the CPG to maintain the phase of locomotor oscillations and cycle timing during deletions and during sensory stimulation. The model provides a basis for functional identification of spinal interneurons involved in generation and control of the locomotor pattern.

Synaptic depression in conjunction with A-current channels promote phase constancy in a rhythmic network.

In many central pattern generators, pairs of neurons maintain an approximately fixed phase despite large changes in the frequency. The mechanisms underlying phase maintenance are not clear. Previous theoretical work suggested that inhibitory synapses that show short-term depression could play a critical role in this respect. In this work we examine how the interaction between synaptic depression and the kinetics of a transient potassium (A-like) current could be advantageous for phase constancy in a rhythmic network. To demonstrate the mechanism in the context of a realistic central pattern generator, we constructed a detailed model of the crustacean pyloric circuit. The frequency of the rhythm was modified by changing the level of a ligand-activated current in one of the pyloric neurons. We examined how the time difference of firing activities between two selected neurons in this circuit is affected by synaptic depression, A-current, and a combination of the two. We tuned the parameters of the model such that with synaptic depression alone, or A-current alone, phase was not maintained between these two neurons. However, when these two components came together, they acted synergistically to maintain the phase across a wide range of cycle periods. This suggests that synaptic depression may be necessary to allow an A-current to delay a postsynaptic neuron in a frequency-dependent manner, such that phase invariance is ensured.

Cyclic eruptive behavior of silicic volcanoes

Silicic volcanism seems chaotic: the styles, magnitudes, and/or timing of successive eruptions may be without apparent pattern, or patterns may merge, fade, or abruptly change. However, geophysical monitoring of recent eruptions shows that some silicic volcanoes can exhibit cyclic eruptive behavior wherein periods of explosive activity or rapid extrusion alternate with periods of repose. The cycles are commonly observed in time-averaged amplitudes of eruption-related seismicity and also have been observed in ground-surface tilt data. When tilt and seismicity are both observed during oscillatory behavior, as at Soufriere Hills volcano on Montserrat, British West Indies, they correlate in time. Cycle periods range from hours to days, and cycle amplitudes and waveforms vary widely. Complex oscillatory behavior is also sometimes observed during high-pressure (tens of MPa) extrusion of industrial polymer melts. With this phenomenon as a guide, we construct a simple dynamic model for the oscillatory behavior of erupting volcanoes. We propose that cyclic eruptions result from Newtonian flow of compressible magma through the volcanic conduit combined with a stick-slip condition along the conduit wall, in analogy to the behavior of industrial polymers. If magma is forced into the conduit at a constant rate, pressure and flow rate rise. If the flow rate through the conduit exceeds a threshold value, the flow resistance abruptly drops as the magma slips along a shallow portion of the conduit wall. This reduces resistance to flow and causes the flow rate to jump to a higher value. If this enhanced flow rate exceeds the supply rate, both pressure and flow rate decline as the compressed magma in the conduit expands. Eventually slip ceases as the magma reattaches to the conduit wall at a flow rate less than the supply rate. Consequently pressure begins to increase, and the cycle begins again. This simple model reconciles a variety of disparate phenomena associated with cyclic silicic volcanism and provides a paradigm to interpret cyclic eruptive behavior.

Crawling motor patterns induced by pilocarpine in isolated larval nerve cords of Manduca sexta.

1. Larval crawling is a bilaterally symmetrical behavior that involves an anterior moving wave of motor activity in the body wall muscles in conjunction with sequential movements of the abdominal prolegs and thoracic legs. The purpose of this study was to determine whether the larval CNS by itself and without phasic sensory feedback was capable of producing patterned activity associated with crawling. To establish the extent of similarity between the output of the isolated nerve cord and crawling, the motor activity produced in isolated larval nerve cords was compared with the motor activity from freely crawling larvae. 2. When exposed to the muscarinic receptor agonist pilocarpine (1.0 mM), isolated larval nerve cords produced long-lasting rhythmic activity in the motor neurons that supply the thoracic leg, abdominal body wall, and abdominal proleg muscles. The rhythmic activity evoked by pilocarpine was abolished reversibly and completely by bath application of the muscarinic-receptor antagonist atropine (0.01 mM) in conjunction with pilocarpine (1.0 mM), suggesting that the response was mediated by muscarinic-like acetylcholine receptors. 3. Similar to crawling in intact animals, the evoked activity in isolated nerve cords involved bilaterally symmetrical motor activity that progressed from the most posterior abdominal segment to the most anterior thoracic segment. The rhythmic activity in thoracic leg, abdominal proleg, and abdominal body wall motor neurons showed intrasegmental and intersegmental cycle-to-cycle coupling. The average cycle period for rhythmic activity in the isolated nerve cord was approximately 2.5 times slower than the cycle period for crawling in intact larvae, but not more variable. 4. Like crawling in intact animals, in isolated nerve cords, bursting activity in the dorsal body wall motor neurons occurred before activity in ventral/lateral body wall motor neurons within an abdominal segment. The evoked bursting activity recorded from the proleg nerve was superimposed on a high level of tonic activity. 5. In isolated nerve cords, bursts of activity in the thoracic leg levator/extensor motor neurons alternated with bursts of activity in the depressor/flexor motor neurons. The burst duration of the levator/extensor activity was brief and remained relatively steady as cycle period increased. The burst duration of the depressor/ flexor activity occupied the majority of an average cycle and increased as cycle period increased. The phase of both levator/extensor motor nerve activity and depressor/flexor motor nerve activity remained relatively stable over the entire range of cycle periods. The timing and patterning of thoracic leg motor neuron activity in isolated nerve cords quantitatively resembled thoracic leg motor activity in freely crawling larvae. 6. The rhythmic motor activity generated by an isolated larval nerve cord resembled a slower version of normal crawling in intact larvae. Because of the many similarities between activity induced in the isolated nerve cord and the muscle activity and movements of thoracic and abdominal segments during crawling, we concluded that central mechanisms can establish the timing and patterning of the crawling motor pattern and that crawling may reflect the output of a central pattern generating network.

Experimentally derived model for the locomotor pattern generator in the Xenopus embryo.

1. Simulations of Xenopus embryo spinal neurons were endowed with Hodgkin-Huxley-style models of voltage-dependent Na+, Ca2+, slow K+ and fast K+ currents together with a Na(+)-dependent K+ current. The parameters describing the activation, inactivation and relaxation of these currents were derived from previous voltage-clamp studies of Xenopus embryo spinal neurons. Each of the currents was present at realistic densities. 2. The model neurons fired repetitively in response to current injection. The Ca2+ current was essential for repetitive firing in response to current injection. The fast K+ current appeared mainly to control spike width, whereas the slow K+ current exerted a powerful influence on the reptitive firing properties of the neurons without markedly affecting spike width. 3. The properties of the model neurons could be made more consistent with those previously reported for Xenopus embryo neurons during intracellular recordings in vivo, if the shunting effect of the sharp microelectrode was incorporated into the model. 4. The model neurons were then used to create a simplified version of the spinal network that controls swimming in the frog embryo. This model network could generate the motor pattern for swimming: the activity between the left and right sides alternated with a cycle period that varied from 50 to 120 ms. This is very similar to the range of cycle periods observed in the real embryo. The shunting effect of the microelectrode was once again taken into account. 5. Reductions of the K+ currents perturbed the motor pattern and gave three forms of aberrant motor activity very similar to those previously seen during the application of K+ channel blockers to the real embryo. The ability to generate the correct motor pattern for swimming in the model depended on the balance between the K+ currents and the inward Na+ and Ca2+ currents rather than their absolute values. 6. The model network could generate a motor pattern for swimming over a very wide range of excitatory (2-10 nS) and inhibitory (2-400 nS) synaptic strengths. Rough estimates of the physiological synaptic strengths in the real circuit (around 20-60 nS for inhibition and 2-5 nS for excitation) fall within the range of synaptic strengths that gave simulation of the swimming motor pattern in the model. 7. The cycle period of the motor activity in the model shortened either as the excitatory synapses were strengthened or as the inhibitory synapses were weakened. 8. The prediction that the strength of the mid-cycle inhibition determines cycle period has been tested by using low levels of strychnine to reduce glycinergic reciprocal inhibition in a graded manner in the real embryo. As the inhibition was reduced, the cycle period of fictive swimming in the embryo shortened by amounts very close to those predicted by the model. 9. This new experimentally derived model can replicate many of the known features of fictive swimming in the real embryo and may be of value as an analytical tool in attempting to understand how the spinal circuitry of the Xenopus embryo and related amphibian embryos control a variety of motor behaviours.

Neural network simulations of coupled locomotor oscillators in the lamprey spinal cord.

The segmental locomotor network in the lamprey spinal cord was simulated on a computer using a connectionist-type neural network. The cells of the network were identical except for their excitatory levels and their synaptic connections. The synaptic connections used were based on previous experimental work. It was demonstrated that the connectivity of the circuit is capable of generating oscillatory activity with the appropriate phase relations among the cells. Intersegmental coordination was explored by coupling two identical segmental networks using only the cells of the network. Each of the possible couplings of a bilateral pair of cells in one oscillator with a bilateral pair of cells in the other oscillator produced stable phase locking of the two oscillators. The degree of phase difference was dependent upon synaptic weight, and the operating range of synaptic weights varied among the pairs of connections. The coupling was tested using several criteria from experimental work on the lamprey spinal cord. Coupling schemes involving several pairs of connecting cells were found which 1) achieved steady-state phase locking within a single cycle, 2) exhibited constant phase differences over a wide range of cycle periods, and 3) maintained stable phase locking in spite of large differences in the intrinsic frequencies of the two oscillators. It is concluded that the synaptic connectivity plays a large role in producing oscillations in this network and that it is not necessary to postulate a separate set of coordinating neurons between oscillators in order to achieve appropriate phase coupling.

The development of swimming rhythmicity in post-embryonic Xenopus laevis

The post-embryonic development of 'fictive' swimming in immobilized Xenopus laevis tadpoles has been examined during the first day of larval life. In Xenopus embryos (stage 37-38; Nieuwkoop & Faber 1956), the rhythmic ventral root activity underlying swimming occurs as single brief (ca. 7 ms) compound impulses on each cycle. However, by stage 42 (about 24 h after hatching), ventral root discharge consists of bursts lasting around 20 ms per cycle. In addition to increased burst duration in each cycle of larval swimming, the range of cycle periods within an episode increases, although mean period values (ca. 70-80 ms) remain similar to those of the younger animal. Consequently, motoneurons at developmental stage 42 are active during swimming for a greater percentage (ca. 25%) of cycle time than at stage 37-38 (ca. 10%). Developmental stage 40 (ca. 12 h post-hatching) is an intermediate stage in rhythm development. Ventral root discharge varies from bursts of 10-20 ms at the start of an episode to embryonic (ca. 7 ms) spikes at the end of an episode. Furthermore, discharge varies from bursts of activity in rostral segments of stage 40 larvae to 7 ms spikes more caudally, as in embryos. The data thus suggest that Xenopus swimming rhythmicity develops relatively rapidly, along a rostrocaudal gradient, and may involve acquisition of multiple spiking in spinal neurons.

Patterns of Synaptic Drive to Ventrally Located Spinal Neurones inRana TemporariaEmbryos During Rhythmic And Non-Rhythmic Motor Responses

1. Intracellular recordings have been made from ventrally located neurones in the spinal cord of Rana temporaria embryos at around the time of hatching. Both short-latency 'reflex' and more prolonged rhythmic motor responses can be elicited by stimulation of the skin in immobilized embryos. Initial responses to single-sided skin stimuli usually involve excitation of neurones on the opposite side and strychnine-sensitive inhibition of neurones on the same side. Less reliable responses to dimming the lights also involve initial excitation on one side associated with inhibition on the opposite side. 2. Intracellular recordings from single neurones during rhythmic activity show that on each cycle the same neurone can fire one or many spikes during the course of a single evoked or spontaneous episode. Bursts occur at longer cycle periods, generally at the start of episodes; single spikes occur at shorter cycle periods, generally later in episodes. 3. During sustained rhythmic responses, neuronal membrane potential is generally depolarised and returns gradually to its resting level at the end of the episode. During the episode, relatively depolarising phases of synaptic excitation alternate with relatively hyperpolarising phases of chloride-dependent synaptic inhibition. Cell input resistance is reduced by around 50% throughout each episode. Within each cycle, input resistance is reduced further during the hyperpolarising phase than during the depolarising phase. 4. Rhythmic excitation and inhibition of ventrally located neurones appears to be similar throughout the whole range of cycle periods, supporting the suggestion that a single rhythm-generating system with a wide 'permissive' range drives rhythmic movements in R. temporaria embryos.

Centrally Generated Rhythmic and Non-Rhythmic Behavioural Responses inRana TemporariaEmbryos

Embryos of the frog Rana temporaria up to and around the time of hatching show a range of rhythmic and non-rhythmic movements. These may occur spontaneously or in response to lightly touching the skin of the trunk or head. The first response to touching one side is usually on the opposite side. Non-rhythmic movements range from weak twitches centred on the mid trunk to strong flexions along much of one side of the body and part of the tail, which result in the animal becoming tightly coiled. Rhythmic movements range from slow, high-amplitude 'lashing' movements to faster, lower-amplitude 'swimming' movements. During rhythmic movements, a wave of bending passes along the animal from head to tail. The longitudinal phase delay in bending is constant for a range of cycle periods (88-193 ms) but is not uniform along the whole body. Bending is maximal along the body and rostral part of the tail, decreases towards the tip of the tail and is lowest at the head. Lateral displacement during rhythmic movements is lowest 0.2 body lengths from the snout, increases rostral and caudal to this level and is highest at the tip of the tail. In animals immobilised with curare, a range of patterns of motor discharge can be recorded in response to stimulation. Non-rhythmic responses range from single spikes to prolonged bursts, usually on the opposite side to the stimulus. Stronger bursts can alternate briefly between the two sides and are never synchronous on both. Episodes of sustained rhythmic activity can be evoked by touch, electrical stimulation of the skin or, rarely, dimming the lights. Cycle periods within each episode can vary considerably but often shorten as activity proceeds. Discharge on the two sides alternates (phase is approximately 0.5). Motor root burst duration correlates with cycle period, bursts being longer at longer cycle periods. Burst onset is delayed caudally, this delay being longer at longer cycle periods. Stimulating one side of the head evokes a large burst of discharge on the opposite side, often followed by sustained rhythmic discharge. These responses in immobilised animals are judged to constitute centrally generated correlates of the main behavioural responses of R. temporaria embryos.

Estimating Amplitudes of Fifth-Order Sea Level Fluctuations from Peritidal Through Basinal Carbonate Deposits, Lower Mississippian, Wyoming-Montana: ABSTRACT

Three types of 1-10-m upward-shallowing cycles are observed in the Lower Mississippian Lodgepole and lower Madison formations of Wyoming and Montana. Typical peritidal cycles have pellet grainstone bases overlain by algal laminites, which are rarely capped by paleosol/regolith horizons. Shallow ramp cycles have burrowed pellet-skeletal wackestone bases overlain by cross-bedded ooid/crinoid grainstone caps. Deep ramp cycles are characterized by sub-wave base limestone/argillite, storm-deposited limestone, overlain by hummocky stratified grainstone caps. Average cycle periods range from 17-155 k.y. This, rhythmically bedded limestone/argillite deposits of basinal facies do not contain shallowing-upward cycles, but do contain 2-4 k.y. limestone/argillite rhythms. These sub-wave base deposit are associated with Waulsortian-type mud mounds which have >50 m synoptic relief. This relief provides minimum water depth estimates for the deposits, and implies storm-wave base was less than 50 m. Two-dimensional computer modeling of cyclic platform through noncyclic basinal deposits allows for bracketing of fifth-order sea level fluctuation amplitudes, thought responsible for cycle formation. Computer models using fifth-order amplitudes less than 20 m do not produce cycles on the deep ramp (assuming a 25-30 m storm-wave base). Amplitudes >30 m produce water depths on the inner ramp that are too deep, and disconformities extend too far into themore » basin. The absence of meter-scale cycles in the basin suggests water depths were too great to record the effects of sea level oscillations occurring on the platform, or climatic fluctuation, associated with glacio-eustatic sea level oscillations, were not sufficient to affect hemipelagic depositional patterns in the tropical basin environment.« less

Periodic operation of a trickle‐bed reactor

AbstractThe influence of periodic water flow on SO2 oxidation in a trickle bed of activated carbon catalyst was investigated, whereby gaseous reactants were introduced into the trickle‐bed reactor continuously, but water was turned on and off. Mean liquid superficial velocities of 0.86 and 1.65 mm/s were used. At the latter, an increase in the oxidation rate of about 30 to 45% was found within a range of cycle periods from 2 to 80 min. A temperature change of up to 7°C was observed in the bed during periodic operation. An explanation of the improved average oxidation rate under periodic operation is developed in terms of the steady‐state rates with and without water flow for symmetrical and asymmetrical cycles.

Sea-level curve for Pennsylvanian eustatic marine transgressive-regressive depositional cycles along midcontinent outcrop belt, North America

At least 55 cycles of marine inundation and withdrawal are recognized in the mid-Desmoinesian to mid-Virgilian Midcontinent outcrop sequence in North America. They range from widespread major cycles (classic cyclothems) with deep-water facies extending across the northern shelf, through intermediate cycles persisting as marine horizons across the shelf, to minor cycles developed on the lower shelf or as parts of major cycles. Biostratigraphic differentiation of the cycles should establish interbasinal correlation on a scale fine enough to allow evaluation of differential tectonics and sedimentation. Sequential groupings of cycles are more irregular than proposed megacyclothems or mesothems, but they may be obscured by the distinctness of the major cyclothems. Estimates of cycle periods range from about 40 to 120 x 10/sup 3/ yr for the minor cycles up to about 235 to 400 x 10/sup 3/ yr for the major cyclothems. The range for all cycles corresponds well to the range of periods of Earth's orbital parameters that constitute the Milankovitch insolation theory for the Pleistocene ice ages, and it further supports Gondwanan glacial control for the Pennsylvanian cycles. Even though the dominant period of the major Pennsylvanian cyclothems is up to four times longer than the dominant 100,000-yr period inmore » the Pleistocene, the shapes of both curves display rapid marine transgression (rapid melting of ice caps) and slow interrupted regression (slow buildup of ice caps), which suggest similar linkages between the climatic effects of the orbital parameters and ice-cap formation and melting, at the two different scales, widely separated in time. 28 references, 3 figures.« less

Swimming rhythm in decerebrated, paralyzed stingrays: normal and abnormal coupling

Rhythmic motoneuronal activity was recorded from decerebrated, paralyzed stingrays and compared with electromyograms recorded from the same animals. Before and after paralysis, a rostral-to-caudal sequence of alternation occurred between dorsal (elevator) and ventral (depressor) efferents. The swimming pattern was thus observed in the absence of phasic afferent input, and this constitutes fictive locomotion. After paralysis, both the intersegmental delay (time between activation at progressively caudal recording sites) and the burst duration remained linearly related to the swim cycle period. In many instances, neither the slope nor the intercept was significantly altered by immobilization. The intercepts all fell near the origin, indicating that the fictive rhythm remains constant phase coupled. Although the swimming rhythm was obtained after paralysis, some differences occurred. These included fewer and shorter spontaneous sequences, a greater range of cycle periods, and longer burst durations. During fictive swimming, the burst duration:cycle period ratio usually increased to 0.53 from 0.39 observed before paralysis. Therefore, the silent periods seen between burst discharges in antagonist efferents during movement were often absent after paralysis. Mechanical stimulation of the tail reduced both cycle periods and burst durations; however, the burst:cycle ratio remained greater than or equal to 0.50. The linear relation between burst duration and cycle period found for spontaneous sequences was not changed by stimulation of the tail. During fictive swimming the inter- and intrasegmental coupling that characterizes stingray swimming becomes labile. Abnormal coupling appears more often during sequences with long swim cycles. Intrasegmental coupling is tighter than intersegmental coupling at any cycle period. Rhythmic activity at one segmental level can be independent of activity at other levels. This suggests that multiple oscillator circuits exist that are not dependent on propriospinal circuits interconnecting different segments. Rhythmicity in elevator and depressor motoneurons is not dependent on reciprocal connections between the circuitry driving the motor nuclei. Therefore, separate oscillators for elevators and depressors appear to be present within one spinal segment.