Differences between the channels, currents and mechanisms of conduction slowing/block and accommodative processes in simulated cases of focal demyelinating neuropathies

To clarify the differences between the mechanisms of conduction slowing/block and accommodative processes in focal demyelinating neuropathies, this computational study presents the kinetics of the ionic, transaxonal and transmyelin currents defining the intracellular and electrotonic potentials in different segments of human motor nerve fibres. The computations use our previous double cable model of the fibres. The simulated fibres have focal demyelination of internodes, paranodes or both together. The intracellular potentials are defined mainly by the Na(+) current, as the contribution of the K(+) fast and K(+) slow currents to the total nodal ionic current is negligible. The paranodal demyelinations cause an increase in the transaxonal current and a decrease in the transmyelin current at the paranodal segments. However, there is an inverse relationship between the transaxonal and transmyelin currents at the same segments in the cases of internodal demyelination. The internodal ionic channels beneath the myelin sheath do not contribute to the intracellular potentials, but they show a high sensitivity to long-lasting pulses. The slow components of the electrotonic potentials depend on the activation of the channel types in the nodal or internodal axolemma, whereas the fast components of the potentials are determined mainly by the passive cable responses. However, the current kinetics changes (defining the investigated electrotonic changes) are relatively weak. The study summarizes the results from these modelling investigations on the mechanisms underlying the conduction slowing/block and accommodative processes in focal demyelinating neuropathies such as Guillain-Barré syndrome and multifocal motor neuropathy.     

A distributed-parameter model of the myelinated human motor nerve fibre: temporal and spatial distributions of action potentials and ionic currents

 A double cable model of the myelinated human motor nerve fibre is presented. The model is based on the nodal and internodal channels in a previous, two-component model of human motor axons (Bostock et al. 1991), added to a complex extended cable structure of nodal, paranodal and internodal segments. The model assumes a high-resistance myelin sheath and a leakage pathway to the internodal axolemma via the paranodal seal resistance and periaxonal space. The parameter values of the model were adjusted to match the recordings of threshold electrotonus in human motor fibres from Bostock et al. (1991). Kirchoff ’s current law was used to derive a system of partial differential equations for the electrical equivalent circuit, and numerical integration was performed with a fixed time increment and non-uniform spatial step sizes, in accordance with the complex structure of the fibre. The model calculations provide estimates of the spatial and temporal distributions of action potentials and their transaxonal and transmyelin components, both in different segments of the fibre and at different moments during action potential propagation. The distribution of transaxonal and transmyelin currents along the fibre and their contributions from different ionic channels are also explored. Received: 14 July 1994/Accepted in revised form: 4 April 1995     

Different effects of blocked potassium channels on action potentials, accommodation, adaptation and anode break excitation in human motor and sensory myelinated nerve fibres: computer simulations

 Action potentials and electrotonic responses to 300-ms depolarizing and hyperpolarizing currents for human motor and sensory myelinated nerve fibres have been simulated on the basis of double cable models. The effects of blocked nodal or internodal potassium (fast or slow) channels on the fibre action potentials, early and late adaptations to 30-ms suprathreshold slowly increasing depolarizing stimuli have been examined. The effects of the same channels on accommodation after the termination of a prolonged (100 ms) hyperpolarizing current pulse have also been investigated. By removing the nodal fast potassium conductance the action potentials of the sensory fibres are considerably broader than those of the motor neurons. For both types of fibres, the blocked nodal slow potassium channels have a substantially smaller effect on the action potential repolarization. When the suprathreshold depolarizing current intensity is increased, the onset of the spike burst occurs sooner, which is common in the behaviour of the fibres. The most striking differences in the burst activity during early adaptation have been found between the fibres when the nodal fast potassium channels are blocked. The results obtained confirm the fact that the motor fibres adapt more quickly to sustained depolarizing current pulses than the sensory ones. The results also show that normal human motor and sensory fibres cannot be excited by a 100-ms hyperpolarizing current pulse, even at the threshold level. When removing the potassium channels in the nodal or internodal axolemma, the posthyperpolarization increase in excitability is small, which is common in the behaviour of the fibres. However, anode break excitation can be simulated in the fibres with simultaneous removal of the potassium channels under the myelin sheath, and this is more pronounced in the human sensory fibres than in motor fibres. This phenomenon can also be found when the internodal and some of the nodal (fast or slow) potassium channels are simultaneously blocked. Received: 8 November 1999 / Accepted in revised form: 29 February 2000     

Spike discharges of skeletomotor neurons during random noise modulated transmembrane current stimulation and muscle stretch

 Spike discharges of skeletomotor neurons innervating triceps surae muscles elicited by white noise modulated transmembrane current stimulation and muscle stretch were studied in decerebrated cats. The white noise modulated current intensity ranged from 4.3 to 63.2 nA peak-to-peak, while muscle stretches ranged from 100 μm to 4.26 mm peak-to-peak. The neuronal responses were studied by averaging the muscle length records centered at the skeletomotor action potentials (peri-spike average, PSA) and by Wiener analysis. Skeletomotor spikes appeared after a sharp peak in PSA of the injected current, preceded by a longer-lasting smaller wavelet of either depolarizing or hyperpolarizing direction. The PSA amplitude was not related to the injected current amplitude nor showed any differences related to the motor unit type. The PSA amplitudes were virtually independent of the stretching amplitude σ, after an initial increase with stretching amplitudes in the range of 15–40 μm (S.D.), or 100–270 μm peak-to-peak.Analyses of cross-spectra indicated a small or absent increase in gain with frequency in response to injected current, but about 20 dB/decade in the range 10–100 Hz in response to muscle stretch. The peaks of both Wiener kernels in response to current injection appear to decrease with the amplitude of injected current, but this decrease was not statistically significant. The narrow first-order kernels suggest that the transfer function between the current input and spike discharge is lowpass with a wide passband, i.e. there is very little change in dynamics. The values of the second-order kernels appear to be nonzero only along the main diagonal. This is characteristic of a simple Hammerstein type cascade, i.e. a zero memory nonlinearity followed by a linear system. Small values of second-order kernels away from the origin and narrow first-order kernels suggest that the linear cascade contributes very little to the overall dynamic response.In contrast to Wiener kernels found in response to current injection, the Wiener kernels in response to stretch showed a decreasing trend with stretch amplitude. The size of the second-order kernels decreased to a somewhat larger extent with input amplitude than that of the first-order kernels, indicating an amplitude-dependent nonlinearity. Overall, the transformation between length and spike output was described as an LNNL cascade with second-order nonlinearities. Received: 1 April 1993/Accepted in revised form: 24 March 1994     

Local development of action potentials in slow muscle fibres after complete or partial denervation.

Pyriformis muscles of Rana temporaria were completely or partially denervated by cutting the sciatic nerve or some of the small nerve branches entering the muscle. One stimulating and one to three recording microelectrodes were inserted along the fibres in order to compare the electrical activity at these points. In an early period following denervation action potentials of variable size and shape could be observed; these action potentials were often composed of two, sometimes of three or four, components. The size of individual components depended on the position of the recording microelectrode. Individual components could occasionally be triggered separately by adjusting the strength of the stimulating current pulse; propagation of these "all or none" responses was absent. In other fibres one component of the action potential could trigger another one several millimetres apart, thus indicating propagation. Conduction velocities were approximately 0.4 m/s. In partially denervated slow fibres, endplate potentials were confined to one lateral segment of the fibres, while the action potential occupied the denervated part of the membrane. The amplitudes of endplate and action potentials varied inversely with distance. Rough estimates of the length constant of the slow fibre membrane were calculated from the spatial decay of action potentials, endplate potentials and hyperpolarizing electrotonic potentials; mean values obtained were 2.5, 4.8 and 7.7 mm respectively. The results suggest that following denervation Na channels are built into discrete areas of the slow fibre membrane and that this process depends on the amount of denervation in individual fibres.     

Electrotonic Interaction between Muscle Fibers in the Rabbit Ventricle

Transmembrane potentials were recorded simultaneously from pairs of ventricular fibers in an isolated, regularly beating preparation. A double-barrelled microelectrode was used to record the potentials from, and to polarize, one fiber. A single microelectrode was used to record from a distant fiber. The existence of two systems of fibers, termed P and V, was confirmed. Histological evidence for the existence of two types of fibers is also presented. Electrotonic current spread was observed within both systems, electrotonic interaction between the two systems was rare and always weak. In the case of those pairs of fibers showing electrotonic interaction, the distance for an e-fold decrease in magnitude of the electrotonic potentials was found to be from 300 to 600 µ in P fibers and from 100 to 300 µ in V fibers. However, no electrotonic interaction could be observed in the majority of V fiber pairs. Moreover, the magnitude of the electrotonic potential did not decay monotonically with distance in any one direction. It is concluded that the rabbit ventricle cannot be regarded as a single freely interconnected syncytium.     

Myelin as longitudinal conductor: a multi-layered model of the myelinated human motor nerve fibre

 The myelin sheath is normally regarded as an electrical insulator. Low values of radial conductance and capacitance have been measured, and in electrical models of myelinated axons the contribution of longitudinal conduction within the sheath has been ignored. According to X-ray diffraction studies, however, myelin sheaths comprise alternate lipid and aqueous layers, and the latter may be expected to have a low resistivity. We propose a new model of myelinated axons in which the aqueous layers within the myelin provide appreciable longitudinal and radial conductance, the latter via a spiral pathway. We have investigated the likely contribution of these conductive paths within the myelin to the electrical properties of a human motor nerve fibre by computer simulation, representing the myelin sheath as a series of interconnecting parallel lamellae. With this new model, action potential conduction has been simulated along a 20-node cable, and the electrotonic responses to 100-ms depolarizing and hyperpolarizing current pulses have been simulated for a uniformly polarized fibre. We have found that the hypothesis of a longitudinally conducting myelin sheath improves our previous model in two ways: it is no longer necessary to make implausible assumptions about the resistivity or width of the periaxonal space to simulate realistic electrotonus, and the conduction velocity is appreciably faster (by 8.6%). Received: 19 April 1999 / Accepted in revised form: 11 September 2000     

Electrophysiological Control of Reversed Ciliary Beating in Paramecium

Quantitative relations between ciliary reversal and membrane responses were examined in electrically stimulated paramecia. Specimens bathed in 1 mM CaCl₂, 1 mM KCl, and 1 mM Tris-HCl, pH 7.2, were filmed at 250 frames per second while depolarizing current pulses were injected. At current intensities producing only electrotonic shifts the cilia failed to respond. Stimuli which elicited a regenerative response were followed by a period of reversed ciliary beating. With increasing stimulus intensities the latency of ciliary reversal dropped from 30 to 4 ms or less, and the duration of reversal increased from 50 ms to 2.4 s or more; the corresponding regenerative responses increased in amplitude and rate of rise. With progressively larger intracellular positive pulses, electric stimulation became less effective, producing responses with a progressive increase in latency and decrease in duration of reversed beating of the cilia. When 100-ms pulses shifted the membrane potential to +70 mV or more, ciliary reversal was suppressed until the end of the pulse. "Off" responses then occurred with a latency of 2–4 ms independent of further increases in positive potential displacement. These results suggest that ciliary reversal is coupled to membrane depolarization by the influx of ions which produces the regenerative depolarization of the surface membrane. According to this view suppression of the ciliary response during stimulation occurs when the membrane potential approaches the equilibrium potential of the coupling ion, thereby retarding its influx. Previous data together with the present findings suggest that this ion is Ca²⁺.     

Bats and frogs and animals in between: evidence for a common central timing mechanism to extract periodicity pitch

Widely divergent vertebrates share a common central temporal mechanism for representing periodicities of acoustic waveform events. In the auditory nerve, periodicities corresponding to frequencies or rates from about 10 Hz to over 1,000 Hz are extracted from pure tones, from low-frequency complex sounds (e.g., 1st harmonic in bullfrog calls), from mid-frequency sounds with low-frequency modulations (e.g., amplitude modulation rates in cat vocalizations), and from time intervals between high-frequency transients (e.g., pulse-echo delay in bat sonar). Time locking of neuronal responses to periodicities from about 50 ms down to 4 ms or less (about 20–300 Hz) is preserved in the auditory midbrain, where responses are dispersed across many neurons with different onset latencies from 4–5 to 20–50 ms. Midbrain latency distributions are wide enough to encompass two or more repetitions of successive acoustic events, so that responses to multiple, successive periods are ongoing simultaneously in different midbrain neurons. These latencies have a previously unnoticed periodic temporal pattern that determines the specific times for the dispersed on-responses.     

Membrane Characteristics of the Canine Papillary Muscle Fiber

Passive and active responses to intracellular and extracellular stimulation were studied in the canine papillary muscle. The electrotonic potential produced by extracellular polarization with the partition chamber method fitted the time course and the spatial decay expected from the cable theory (the time constant, 3.3 msec; the space constant, 1.2 mm). Contrariwise, spatial decay of the electrotonic potentials produced by intracellular polarization was very short and did not fit the decay curve expected for a simple cable, although only a small difference of time course in the electrotonic potentials produced by intracellular and extracellular polarizations was observed. A similar time course might result from the fact that when current flow results from intracellular polarization, the input resistance is less dependent on the membrane resistance. The foot of the propagated action potential rose exponentially with a time constant of 1.1 msec and a conduction velocity of 0.68 m/sec. The membrane capacity was calculated from the time constant of the foot potential and the conduction velocity to be 0.76 µF/cm². The responses of the papillary muscle membrane to intracellular stimulation differed from those to extracellular stimulation applied with the partition method in the following ways: higher threshold potential, shorter latency for the active response, linearity of the current-voltage relationship, and no reduction in the membrane resistance at the crest of the action potential during current flow.     

Temporal resolution for calling song signals by female crickets, <Emphasis Type="Italic">Gryllus bimaculatus</Emphasis>

A behavioural gap detection paradigm was used to determine the temporal resolution for song patterns by female crickets, Gryllus bimaculatus. For stimuli with a modulation depth of 100% the critical gap duration was 6–8 ms. A reduction of the modulation depth of gaps to 50% led either to an increase or a decrease of the critical gap duration. In the latter case, the critical gap duration dropped to 3–4 ms indicating a higher sensitivity of auditory processing. The response curve for variation of pulse period was not limited by temporal resolution. However, the reduced response to stimuli with a high duty cycle, and thus short pause durations, was in accordance with the limits of temporal resolution. The critical duration of masking pulses inserted into pauses was 4–6 ms. An analysis of the songs of males revealed that gaps (5.8 ms) and masking pulses (6.9 ms) were at detectable time scales for the auditory pathway of female crickets. However, most of the observed temporal variation of song patterns was tolerated by females. Critical cues such as pulse period and pulse duty cycle provided little basis for inter-individual selection by females.     

A dynamical model for reflex activated head movements in the horizontal plane

 We present a controls systems model of horizontal-plane head movements during perturbations of the trunk, which for the first time interfaces a model of the human head with neural feedback controllers representing the vestibulocollic (VCR) and the cervicocollic (CCR) reflexes. This model is homeomorphic such that model structure and parameters are drawn directly from anthropomorphic, biomechanical and physiological studies. Using control theory we analyzed the system model in the time and frequency domains, simulating neck movement responses to input perturbations of the trunk. Without reflex control, the head and neck system produced a second-order underdamped response with a 5.2 dB resonant peak at 2.1 Hz. Adding the CCR component to the system dampened the response by approximately 7%. Adding the VCR component dampened head oscillations by 75%. The VCR also improved low-frequency compensation by increasing the gain and phase lag, creating a phase minimum at 0.1 Hz and a phase peak at 1.1 Hz. Combining all three components (mechanics, VCR and CCR) linearly in the head and neck system reduced the amplitude of the resonant peak to 1.1 dB and increased the resonant frequency to 2.9 Hz. The closed loop results closely fit human data, and explain quantitatively the characteristic phase peak often observed. Received: 15 April 1996 / Accepted in revised form: 1 July 1996     

Simulation of electrocortical waves

 We report simulations of the electrocorticogram of the cat and human, based on estimates of fibre range, fibre density, axonal and dendritic delays, and cortical synaptic density. The long-range cort ical connections of real cortex were simplified to couplings of symmetric density, decreasing in density with range, on a closed (toroidal) surface. Non-specific cortical activation was modelled as a diffuse global input and specific sensory input as a lo calised white noise input. Spectral properties of output included peak densities at the frequencies of the major cerebral rhythms, a ‘1/f ’ spectral envelope and ‘shift to the right’ with increasing total power as n on-specific activation increased. Steady-state travelling waves with a velocity of 5–7 m/s (human) and <1 m/s (cat) were produced. Frequency/wavenumber analysis revealed an additional class of activity with wavenumbers independent o f temporal frequency. All these findings accord qualitatively and quantitatively with existing physiological results. Global resonant modes were not prominent, but the simulations obey a restricted case of the analytical results of Nunez (1994). Wave/puls e relations resemble the findings of Freeman (1975). Received: 8 July 1994/Accepted in revised form: 15 September 1994     

Neurophysiological responses of pheromone-sensitive receptor neurons on the antenna of Trichoplusia ni (Hubner) to pulsed and continuous stimulation regimens

Both the frequency and the temporal pattern of action potentialproduction in an insect olfactory receptor neuron are stronglyaffected by odorant composition and the time course over whichstimulus concentration varies. To investigate the temporal characteristicsof the neurophysiological responses of these neurons, we deviseda stimulus delivery system that allows us to repeatedly presentwell-mixed, constant concentration odor pulses with relativelysharp onsets and offsets. Here we compare neurophysiologicalresponses to several different stimulation regimens, includingpulses of different durations and repetition rates. During stimulationwith high concentrations of pheromone, the temporal patternof neural activity from olfactory receptor neurons on the antennaof Trichoplusia ni (Hübner) is characterized by an initialphasic period (100–200 ms), followed by a tonic periodwhich is typically maintained for the remaining duration ofthe stimulus. Different olfactory receptor neurons appear tovary among themselves in the relative distribution between thephasic and tonic portions of the overall discharge. During stimulationregimens involving rapid repeated pulses of odorants, a portionof the phasic response levels is preserved during each pulse.Consequently, T. ni males probably detect much of the fluctuationin concentration of pheromone that may normally occur downwindfrom the site of pheromone release.     

Neural modeling of the dorsal cochlear nucleus: cross-correlation analysis of short-duration tone-burst responses

A conceptual model of a portion of dorsal cochlear nucleus (DCN) neural circuitry has emerged over the past two decades. This model suggests that the response properties of the DCN’s major projection neurons, called type IV units, are due, in part, to the behavior of local circuit inhibitory interneurons called type II units (Young and Brownell 1976). Cross-correlation studies of simultaneously recorded pairs of DCN units in decerebrate cat derived from 50-s best frequency (BF) stimuli are consistent with and have extended this conceptual model (Voigt and Young 1980, 1985, 1988, 1990). Interestingly, Gochin et al. (1989) found no signs of inhibition in the anesthetized rat DCN in cross-correlograms derived from 55-ms short-duration BF tone bursts. This seemingly contradictory result has motivated this study. Computer simulations were run using our network model of the intrinsic DCN neural circuitry. This model has previously been shown to reproduce the major features of both type II and type IV rate-level curves and the inhibitory trough (IT) observed in cross-correlograms derived from long-duration stimuli (Voigt and Davis 1994). The goal was to study the stimulus-duration-dependent strength of ITs in the cross-correlograms derived from short-duration BF tone-burst stimuli. The results suggest that ITs may not be detectable when the stimulus duration is 50 ms but may be detectable when the stimulus duration is 200 ms or greater. Furthermore, when the ITs are detected in cross-correlograms derived from 200-ms data sets, the strength of the IT, as measured by effectiveness, is comparable to the strength of ITs measured when the stimulus duration is 50 s. Received: 16 March 1994/Accepted in revised form: 31 May 1994     

Temporal resolution in olfaction III: flicker fusion and concentration-dependent synchronization with stimulus pulse trains of antennular chemoreceptor cells in the American lobster

To understand how chemoreceptor organs may extract temporal information from odor plumes, we investigated the frequency filter properties of lobster chemoreceptor cells. We used rapid stimulation and high-resolution stimulus measurement for accurate stimulus control and recorded extracellular responses from chemoreceptors in the lobster lateral antennule in situ. We tested 16 hydroxyproline-sensitive cells with a series of ten 100-ms pulses at 10, 100 and 1000 μmol l⁻¹ at stimulation frequencies from 0.5 Hz to 4 Hz. Receptor cell responses could accurately encode 10 μmol l⁻¹, but not 100 or 1000 μmol l⁻¹ pulses, delivered at rates of 4 Hz. Flicker-fusion frequency and synchronization with the stimulus pulse train were concentration dependent: performance rates above 1 Hz became poorer both with increasing pulse amplitude and frequency. Flicker fusion frequency was 3 Hz for 100 μmol l⁻¹ pulses and 2 Hz for 1000 μmol l⁻¹ pulses. Individual cells showed differences in their stimulus pulse following capabilities, as measured by the synchronization coefficient. These individual differences may form a basis for coding temporal features of an odor plume in an across-fiber pattern. Accepted: 7 July 1999     

Tropical zooplankton in the highly-enclosed lagoon of Taiaro Atoll (Tuamotu Archipelago, French Polynesia)

 Nocturnal zooplankton assemblages around Taiaro Atoll were sampled over six nights during February 1994. Replicate zooplankton samples were collected at windward and leeward locations in the enclosed lagoon and adjacent ocean with a metered net (85 cm diameter, 500 μm mesh) towed for 15 min at 5 m depth. The zooplankton community in the lagoon was very different from that in the ocean. Oceanic samples contained 50 mostly holoplanktonic taxa (diversity index, H′=2.62; evenness index, J′=0.67). Lagoonal samples contained 19 mostly meroplanktonic taxa (H′=1.54, J′=0.52) with three taxa (decapod larvae; an ostracod, Cypridina sp.; a copepod, Acartia fossae) contributing >90% of the individuals. Unlike the ocean, zooplankton distributions in the lagoon were not homogenous; instead spatial patterns were apparently formed by the interaction between hydrodynamic processes and species-specific behaviour. Accepted: 28 August 1997     

A distributed-parameter model of the myelinated nerve fiber

This paper presents a new model for the characterization of electrical activity in the nodal, paranodal and internodal regions of isolated amphibian and mammalian myelinated nerve fibers. It differs from previous models in the following ways: (1) in its ability to incorporate detailed anatomical and electrophysiological data; (2) in its approach to the myelinated nerve fiber as a multi-axial cable; and (3) in the numerical algorithm used to obtain distributed model equation solutions for potential and current. The morphometric properties are taken from detailed electron microscopic anatomical studies (Berthold & Rydmark, 1983a, Experientia 39, 964-976). The internodal axolemma is characterized as an excitable membrane and model-generated nodal and internodal membrane action potentials are presented. A system of describing equations for the equivalent network model is derived, based on the application of Kirchoff's Current Law, which take the form of multiple cross-coupled parabolic partial differential equations. An implicit numerical integration method is developed and the numerical solution implemented on a parallel processor. Non-uniform spatial step sizes are used, enabling detailed representation of the nodal region while minimizing the number of total segments necessary to represent the overall fiber. Conduction velocities of 20.2 m sec-1 at 20 degrees C for a 15 microns diameter amphibian fiber and 57.6 m sec-1 at 37 degrees C for a 17.5 microns diameter mammalian fiber are achieved, which agrees qualitatively with published experimental data at similar temperatures (Huxley & St?mpfli, 1949, J. Physiol., Lond. 108, 315-339; Rasminsky, 1973, Arch, Neurol. 28, 287-292). The simulation results demonstrate the ability of this model to produce detailed representations of the transaxonal, transmyelin and transfiber potentials and currents, as well as the longitudinal extra-axonal, periaxonal and intra-axonal currents. Also indicated is the potential contribution of the paranodal axolemma to nodal activity as well as the presence of significant longitudinal currents in the periaxonal space adjacent to the node of Ranvier.     

Brittle star fauna (Echinodermata: Ophiuroidea) of the arctic northwestern Barents sea: composition,abundance, biomass and spatial distribution

 Epibenthic brittle star assemblages were investigated on the northwestern Barents Sea shelf between 81° and 77°N in July 1991. At 9 drift stations in water depths between 80 and 360 m, series of 35–71 photographs, each depicting about 1 m² of the seabed, were taken along transects of about 150- to 300-m length to assess abundances and spatial distribution patterns of adult brittle stars (disc diameter ≥1 mm). Biomass values were derived by combining abundances with size-weight relationships and size frequencies established using specimens from trawl catches. Six brittle star species were identified on the seabed images. Ophiocten sericeum was the most abundant species on shallow shelf banks (≤100 m). Up to 2,800 individuals were counted on a single photograph; median abundances per station ranged from 32 to 524 ind.m^-2 and biomass from 0.3 to 5.0 g ash-free dry weight (AFDW) m^-2. The spatial distribution along the transects (i.e. on the 100-m scale) was, however, extremely patchy. Disc diameters of O. sericeum ranged between 1.6 mm and 15.4 mm. In deeper shelf habitats (>150 m), O. sericeum was rare or absent, and Ophiacantha bidentata dominated the brittle star fauna with median densities and biomasses of 2–49 ind.m^-2 and 0.07–1.9 g AFDW m^-2, respectively. Its disc diameters ranged from 2.9 to 14.4 mm. The other species (Ophiura sarsi, Ophiopholis aculeata, Ophioscolex glacialis, Ophiopleura borealis) occurred in distinctly lower numbers. Our findings provide further evidence that brittle stars dominate epibenthic communities on Arctic shelves and locally reach very high abundances. Dense beds of Ophiocten sericeum seem to be a general phenomenon on high-Arctic shallow shelf banks. Received: 30 March 1995/Accepted: 30 June 1995     

Development of a new coulter counter system: Measurement of the volume,internal conductivity,and dielectric breakdown voltage of a single guard cell protoplast ofVicia faba

Summary Two intracellular microelectrodes were used to study electrotonic interaction between cultured embryonic (16- to 20-day-old) chick myocardial cells reaggregated into small spheresin vitro. Under different culture conditions, reaggregates with two types of functional membrane properties were produced: (i) highly differentiated reaggregates, and (ii) reverted reaggregates. In the highly differentiated state, the cells had high stable resting potentials and produced rapidly-rising tetrodotoxin (TTX)-sensitive action potentials in response to electric field stimulation. In the reverted state, the cells exhibited slowrising spontaneous action potentials having prominent pacemaker potentials and TTX-insensitive upstrokes. These states resemble electrophysiological properties of the highly differentiated (18 daysin ovo) and less fully differentiated (3 daysin ovo) intact embryonic chick heart, respectively. Both types of reaggregates had similar ultrastructural appearance, with many elongated cells and intercalated disc-like structures; gap-like junctions were not seen. The highly differentiated cells had input resistances of about 5 M, and exhibited only little electrotonic interaction in response to intracellular current injection either when the cells were at rest or during the action potential plateau. Intracellular stimulation produced propagating action potentials which triggered contraction of the entire reaggregate. Large hyperpolarizing current pulses applied during the action potential plateau caused premature repolarization which also propagated to the other impaled cell. In the reverted reaggregates, electrotonic interaction was weak or absent in about 52% of the impaled cell pairs, moderate in 30%, and strong in 18% (encountered only at interelectrode distances of less than 100 m). The difference in degree of electrotonic interaction may be due to the state of differentiation with respect to the membrane electrical properties.