Changes in cannabis use among psychotic clients without specialised substance use treatment

The need to address substance use among people with psychosis has been well established. However, treatment studies targeting substance use in this population have reported mixed results. Substance users with psychosis in no or minimal treatment control groups achieve similar reductions in substance use compared to those in more active substance use treatment, suggesting a role for natural recovery from substance use. This meta-analysis aims to quantify the amount of natural recovery from substance use within control groups of treatment studies containing samples of psychotic substance users, with a particular focus on changes in cannabis use. A systematic search was conducted to identify substance use treatment studies. Meta-analyses were performed to quantify reductions in the frequency of substance use in the past 30 days. Significant but modest reductions (mean reduction of 0.3–0.4 SD across the time points) in the frequency of substance use were found at 6 to 24 months follow up. The current study is the first to quantify changes in substance use in samples enrolled in no treatment or minimal treatment control conditions. These findings highlight the potential role of natural recovery from substance use among individuals with psychosis, although they do not rule out effects of regression to the mean. Additionally, the results provide a baseline from which to estimate likely changes or needed effects sizes in intervention studies. Future research is required to identify the processes underpinning these changes, in order to identify strategies that may better support self-management of substance use in people with psychosis.

Low-frequency resistance fluctuations in metal films under current stressing at low temperature (T<0.3Tmelting)

In this paper, we report a systematic study of low frequency 1∕fα resistance fluctuation in thin metal films (Ag on Si) at different stages of damage process when the film is subjected to high current stressing. The resistance fluctuation (noise) measurement was carried out in situ using a small ac bias that has been mixed with the dc stressing current. The experiment has been carried out as a function of temperature in the range of 150–350 K. The experiment establishes that the current stressed film, as it undergoes damage due to various migration forces, develops an additional low-frequency noise spectral power that does not have the usual 1∕f spectral shape. The magnitude of extra term has an activated temperature dependence (activation energy of ≈0.1 eV) and has a 1∕f1.5 spectral dependence. The activation energy is the same as seen from the temperature dependence of the lifetime of the film. The extra 1∕f1.5 spectral power changes the spectral shape of the noise power as the damage process progress. The extra term likely arising from diffusion starts in the early stage of the migration process during current stressing and is noticeable much before any change can be detected in simultaneous resistance measurements. The experiment carried out over a large temperature range establish a strong correlation between the evolution of the migration process in a current stressed film and the low-frequency noise component that is not a 1∕f noise.

Electromagnetic signatures of lightning near the HF frequency band

The dissertation deals with remote narrowband measurements of the electromagnetic radiation emitted by lightning flashes. A lightning flash consists of a number of sub-processes. The return stroke, which transfers electrical charge from the thundercloud to to the ground, is electromagnetically an impulsive wideband process; that is, it emits radiation at most frequencies in the electromagnetic spectrum, but its duration is only some tens of microseconds. Before and after the return stroke, multiple sub-processes redistribute electrical charges within the thundercloud. These sub-processes can last for tens to hundreds of milliseconds, many orders of magnitude longer than the return stroke. Each sub-process causes radiation with specific time-domain characteristics, having maxima at different frequencies. Thus, if the radiation is measured at a single narrow frequency band, it is difficult to identify the sub-processes, and some sub-processes can be missed altogether. However, narrowband detectors are simple to design and miniaturize. In particular, near the High Frequency band (High Frequency, 3 MHz to 30 MHz), ordinary shortwave radios can, in principle, be used as detectors. This dissertation utilizes a prototype detector which is essentially a handheld AM radio receiver. Measurements were made in Scandinavia, and several independent data sources were used to identify lightning sub-processes, as well as the distance to each individual flash. It is shown that multiple sub-processes radiate strongly near the HF band. The return stroke usually radiates intensely, but it cannot be reliably identified from the time-domain signal alone. This means that a narrowband measurement is best used to characterize the energy of the radiation integrated over the whole flash, without attempting to identify individual processes. The dissertation analyzes the conditions under which this integrated energy can be used to estimate the distance to the flash. It is shown that flash-by-flash variations are large, but the integrated energy is very sensitive to changes in the distance, dropping as approximately the inverse cube root of the distance. Flashes can, in principle, be detected at distances of more than 100 km, but since the ground conductivity can vary, ranging accuracy drops dramatically at distances larger than 20 km. These limitations mean that individual flashes cannot be ranged accurately using a single narrowband detector, and the useful range is limited to 30 kilometers at the most. Nevertheless, simple statistical corrections are developed, which enable an accurate estimate of the distance to the closest edge of an active storm cell, as well as the approach speed. The results of the dissertation could therefore have practical applications in real-time short-range lightning detection and warning systems.

Multi frequency interrogation of polypyrrole based gas sensors for organic vapors

Polypyrrole exhibits reversible changes in their direct current resistance on exposure to organic volatiles. However, one needs to employ an array of such sensors to discriminate organic volatiles present in a mixture. Hence, polypyrrole based gas sensor is designed for the detection and discrimination of different organic volatiles. Multi frequency impedance measurement technique is used to detect the organic vapors, such as acetone, ethanol and Isopropyl alcohol, in the gas phase, over a frequency range 10 Hz to 2 MHz. The sensor response is monitored by measuring the changes in its capacitance, resistance and the dissipation factor upon exposure to organic volatiles. It is observed that the capacitive property of the sensor is more sensitive to these volatiles than its resistive property. Each volatile responds to the sensor in terms of dissipation factor at specific frequency and found that the peak magnitude has a linear relationship with their concentrations.

Matching sender and receiver: poikilothermy and frequency tuning in a tree cricket

Animals communicate in non-ideal and noisy conditions. The primary method they use to improve communication efficiency is sender-receiver matching: the receiver's sensory mechanism filters the impinging signal based on the expected signal. In the context of acoustic communication in crickets, such a match is made in the frequency domain. The males broadcast a mate attraction signal, the calling song, in a narrow frequency band centred on the carrier frequency (CF), and the females are most sensitive to sound close to this frequency. In tree crickets, however, the CF changes with temperature. The mechanisms used by female tree crickets to accommodate this change in CF were investigated at the behavioural and biomechanical level. At the behavioural level, female tree crickets were broadly tuned and responded equally to CFs produced within the naturally occurring range of temperatures (18 to 27 degrees C). To allow such a broad response, however, the transduction mechanisms that convert sound into mechanical and then neural signals must also have a broad response. The tympana of the female tree crickets exhibited a frequency response that was even broader than suggested by the behaviour. Their tympana vibrate with equal amplitude to frequencies spanning nearly an order of magnitude. Such a flat frequency response is unusual in biological systems and cannot be modelled as a simple mechanical system. This feature of the tree cricket auditory system not only has interesting implications for mate choice and species isolation but may also prove exciting for bio-mimetic applications such as the design of miniature low frequency microphones.

Energy Reduction in SRAM using Dynamic Voltage and Frequency Management

This paper describes a dynamic voltage frequency control scheme for a 256 X 64 SRAM block for reducing the energy in active mode and stand-by mode. The DVFM control system monitors the external clock and changes the supply voltage and the body bias so as to achieve a significant reduction in energy. The behavioral model of the proposed DVFM control system algorithm is described and simulated in HDL using delay and energy parameters obtained through SPICE simulation. The frequency range dictated by an external controller is 100 MHz to I GHz. The supply voltage of the complete memory system is varied in steps of 50 mV over the range of 500 mV to IV. The threshold voltage range of operation is plusmn100 mV around the nominal value, achieving 83.4% energy reduction in the active mode and 86.7% in the stand-by mode. This paper also proposes a energy replica that is used in the energy monitor subsystem of the DVFM system.

Time-frequency analysis of human motion during rhythmic exercises

Biomechanical signals due to human movements during exercise are represented in time-frequency domain using Wigner Distribution Function (WDF). Analysis based on WDF reveals instantaneous spectral and power changes during a rhythmic exercise. Investigations were carried out on 11 healthy subjects who performed 5 cycles of sun salutation, with a body-mounted Inertial Measurement Unit (IMU) as a motion sensor. Variance of Instantaneous Frequency (I.F) and Instantaneous Power (I.P) for performance analysis of the subject is estimated using one-way ANOVA model. Results reveal that joint Time-Frequency analysis of biomechanical signals during motion facilitates a better understanding of grace and consistency during rhythmic exercise.

Nearly constant switching frequency hysteresis current controller for general n-level inverter fed induction motor drive

A current-error space phasor based hysteresis controller with nearly constant switching frequency is proposed for a general n-level voltage source inverter fed three-phase induction motor drive. Like voltage-controlled space vector PWM (SVPWM), the proposed controller can precisely detect sub-sector changes and for switching it selects only the nearest switching voltage vectors using the information of the estimated fundamental stator voltages along α and β axes. It provides smooth transition between voltage levels, including operation in over modulation region. Due to adjacent switching amongst the nearest switching vectors forming a triangular sub-sector, in which tip of the fundamental stator voltage vector of the machine lies, switching loss is reduced while keeping the current-error space phasor within the varying parabolic boundary. Appropriate dimension and orientation of this parabolic boundary ensures similar switching frequency spectrum like constant switching frequency SVPWM-based induction motor (IM) drive. Inherent advantages of multi-level inverter and space phasor based current hysteresis controller are retained. The proposed controller is simulated as well as implemented on a 5-level inverter fed 7.5 kW open-end winding IM drive.

Time resolved terahertz spectroscopy of low frequency electronic resonances and optical pump-induced terahertz photoconductivity in reduced graphene oxide membrane

Towards ultrafast optoelectronic applications of single and a few layer reduced graphene oxide (RGO), we study time domain terahertz spectroscopy and optical pump induced changes in terahertz conductivity of self-supported RGO membrane in the spectral window of 0.5-3.5 THz. The real and imaginary parts of conductivity spectra clearly reveal low frequency resonances, attributed to the energy gaps due to the van Hove singularities in the density of states flanking the Dirac points arising due to the relative rotation of the graphene layers. Further, optical pump induced terahertz conductivity is positive, pointing to the dominance of intraband scattering processes. The relaxation dynamics of the photo-excited carriers consists of three cooling pathways: the faster (similar to 450 fs) one due to optical phonon emission followed by disorder mediated large momentum and large energy acoustic phonon emission with a time constant of a few ps (called the super-collision mechanism) and a very large time (similar to 100 ps) arising from the deep trap states. The frequency dependence of the dynamic conductivity at different delay times is analyzed in term of Drude-Smith model. (C) 2014 Published by Elsevier Ltd.

Modeling non-stationarity in intensity, duration and frequency of extreme rainfall over India

Significant changes are reported in extreme rainfall characteristics over India in recent studies though there are disagreements on the spatial uniformity and causes of trends. Based on recent theoretical advancements in the Extreme Value Theory (EVT), we analyze changes in extreme rainfall characteristics over India using a high-resolution daily gridded (1 degrees latitude x 1 degrees longitude) dataset. Intensity, duration and frequency of excess rain over a high threshold in the summer monsoon season are modeled by non-stationary distributions whose parameters vary with physical covariates like the El-Nino Southern Oscillation index (ENSO-index) which is an indicator of large-scale natural variability, global average temperature which is an indicator of human-induced global warming and local mean temperatures which possibly indicate more localized changes. Each non-stationary model considers one physical covariate and the best chosen statistical model at each rainfall grid gives the most significant physical driver for each extreme rainfall characteristic at that grid. Intensity, duration and frequency of extreme rainfall exhibit non-stationarity due to different drivers and no spatially uniform pattern is observed in the changes in them across the country. At most of the locations, duration of extreme rainfall spells is found to be stationary, while non-stationary associations between intensity and frequency and local changes in temperature are detected at a large number of locations. This study presents the first application of nonstationary statistical modeling of intensity, duration and frequency of extreme rainfall over India. The developed models are further used for rainfall frequency analysis to show changes in the 100-year extreme rainfall event. Our findings indicate the varying nature of each extreme rainfall characteristic and their drivers and emphasize the necessity of a comprehensive framework to assess resulting risks of precipitation induced flooding. (C) 2014 Elsevier B.V. All rights reserved.

HCN channels enhance spike phase coherence and regulate the phase of spikes and LFPs in the theta-frequency range

What are the implications for the existence of subthreshold ion channels, their localization profiles, and plasticity on local field potentials (LFPs)? Here, we assessed the role of hyperpolarization-activated cyclic-nucleotide-gated (HCN) channels in altering hippocampal theta-frequency LFPs and the associated spike phase. We presented spatiotemporally randomized, balanced theta-modulated excitatory and inhibitory inputs to somatically aligned, morphologically realistic pyramidal neuron models spread across a cylindrical neuropil. We computed LFPs from seven electrode sites and found that the insertion of an experimentally constrained HCN-conductance gradient into these neurons introduced a location- dependent lead in the LFP phase without significantly altering its amplitude. Further, neurons fired action potentials at a specific theta phase of the LFP, and the insertion of HCN channels introduced large lags in this spike phase and a striking enhancement in neuronal spike-phase coherence. Importantly, graded changes in either HCN conductance or its half-maximal activation voltage resulted in graded changes in LFP and spike phases. Our conclusions on the impact of HCN channels on LFPs and spike phase were invariant to changes in neuropil size, to morphological heterogeneity, to excitatory or inhibitory synaptic scaling, and to shifts in the onset phase of inhibitory inputs. Finally, we selectively abolished the inductive lead in the impedance phase introduced by HCN channels without altering neuronal excitability and found that this inductive phase lead contributed significantly to changes in LFP and spike phase. Our results uncover specific roles for HCN channels and their plasticity in phase-coding schemas and in the formation and dynamic reconfiguration of neuronal cell assemblies.

An evaluation of length-frequency sampling procedures and subsequent data analysis for purse seine-caught yellowfin tuna in the Eastern Pacific Ocean

ENGLISH: A two-stage sampling design is used to estimate the variances of the numbers of yellowfin in different age groups caught in the eastern Pacific Ocean. For purse seiners, the primary sampling unit (n) is a brine well containing fish from a month-area stratum; the number of fish lengths (m) measured from each well are the secondary units. The fish cannot be selected at random from the wells because of practical limitations. The effects of different sampling methods and other factors on the reliability and precision of statistics derived from the length-frequency data were therefore examined. Modifications are recommended where necessary. Lengths of fish measured during the unloading of six test wells revealed two forms of inherent size stratification: 1) short-term disruptions of existing pattern of sizes, and 2) transition zones between long-term trends in sizes. To some degree, all wells exhibited cyclic changes in mean size and variance during unloading. In half of the wells, it was observed that size selection by the unloaders induced a change in mean size. As a result of stratification, the sequence of sizes removed from all wells was non-random, regardless of whether a well contained fish from a single set or from more than one set. The number of modal sizes in a well was not related to the number of sets. In an additional well composed of fish from several sets, an experiment on vertical mixing indicated that a representative sample of the contents may be restricted to the bottom half of the well. The contents of the test wells were used to generate 25 simulated wells and to compare the results of three sampling methods applied to them. The methods were: (1) random sampling (also used as a standard), (2) protracted sampling, in which the selection process was extended over a large portion of a well, and (3) measuring fish consecutively during removal from the well. Repeated sampling by each method and different combinations indicated that, because the principal source of size variation occurred among primary units, increasing n was the most effective way to reduce the variance estimates of both the age-group sizes and the total number of fish in the landings. Protracted sampling largely circumvented the effects of size stratification, and its performance was essentially comparable to that of random sampling. Sampling by this method is recommended. Consecutive-fish sampling produced more biased estimates with greater variances. Analysis of the 1988 length-frequency samples indicated that, for age groups that appear most frequently in the catch, a minimum sampling frequency of one primary unit in six for each month-area stratum would reduce the coefficients of variation (CV) of their size estimates to approximately 10 percent or less. Additional stratification of samples by set type, rather than month-area alone, further reduced the CV's of scarce age groups, such as the recruits, and potentially improved their accuracy. The CV's of recruitment estimates for completely-fished cohorts during the 198184 period were in the vicinity of 3 to 8 percent. Recruitment estimates and their variances were also relatively insensitive to changes in the individual quarterly catches and variances, respectively, of which they were composed. SPANISH: Se usa un diseño de muestreo de dos etapas para estimar las varianzas de los números de aletas amari11as en distintos grupos de edad capturados en el Océano Pacifico oriental. Para barcos cerqueros, la unidad primaria de muestreo (n) es una bodega de salmuera que contenía peces de un estrato de mes-área; el numero de ta11as de peces (m) medidas de cada bodega es la unidad secundaria. Limitaciones de carácter practico impiden la selección aleatoria de peces de las bodegas. Por 10 tanto, fueron examinados los efectos de distintos métodos de muestreo y otros factores sobre la confiabilidad y precisión de las estadísticas derivadas de los datos de frecuencia de ta11a. Se recomiendan modificaciones donde sean necesarias. Las ta11as de peces medidas durante la descarga de seis bodegas de prueba revelaron dos formas de estratificación inherente por ta11a: 1) perturbaciones a corto plazo en la pauta de ta11as existente, y 2) zonas de transición entre las tendencias a largo plazo en las ta11as. En cierto grado, todas las bodegas mostraron cambios cíclicos en ta11a media y varianza durante la descarga. En la mitad de las bodegas, se observo que selección por ta11a por los descargadores indujo un cambio en la ta11a media. Como resultado de la estratificación, la secuencia de ta11as sacadas de todas las bodegas no fue aleatoria, sin considerar si una bodega contenía peces de un solo lance 0 de mas de uno. El numero de ta11as modales en una bodega no estaba relacionado al numero de lances. En una bodega adicional compuesta de peces de varios lances, un experimento de mezcla vertical indico que una muestra representativa del contenido podría estar limitada a la mitad inferior de la bodega. Se uso el contenido de las bodegas de prueba para generar 25 bodegas simuladas y comparar los resultados de tres métodos de muestreo aplicados a estas. Los métodos fueron: (1) muestreo aleatorio (usado también como norma), (2) muestreo extendido, en el cual el proceso de selección fue extendido sobre una porción grande de una bodega, y (3) medición consecutiva de peces durante la descarga de la bodega. EI muestreo repetido con cada método y distintas combinaciones de n y m indico que, puesto que la fuente principal de variación de ta11a ocurría entre las unidades primarias, aumentar n fue la manera mas eficaz de reducir las estimaciones de la varianza de las ta11as de los grupos de edad y el numero total de peces en los desembarcos. El muestreo extendido evito mayormente los efectos de la estratificación por ta11a, y su desempeño fue esencialmente comparable a aquel del muestreo aleatorio. Se recomienda muestrear con este método. El muestreo de peces consecutivos produjo estimaciones mas sesgadas con mayores varianzas. Un análisis de las muestras de frecuencia de ta11a de 1988 indico que, para los grupos de edad que aparecen con mayor frecuencia en la captura, una frecuencia de muestreo minima de una unidad primaria de cada seis para cada estrato de mes-área reduciría los coeficientes de variación (CV) de las estimaciones de ta11a correspondientes a aproximadamente 10% 0 menos. Una estratificación adicional de las muestras por tipo de lance, y no solamente mes-área, redujo aun mas los CV de los grupos de edad escasos, tales como los reclutas, y mejoró potencialmente su precisión. Los CV de las estimaciones del reclutamiento para las cohortes completamente pescadas durante 1981-1984 fueron alrededor de 3-8%. Las estimaciones del reclutamiento y sus varianzas fueron también relativamente insensibles a cambios en las capturas de trimestres individuales y las varianzas, respectivamente, de las cuales fueron derivadas. (PDF contains 70 pages)

Damage detection in civil structures using high-frequency seismograms

The dynamic properties of a structure are a function of its physical properties, and changes in the physical properties of the structure, including the introduction of structural damage, can cause changes in its dynamic behavior. Structural health monitoring (SHM) and damage detection methods provide a means to assess the structural integrity and safety of a civil structure using measurements of its dynamic properties. In particular, these techniques enable a quick damage assessment following a seismic event. In this thesis, the application of high-frequency seismograms to damage detection in civil structures is investigated.

Two novel methods for SHM are developed and validated using small-scale experimental testing, existing structures in situ, and numerical testing. The first method is developed for pre-Northridge steel-moment-resisting frame buildings that are susceptible to weld fracture at beam-column connections. The method is based on using the response of a structure to a nondestructive force (i.e., a hammer blow) to approximate the response of the structure to a damage event (i.e., weld fracture). The method is applied to a small-scale experimental frame, where the impulse response functions of the frame are generated during an impact hammer test. The method is also applied to a numerical model of a steel frame, in which weld fracture is modeled as the tensile opening of a Mode I crack. Impulse response functions are experimentally obtained for a steel moment-resisting frame building in situ. Results indicate that while acceleration and velocity records generated by a damage event are best approximated by the acceleration and velocity records generated by a colocated hammer blow, the method may not be robust to noise. The method seems to be better suited for damage localization, where information such as arrival times and peak accelerations can also provide indication of the damage location. This is of significance for sparsely-instrumented civil structures.

The second SHM method is designed to extract features from high-frequency acceleration records that may indicate the presence of damage. As short-duration high-frequency signals (i.e., pulses) can be indicative of damage, this method relies on the identification and classification of pulses in the acceleration records. It is recommended that, in practice, the method be combined with a vibration-based method that can be used to estimate the loss of stiffness. Briefly, pulses observed in the acceleration time series when the structure is known to be in an undamaged state are compared with pulses observed when the structure is in a potentially damaged state. By comparing the pulse signatures from these two situations, changes in the high-frequency dynamic behavior of the structure can be identified, and damage signals can be extracted and subjected to further analysis. The method is successfully applied to a small-scale experimental shear beam that is dynamically excited at its base using a shake table and damaged by loosening a screw to create a moving part. Although the damage is aperiodic and nonlinear in nature, the damage signals are accurately identified, and the location of damage is determined using the amplitudes and arrival times of the damage signal. The method is also successfully applied to detect the occurrence of damage in a test bed data set provided by the Los Alamos National Laboratory, in which nonlinear damage is introduced into a small-scale steel frame by installing a bumper mechanism that inhibits the amount of motion between two floors. The method is successfully applied and is robust despite a low sampling rate, though false negatives (undetected damage signals) begin to occur at high levels of damage when the frequency of damage events increases. The method is also applied to acceleration data recorded on a damaged cable-stayed bridge in China, provided by the Center of Structural Monitoring and Control at the Harbin Institute of Technology. Acceleration records recorded after the date of damage show a clear increase in high-frequency short-duration pulses compared to those previously recorded. One undamage pulse and two damage pulses are identified from the data. The occurrence of the detected damage pulses is consistent with a progression of damage and matches the known chronology of damage.

Response of linear, viscous damped systems to excitations having time-varying frequency

The response of linear, viscous damped systems to excitations having time-varying frequency is the subject of exact and approximate analyses, which are supplemented by an analog computer study of single degree of freedom system response to excitations having frequencies depending linearly and exponentially on time.

The technique of small perturbations and the methods of stationary phase and saddle-point integration, as well as a novel bounding procedure, are utilized to derive approximate expressions characterizing the system response envelope—particularly near resonances—for the general time-varying excitation frequency.

Descriptive measurements of system resonant behavior recorded during the course of the analog study—maximum response, excitation frequency at which maximum response occurs, and the width of the response peak at the half-power level—are investigated to determine dependence upon natural frequency, damping, and the functional form of the excitation frequency.

The laboratory problem of determining the properties of a physical system from records of its response to excitations of this class is considered, and the transient phenomenon known as “ringing” is treated briefly.

It is shown that system resonant behavior, as portrayed by the above measurements and expressions, is relatively insensitive to the specifics of the excitation frequency-time relation and may be described to good order in terms of parameters combining system properties with the time derivative of excitation frequency evaluated at resonance.

One of these parameters is shown useful for predicting whether or not a given excitation having a time-varying frequency will produce strong or subtle changes in the response envelope of a given system relative to the steady-state response envelope. The parameter is shown, additionally, to be useful for predicting whether or not a particular response record will exhibit the “ringing” phenomenon.

The ionic basis of frequency selectivity in hair cells of the bullfrog's sacculus

Hair cells from the bull frog's sacculus, a vestibular organ responding to substrate-borne vibration, possess electrically resonant membrane properties which maximize the sensitivity of each cell to a particular frequency of mechanical input. The electrical resonance of these cells and its underlying ionic basis were studied by applying gigohm-seal recording techniques to solitary hair cells enzymatically dissociated from the sacculus. The contribution of electrical resonance to frequency selectivity was assessed from microelectrode recordings from hair cells in an excised preparation of the sacculus.

Electrical resonance in the hair cell is demonstrated by damped membrane-potential oscillations in response to extrinsic current pulses applied through the recording pipette. This response is analyzed as that of a damped harmonic oscillator. Oscillation frequency rises with membrane depolarization, from 80-160 Hz at resting potential to asymptotic values of 200-250 Hz. The sharpness of electrical tuning, denoted by the electrical quality factor, Q_e, is a bell-shaped function of membrane voltage, reaching a maximum value around eight at a membrane potential slightly positive to the resting potential.

In whole cells, three time-variant ionic currents are activated at voltages more positive than -60 to -50 mV; these are identified as a voltage-dependent, non-inactivating Ca current (I_ca), a voltage-dependent, transient K current (I_a), and a Ca-dependent K current (I_c). The C channel is identified in excised, inside-out membrane patches on the basis of its large conductance (130-200 pS), its selective permeability to Kover Na or Cl, and its activation by internal Ca ions and membrane depolarization. Analysis of open- and closed-lifetime distributions suggests that the C channel can assume at least two open and three closed kinetic states.

Exposing hair cells to external solutions that inhibit the Ca or C conductances degrades the electrical resonance properties measured under current-clamp conditions, while blocking the A conductance has no significant effect, providing evidence that only the Ca and C conductances participate in the resonance mechanism. To test the sufficiency of these two conductances to account for electrical resonance, a mathematical model is developed that describes I_ca, I_c, and intracellular Ca concentration during voltage-clamp steps. I_ca activation is approximated by a third-order Hodgkin-Huxley kinetic scheme. Ca entering the cell is assumed to be confined to a small submembrane compartment which contains an excess of Ca buffer; Ca leaves this space with first-order kinetics. The Ca- and voltage-dependent activation of C channels is described by a five-state kinetic scheme suggested by the results of single-channel observations. Parameter values in the model are adjusted to fit the waveforms of I_ca and I_c evoked by a series of voltage-clamp steps in a single cell. Having been thus constrained, the model correctly predicts the character of voltage oscillations produced by current-clamp steps, including the dependencies of oscillation frequency and Q_e on membrane voltage. The model shows quantitatively how the Ca and C conductances interact, via changes in intracellular Ca concentration, to produce electrical resonance in a vertebrate hair cell.