The phase response curve is measured by delivering a precisely timed perturbation to an oscillation and measuring the effect on the cycle period. The infinitesimal phase response curve iprc is a fundamental tool in the study of neuronal synchrony schultheiss et al. We rely upon the direct measurement of the phase response curves prcs to a stim. The frequency response hjw is in general is complex, with real and imaginary parts. The transient prcs are closely matched by steadystate prcs if the stimulus changes sufficiently slowly. The amount of phase delay increases until about eight hours after wakeup time, when the effect swings abruptly from. The benchmark response is defined as 10% or 5%, or 1%.
Estimation of the phase response curve from parkinsonian tremor tabish a. Prc describes the phase shift of the oscillation in re sponse to a perturbing pulse at. Cholinergic neuromodulation changes phase response curve shape and type in cortical pyramidal neurons klaus m. Phase response curve and phase transition curve to understand circadian entrainment, it is useful to understand the perturbing effect of a single stimulus e. Prc estimates, in turn, provide an intuitive insight into how oscillatory systems become entrained and how they can be desynchronized. Oct 21, 2011 a phase response curve prc, also called a phase resetting curve, is measured by delivering a precisely timed perturbation to an oscillation and measuring the effect on the cycle period. The main difference between the methods lies in the reliability which is.
Strong phase response curve for the circadian rhythm of. The response of a system to sinusoidal input gives valuable information about its behavior in the frequency domain, similar to convolution in the time domain. The prc describes the phase shift of the oscillation in response to a perturbing pulse at each phase of the oscillation. Winfrees analysis involves the appli cation of perturbing stimuli at different points in the pacemakercycle. Cholinergic neuromodulation changes phase response curve. Phase response curve prc, also known as phase resetting or phase sensitivity curve, is a basic characteristic of a limit cycle oscillator. For example, if pij represents the effect of 53bp1 on s phase, then all of the s phase transition reactions e. An interactive tool for visualizing, simulating, and perturbing cellcycle progression david borland,1,6 hong yi,1,6 gavin d. An important characteristic of the limit cycle oscillator resulting from the phase reduction procedure is its phaseresponse or phaseresetting curve prc.
Discontinuities in phase resetting experiments leon glass and a. Stiefel abstract the phase response curve prc is an important tool to determine the excitability type of single neurons which reveals consequences for their synchronizing properties. Waterhouse i and anna wirzjusticg 1department of physiological sciences, university of manchester, manchester u. The mouse clock mutation reduces circadian pacemaker. Computation of phase response curves via a direct method adapted. Amplifiers produce gain while filters alter the amplitude andor phase.
If the phase shifts obtained by a perturbing signal are plotted against circadian time, the result is a phase response curve. Craig 6 if the input to this system is a sine wave, the steadystate output after the transients have died out is also a sine wave with the same frequency, but with a different amplitude and phase angle. A new approach for determining phase response curves. Phase response curve can be applied to any oscillatory system to describe how a perturbation changes the phase of the oscillation. Impulse responses of automaticity in the purkinje fiber.
Toourknowledge,noother drughasbeenfoundthat mimickstheeffects oflight onthe clocksocloselyin gonyaulax. Spectral reconstruction of phase response curves reveals the synchronization properties of mouse globus pallidus neurons joshua a. Phase response curves prcs are widely used in circadian clocks, neuroscience, and heart physiology. Amplifiers and filters are widely used electronic circuits that have the properties of amplification and filtration, hence their names. By statistical calculation an upper 95% confidence limit of the curve is determined 3. Thus, large amounts of data are necessary to determine the prc with the direct method. Effects of synaptic plasticity on phase and period locking in. Prcs measure the phaseshift resulting from perturbing the oscillator with a brief stimulus at different times of the cycle. This paper will focus on applying stimulation to modulate synchrony in chemical oscillators. Inferring the phase response curve from observation of a. They have been extensively used to understand the synchronous activity patterns emerging from a network of weakly coupled oscillators. To understand and explore this process, we developed the cell cycle browser ccb, an interactive web interface based on realtime reporter data collected in proliferating human cells.
The phase response curve prc is a tool used in neuroscience that measures the. A comparison of methods to determine neuronal phaseresponse. The phase and amplitude of the resulting curves depended on the variable affected, the sign of the perturbing stimulus and its strength. Phase resetting and annihilation pacemaker activity in. A new approach for determining phase response curves reveals that purkinje cells can act as perfect integrators elena phoka1,2. Desynchronization of stochastically synchronized chemical. Edwards, panagiotis kassavetis, and tom gilbertson2,3 1sobell department of motor neuroscience and movement disorders, university college london institute of neurology, london, united kingdom. Jun, 2006 the mouse clock gene encodes a basic helixloophelixpas transcription factor, clock, that acts in concert with bmal1 to form the positive elements of the circadian clock mechanism in mammals.
The phaseresponse curve prc is an important tool to determine the excitability type of single neurons which reveals consequences for their synchronizing properties. The function zis called the phase response curve prc and describes the magnitude of phase changes after perturbing an oscillatory. Phasedependent transient responses of respiratory motor. Phase response curves obtained by perturbing different variables. The benchmark dose corresponds to the bench mark response on the upper confidence limit curve. Computation of phase response curves via a direct method. Phase response curves prcs are widely used in circadian clocks, neurosci ence, and heart. Phase response curve circadian sleep disorders network. Spectral reconstruction of phase response curves reveals. This curve describes variation of the phase j of the system in response to a weak external perturbation p t. A useful measurable property of a neural oscillator is its phase response curve prc. Frequency response of an amplifier or filter shows how the gain of the output responds to input signals at different frequencies. Apparent discontinuities in the phase resetting response of cardiac pacemakers t. Phase response curves in neuroscience springerlink.
A phase response curves prcs of rat1 mper2luc cells treated with five different. Neuronal phase response curves prcs summarize the relationship between the timing of inputs within a neurons spike cycle and the consequent shifts in output spike timing. A comparison of methods to determine neuronal phase response curves ben torbennielsen, marylka uusisaari, klaus m. The consequences of these results for interpreting the molecular mechanism of circadian clocks on the basis of phase response curves to. The phase response curve prc is a measure of the spike time shift caused by perturbations of the membrane potential as a function of the phase of the spike cycle of a neuron. Edwards, panagiotis kassavetis, and tom gilbertson2,3 1sobell department of motor neuroscience and movement disorders, university college london institute of neurology.
A comparison of methods to determine neuronal phaseresponse curves. Above the critical line red dashed line of fig 4b, the perturbed phases are. The resulting curve is called a phase response curve prc. Here we will give rules that can be used to eyeball phase, given only a polezero plot. A phaseresponse curve prc is a plot of the magnitude and direction of the time shift induced by an environmental perturbation as a function of the circadian time at which the perturbation is given. The phase response curves of perturbations of form b resemble very much the perturbations of form a when applying the perturbation during the middle of the delay or advance part of the oscillation. Kinetics the cycloheximideinduced phase changesinthe. University of groningen latitudinal differences in the.
The phase response curve prc is a curve describing the relationship between a stimulus, such as light exposure, and a response, in this case a shift in circadian rhythm phase shift. Increasing the length of the pulse has nearly the same effect as increasing the strength of the perturbation. A phase response curve prc tabulates the transient change in the cycle period of an oscillator induced by a perturbation as a function of the phase at which it is. Characteristics the discrete stimuli often come in the form of light pulses, which induce phase delays and phase advances during early and late subjective night, respectively, but produce negligible phase shifts. Description of phase resetting characteristics have focused on only one phase of the respiratory oscillator, i. The form of a neurons prc reflects its mechanism of spike initiation or excitability as well as other influences of membrane conductances on synaptic integration. The cell cycle is driven by precise temporal coordination among many molecular activities. This approach complements the infinitesimal phase response curve approach. Labelfree noninvasive imaging technique non perturbing. Pdf phase response curves prcs are widely used in circadian clocks, neuroscience, and heart physiology. A phase response curve and b phase transition curve described by eqs. Principles and practice of sleep medicine fifth edition, 2011. An examplephase response curve of cellular circadian rhythms 2 1. Phase response curve prc in a graph that plots the magnitude of phase shifts resulting from perturbations from discrete stimuli.
At some points in the subjective day there are little or no shifts in the phase. Dose response curve phase fluorescence correlation and more. A phase response curve prc tabulates the transient change in the cycle. Abstract the phaseresponse curve prc is an important tool to determine the excitabil ity type of single neurons which reveals consequences for their synchronizing prop erties.
This is often more useful and intuitive when expressed in polar coordinate. Phase response curves prcs, characterizing the response of an oscillator to weak external perturbation, have been estimated from a broad range of biological oscillators, including single neurons in vivo. A geometric approach to phase response curves and its. Phase response curves obtained by perturbing different. Spectral reconstruction of phase response curves reveals the. A phase response curve prc illustrates the transient change phase response in the cycle period of an oscillation induced by a perturbation as a function of the phase at which it is received. Frequency response analysis of amplifiers and filters. The prc describes the dependence of the phase shift of the oscillation in response to a small perturbing pulse at each phase of the oscillation.
Mar 07, 2016 this video was made by two trainees at ucsds center for circadian biology ben sheredos and liz harrison for use in the undergraduate class on circadian rhythms and biological clocks at ucsd. Craig 9 when one has the frequency response curves for any system and is given a specific sinusoidal input, it is an easy calculation to get the sinusoidal output. The relationship between old phase and phase shift characterizes the respiratory oscillator, and called as a phase response curve. Phase response curve prc, also known as phase resetting or phase sensitivity curve, is a basic characteristic of a limit cycle oscillator14. Butera 1laboratory for neuroengineering, georgia institute of technology, atlanta, georgia. The original clock mutant allele is a dominant negative antimorphic mutation that deletes exon 19 and causes an internal deletion of 51 aa in the cterminal activation domain of the clock protein. The effect of such single stimulus depends on the phase. Sensitivity analysis of oscillator models in the space of phase.
They quantify the response of an oscillator to pulselike perturbations. Prcs measure the phase shift resulting from perturbing the oscillator with a brief stimulus at different times of the cycle. Phase response curves are also a clear way to summarize the interactions between the intrinsic membrane properties, the mean firing rate, and the effects of transient inputs on spike timing. Singularly perturbed phase response curves for relaxation. The phase response curve as a measure of neural response. We would like to show you a description here but the site wont allow us. James surmeier1 1department of physiology, feinberg school of medicine, northwestern university, chicago, illinois. Introduction the phaseresponse curve prc of a regularly. A skeleton protocola minimal phase response curve recipe 12 3. Eigenfunctions and vector plots are used to explore this frequency response. Phase response curves if organisms kept in constant conditions are exposed to a single perturbing signals a light pulse introduced to organisms kept in constant dark will be used as an example at different times over a 24hour period, the phase shift caused is not always the same. Rensing department of biology, university of bremen, d2800 bremen, federal republic of germany received 23 april 1981 we have used computer analysis of a modified version of. Phase response curve an overview sciencedirect topics.
Inferring the phase response curve from observation of a continuously perturbed oscillator. The influence of stationary synaptic activity on the prc. Estimation of the phase response curve from parkinsonian tremor. When the oscillator is perturbed with a transient external stimulus, the.
Spike generation in cortical neurons depends on the interplay between diverse intrinsic conductances. Neural oscillators are usually perturbed by a change in a. A comparison of methods to determine neuronal phase. In general, the prc can be computed numerically for model neurons or experimentally for biological neurons by injecting a brief.
Examples of biological oscillations are the heartbeat, circadian rhythms, and the regular, repetitive firing observed in some neurons in the absence of noise. Histheory, in which techniques of differential topology are used, predicts that if the phase resetting of the pacemaker in re sponse to the perturbation follows cer. The phase response curve prc is an important tool to determine the excitability type of single neurons which reveals consequences for their synchronizing properties. It isproposedthat the early phase changes are caused byalterations in the ion. It is not common to present exper imental results in this format.
Estimation of the phase response curve from parkinsonian. Pdf phase response curves prc, characterising the response of an oscillator to weak external perturbation, have been estimated from a broad range of. That is, we can separate hjw into its magnitude called amplitude response and its phase component called phase response. In physics and computational neuroscience, there have been quite a bit of work for prcs. Near the rheobase, purely positive type i phase response curves are associated with. Earlyphasechangesvaryfrom5hr advanceat circadiantimect20to 1 hrdelayat ct12. Suppose that a neuron is firing regularly with an interspike interval isi t. At spontaneous wakeup time, exogenous externally administered melatonin has a slight phase delaying effect. Phase response curves provide valuable information on the properties of oscillators and their synchronization. Phaseresponse curves give the responses of neurons to.
I do not know the history of the terminology, but it would be nice if someone can fill that up. The study and applications of the prcs are currently receiving growing. The phase response curve prc provides a useful description of the neural response in particular and of nonlinear oscillators in general ermentrout 1996. A phase response curves prcs of rat1 mper2luc cells treated with five. It describes how the phase of an oscillator is affected by brief and vanishingly weak perturbations as a function of the phase at which they were delivered. For each element pij in array p, add pij to each of the reactions for phase transition j. A method for understanding synchronization of adapting neurons jianxia cui, 1carmen c. The phase shifting response of an oscillator is likely to be a characteristic of the phase when the stimulus begins, that is, of the first unperturbed phase to be presented with a stimulus.
The phase response curve prc describes this dependence. Although the prcs for each different species is different, examination of prcs show that there are certain common features. Apparent discontinuities in the phaseresetting response. Transition of phase response properties and singularity in the. The phase response curve for melatonin is roughly twelve hours out of phase with the phase response curve for light. Jun 15, 2003 the phase delays in this study are also consistent in magnitude with those reported to single light pulses in an intensity response curve recently reported by zeitzer et al. Pdf singularly perturbed phase response curves alessio. The consequences of these results for interpreting the molecular mechanism of circadian clocks on the basis of phase response curves to various treatments are discussed.
A plot of such phase changes, both magnitude and sign, as a function of the phase of the oscillation so perturbed, is called a phase response curve prc refs 1, 2. A phase shift in your circadian rhythms means that your bedtime and wakeup time will move earlier in the day phase advance or later in the day phase delay. Effects of synaptic plasticity on phase and period locking. If one uses the midpoint or end of the stimulus as the marker, then one is choosing a phase which has already been perturbed. What is not obvious, but extremely important, is that the frequency response curves are really a.
382 368 470 1443 800 264 984 1319 1190 1194 1186 311 821 755 127 1357 203 407 321 1215 412 775 346 124 615 128 1454 619 529 1317 1494 1513 695 261 526 260 1412 1165 970 121 458 180 1342 768 1023 1156 355 1046 168