The sensory world of the platypus

Vision, audition and somatic sensation in the platypus are reviewed. Recent work on the eye and retinal ganglion cell layer of the platypus is presented that provides an estimate of visual acuity and suggests that platypus ancestors may have used vision, as well as the bill organ, for underwater predation. The combined electroreceptor and mechanoreceptor array in the bill is considered in detail, with special reference to the elaborate cortical structure, where inputs from these two sensory arrays are integrated in a manner that is astonishingly similar to the stripe-like ocular dominance array in primate visual of cortex, that integrates input from the two eyes. A new hypothesis, along with supporting data, is presented for this combined mechanoreceptive-electroreceptive complex in platypus cortex. Bill mechanoreceptors are shown to be capable of detecting mechanical waves travelling through the water from moving prey. These mechanical waves arrive after the electrical activity from the same prey, as a function of distance. Bimodal cortical neurones, sensitive to combined mechanical and electrical stimulation, with a delay, can thus signal directly the absolute distance of the prey. Combined with the directional information provided by signal processing of the thousands of receptors on the bill surface, the stripe-like cortical array enables the platypus to use two different sensory systems in its bill to achieve a complete, three-dimensional 'fix' on its underwater prey.     

Distribution and putative function of autonomic nerve fibres in the bill skin of the platypus (Ornithorhynchus anatinus)

The electroreceptors located in the bill skin of the platypus are modified secretory glands. The electroreceptive nerve terminals form bare endings in close proximity to the duct of these glands. In this study, we describe the autonomic innervation of the glands and a separate specialized autonomic innervation of the epidermal portion of the glandular duct. A range of immunohistochemical labels showed that the gland cells of the electroreceptors have a non-noradrenergic (putative parasympathetic) innervation. Phalloidin labelling revealed a 'sphincter' of epidermal luminal cells that labelled strongly for actin. These actin-dense keratinocytes were seen to have a noradrenergic (putative sympathetic) innervation. Fine-diameter sensory fibres containing substance P (presumably C-fibre thermoreceptors or polymodal nociceptors) were observed to terminate in the superficial epidermis surrounding the pore of the gland. When the bill of the platypus is dry these pores were closed. However, when room temperature water was washed over the bill, the pores opened. It is proposed that this autonomic and sensory innervation, along with the actin sphincter, mediates the opening and closing of the pores. By doing this, the platypus prevents the desiccation of the bare electrosensory nerve terminals when it is out of the water, and it may also be a way to regulate the impedance of the internal electrical circuit presented to the water at the pores.     

Integration of free pulses in electrical self-stimulation of the rat brain.

Frequency thresholds for electrical self-stimulation of the medial forebrain bundle were estimated in rats while low frequencies of pulses were applied continuously. When continuous pulses were delivered to the same electrode that received the 0.5-sec trains of response-initiated stimulation, thresholds decreased by the free-pulse frequency (Experiment 1), consistently across current (Experiment 2). Estimates of the reward added by concurrent, response-contingent stimulation of the opposite electrode of a bilateral pair predicted the drop in threshold caused by the noncontingent pulses applied to the opposite hemisphere (Experiment 3), again, robustly across test current (Experiment 4). Continuous pulses restricted to times between self-initiated trains lost their effect (Experiment 5). The perception of reward was invariant despite changes in the overall activity of the self-stimulation substrate. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Temporal selectivity of evoked vocal responses of Batrachyla antartandica (Amphibia: Leptodactylidae)

The advertisement call of the leptodactylid frog Batrachyla antartandica from southern Chile consists of a train of brief percussive tone pulses whose energy is centred at about 2 kHz. To gain an understanding of the temporal features that are essential for call recognition, playback experiments were conducted with 11 males. Subjects were presented with a synthetic imitation of this signal and variants for which different temporal call parameters were modified systematically. The number of pulses, pulse rate and latency of evoked vocal responses (EVRs) to stimuli having high pulse repetition rates (i.e. 8 and 16 pulses/s) were significantly weaker relative to responses to stimuli having an equal number of pulses but lower pulse rates. A similar, non-significant tendency was observed for a series of stimuli with different pulse rates for which the total stimulus duration was held constant. EVRs also decreased significantly for stimuli having long pulse durations (i.e. 48 and 96 ms) relative to stimuli comprising shorter pulses. No significant differences were observed between EVRs to stimuli for which pulse rise and fall times were varied from 1-20 ms. Responses to calls comprising trains of 10 pulses were weaker compared with stimuli having fewer pulses per train. The selective EVRs of B. antartandica for different temporal parameters contributes to an understanding of the mechanisms involved in call recognition and stress the relevance of temporal processing of sound by males for the emergence of specific patterns of vocal behaviour in anurans.Copyright 1997 The Association for the Study of Animal Behaviour1997The Association for the Study of Animal Behaviour     

Numerical analysis of a model of ligand-induced B-cell antigen-receptor clustering. Implications for simple models of B-cell activation in an immune network

Experimental measures of responsiveness to painful and non-painful stimuli as well as measures of typical and present clinical pain were assessed in 26 female patients with fibromyalgia and in an equal number of age-matched healthy women. Pressure pain thresholds, determined by means of a dolorimeter, were lower in the patients compared to the control subjects both at a tender point (trapezius) and at a non-tender control point (inner forearm). The same was true for the heat pain thresholds, measured using a contact thermode. In contrast, the pain thresholds for electrocutaneous stimuli were decreased only at the tender point. The detection thresholds for non-painful stimuli (warmth, cold and electrical stimuli) seemed to be less affected in the fibromyalgia patients, with only the detection threshold for cold being lower at both sites. Tender points were more sensitive than control points for mechanical pressure. The reverse was found for the other modalities which were tested. Although the 3 experimental pain thresholds showed patterns of either generalized or site-specific pain hyperresponsiveness, the between-methods correlations were not very high. While the correlations between the experimental pain thresholds and the various measures of clinical pain (Localized Pain Rating, McGill Pain Questionnaire) in the patients were generally low, there were significant negative correlations between pressure pain thresholds at the two sites and the level of present pain assessed by the Localized Pain Rating. We conclude that a pattern of pain hyperresponsiveness, generalized across the site of noxious stimulation and across the physical nature of the stressor, is associated with fibromyalgia.(ABSTRACT TRUNCATED AT 250 WORDS)     

Morphology of pyramidal neurones in cytochrome oxidase modules of the S-I bill representation of the platypus

The primary somatosensory cortex of the platypus (Ornithorhynchus anatinus) is characterized by a distinct array of functionally specific cytochrome oxidase (CO) modules, forming alternate CO-rich and CO-poor bands. In the current study, we undertook to establish whether the cellular morphology of layer V pyramidal neurones reflects this modular organization. To this end, we injected neurones with Lucifer Yellow in 250 microm thick, flat-mounted cortical slices and processed the tissue to reveal a light-stable reaction product. By aligning blood vessels in serial sections in which we injected individual neurones with sections processed for CO, we were able to establish the exact location of injected cells with respect to the pattern of CO bands. Pyramidal neurones in the CO-poor bands (which are responsive to both mechano- and electroreceptive stimuli) had basal dendritic fields that were larger than those in the CO-rich bands. The large basal dendritic fields of layer V pyramidal neurones in the CO-poor bands may allow for integration of a greater number of more diverse inputs, thus allowing for averaging of stimuli to improve the signal-to-noise ratio or enhance spatial discrimination of peripheral stimuli. In some instances, neurones located within approximately 100 microm of the boundaries of the CO bands had dendritic fields that appeared to conform to the CO bands, the dendrites preferentially arborizing within a single band and avoiding the neighbouring band. However, the bias was not absolute, as we observed many examples of cells with dendrites that crossed the boundary between bands. Furthermore, many cells had dendrites that showed distinct dendritic bias that bore no obvious relationship to the CO boundaries.     

Effects of pulse separation on detection thresholds for electrical stimulation of the human cochlea

Effects of pulse separation on detection of electrical stimulation of the cochlea were studied in 12 profoundly deaf human subjects with Nucleus 22 cochlear implants. Biphasic symmetric pulses were used. Pulse separation is the time from offset of one biphasic pulse to the onset of the next biphasic pulse in the train. Effects of pulse separation were studied in the context of different covariables in four stages of the experiment. Effects of pulse separation seen in the different stages were similar, despite the different covariables. Both pulse separation and the total number of pulses per stimulus seem to be important variables affecting stimulus detection. For 0.5 ms/phase pulses, thresholds were lowest at the shortest pulse separations tested (0.2-1.1 ms) and increased as a function of pulse separation. For 2 ms/phase pulses, detection thresholds were lowest at pulse separations around 7.5 ms, in most cases, and higher at both longer and shorter pulse separations. These results suggest that interactions among adjacent pulses can either hinder or facilitate detection of the signal depending on the magnitudes of pulse separation and phase duration. Pulse separations at which thresholds measured for 2 ms/phase pulses were minimum were fairly consistent across subjects and did not correlate well with speech recognition scores. However, significant variation in this measure across species has been seen.     

Effects of spatial frequency, duration, and contrast on discriminating motion directions

The minimum speed required for discriminating the direction of drifting gratings was measured at a variety of spatial frequencies, display durations, and contrasts. As was reported previously, speed thresholds were relatively constant for middle and high spatial frequencies, but speed threshold was found to be almost inversely proportional to spatial frequency in the range of 0.25 to 1.0 c/deg. Speed threshold was also found to be inversely proportional to duration between 73 and 40 ms. These results at low frequencies and short durations are shown to be consistent with limits set by the spread of energy in the stimuli, producing velocity uncertainty. A quantitative model of temporal filtering is presented that largely accounts for results at all spatial frequencies and durations by the inclusion of constant positional noise. A discussion includes the possible roles of magnocellular and parvocellular mechanisms in mediating speed thresholds.     

Brief, prolonged and repeated stimuli applied to hyperalgesic skin areas: a psychophysical study

In this experimental study brief/prolonged and single/repeated, nociceptive stimuli (laser, thermode and electrical) were used to investigate sensory changes in capsaicin-induced primary and secondary hyperalgesia. The pain threshold to prolonged thermode stimulation was reduced in the primary area and remained constant in the secondary area. The pain thresholds to brief laser and electrical stimuli remained constant in the primary but reduced in the secondary area. The summation pain threshold to repeated (five stimuli delivered at 0.5, 1, 2 and 3 Hz) laser and electrical stimuli was reduced in the secondary area. The stimulus response functions to single laser and electrical stimuli were increased in the secondary area.     

Epidural epinephrine and clonidine: segmental analgesia and effects on different pain modalities

BACKGROUND: It is not known whether epidural epinephrine has an analgesic effect per se. The segmental distribution of clonidine epidural analgesia and its effects on temporal summation and different types of noxious stimuli are unknown. The aim of this study was to clarify these issues. METHODS: Fifteen healthy volunteers received epidurally (L2-L3 or L3-L4) 20 ml of either epinephrine, 100 microg, in saline; clonidine, 8 microg/kg, in saline; or saline, 0.9%, alone, on three different days in a randomized, double-blind, cross-over fashion. Pain rating after electrical stimulation, pinprick, and cold perception were recorded on the dermatomes S1, L4, L1, T9, T6, T1, and forehead. Pressure pain tolerance threshold was recorded at S1, T6, and ear. Pain thresholds to single and repeated (temporal summation) electrical stimulation of the sural nerve were determined. RESULTS: Epinephrine significantly reduced sensitivity to pinprick at L1-L4-S1. Clonidine significantly decreased pain rating after electrical stimulation at L1-L4 and sensitivity to pinprick and cold at L1-L4-S1, increased pressure pain tolerance threshold at S1, and increased thresholds after single and repeated stimulation of the sural nerve. CONCLUSIONS: Epidural epinephrine and clonidine produce segmental hypoalgesia. Clonidine bolus should be administered at a spinal level corresponding to the painful area. Clonidine inhibits temporal summation elicited by repeated electrical stimulation and may therefore attenuate spinal cord hyperexcitability.     

Effects of electrode configuration on psychophysical strength-duration functions for single biphasic electrical stimuli in cats

The interface between electrode and neural target tissue is thought to influence certain characteristics of neural and behavioral responses to electrical stimulation of the auditory system. At present, the biophysical properties of this interface are not well understood. Here the effects of biphasic phase duration and electrode configuration on psychophysical threshold in response to electrical stimulation in cats are described. Five cats were trained to respond to acoustic stimuli using food as a reward in an operant reinforcement paradigm. After training, the animals were unilaterally deafened and implanted with a multicontact intracochlear electrode array. Thresholds for single presentations of biphasic current pulses were measured as a function of phase duration and electrode arrangement. Statistical analyses of the data indicated that strength-duration function slopes between 200 and 1600 microseconds/phase were significantly different for the different electrode configurations and, overall, were unrelated to the absolute level of the strength-duration function (i.e., were independent of absolute threshold). For all subjects, the slope of this function for intermediate pulse durations was dependent on electrode configuration and most shallow for radial-bipolar configurations (-3.4 dB/doubling), was steepest for monopolar arrangements (-5.9 dB/doubling), and was intermediate for longitudinal-bipolar pairings. (-4.4 dB/doubling). Slopes for both shorter and longer phase duration stimuli were not significantly different. The underlying mechanisms for these effects may include, or be a combination of altered electrical field patterns, integrated activity across multiple fibers, and stochastic behavior of individual auditory neurons to electrical stimulation.     

Electropermeabilization of mammalian cells to macromolecules: control by pulse duration

Membrane electropermeabilization to small molecules depends on several physical parameters (pulse intensity, number, and duration). In agreement with a previous study quantifying this phenomenon in terms of flow (Rols and Teissié, Biophys. J. 58:1089-1098, 1990), we report here that electric field intensity is the deciding parameter inducing membrane permeabilization and controls the extent of the cell surface where the transfer can take place. An increase in the number of pulses enhances the rate of permeabilization. The pulse duration parameter is shown to be crucial for the penetration of macromolecules into Chinese hamster ovary cells under conditions where cell viability is preserved. Cumulative effects are observed when repeated pulses are applied. At a constant number of pulses/pulse duration product, transfer of molecules is strongly affected by the time between pulses. The resealing process appears to be first-order with a decay time linearly related to the pulse duration. Transfer of macromolecules to the cytoplasm can take place only if they are present during the pulse. No direct transfer is observed with a postpulse addition. The mechanism of transfer of macromolecules into cells by electric field treatment is much more complex than the simple diffusion of small molecules through the electropermeabilized plasma membrane.     

Changing echocardiography request patterns between 1988 and 1993

OBJECTIVES: This article evaluates the concept of auditory threshold and discusses the limitations of assessing threshold in human neonates. The advantages and limitations of assessing neonatal threshold by means of auditory brain stem response (ABR) are discussed, and data from several studies of newborn ABR threshold are compared. The authors report data from their own study designed to compare adult and neonatal ABR threshold using tonal stimuli. EXPERIMENTAL DESIGN: Several studies are compared. Data from the authors are ABR thresholds for tone bursts of 0.5, 1.5, 4, and 8 kHz, determined from 2-channel recordings in full-term neonates and adults. Stimuli were calibrated in SPL by means of a probe microphone inserted into the ear canal along with the insert transducer of each subject. RESULTS: All studies find a degree of threshold elevation in neonates relative to adult threshold. Neonatal ABR thresholds from our laboratory for stimuli from 500 through 8000 Hz are elevated relative to adult thresholds by 5 to 25 dB. Threshold elevation in our data and in other studies has found that neonatal ABR thresholds to high-frequency stimuli show the largest elevation relative to adults and low-frequency stimuli the most mature. CONCLUSIONS: Thresholds of neonates, as measured by the ABR, are immature especially for high-frequency stimuli. Proper stimulus calibrations, which removes the influence of ear canal resonance, are important for comparisons of data across age groups. Developmental differences in the conductive mechanism and neural immaturity are the most harmonious explanations for elevation of neonatal ABR thresholds.     

COAE thresholds: 1. Effects of equal-amplitude versus subtraction methods

Although research has demonstrated that click-evoked otoacoustic emissions (COAEs) elicited by high-level stimuli are useful for identifying hearing loss, the ability of COAEs to predict behavioral thresholds has not been adequately tested. Results of studies comparing COAE thresholds and behavioral thresholds have been equivocal, perhaps due to the need for a more rigorous approach to COAE threshold estimation. The present study was designed to address several methodological concerns in COAE threshold testing, particularly the effects of two methods of stimulus presentation on COAE testing and threshold calculation. In an attempt to make COAE threshold estimation consistent across participants, COAE threshold calculations were based on mean noise floor levels across participants. COAE and noise floor levels were measured in 15 participants using both equal-amplitude clicks and a subtraction method. Broadband COAEs were analyzed into 1/3 octave bands, so that input/output functions could be examined and COAE thresholds could be calculated for each 1/3 octave band. Comparison of the two stimulus methods indicated several differences. Mean noise floor levels for the equal-amplitude method were approximately 6 dB lower than those measured for the subtraction method across frequency. In many cases COAEs evoked using the equal-amplitude method were higher in amplitude than those evoked using the subtraction method. COAE thresholds measured using the equal-amplitude click stimuli were significantly lower than those measured using the subtraction method. The significantly higher thresholds obtained using the subtraction method may be attributed in part to the reduction of COAE amplitude by the subtraction procedure, and not merely to the higher noise level. Slopes of the input/output functions were not significantly different between the two stimulus methods. These results suggest that the equal-amplitude method is preferable for COAE threshold testing because lower noise floor and larger amplitude COAEs may be obtained in the same test time.     

Electrical stimulation of the auditory nerve. I. Correlation of physiological responses with cochlear status

The purpose of the present study was to evaluate evoked potential and single fibre responses to biphasic current pulses in animals with varying degrees of cochlear pathology, and to correlate any differences in the physiological response with status of the auditory nerve. Six cats, whose cochleae ranged from normal to a severe neural loss (< 5% spiral ganglion survival), were used. Morphology of the electrically evoked auditory brainstem response (EABR) was similar across all animals, although electrophonic responses were only observed from the normal animal. In animals with extensive neural pathology, EABR thresholds were elevated and response amplitudes throughout the dynamic range were moderately reduced. Analysis of single VIIIth nerve fibre responses were based on 207 neurons. Spontaneous discharge rates among fibres depended on hearing status, with the majority of fibres recorded from deafened animals exhibiting little or no spontaneous activity. Electrical stimulation produced a monotonic increase in discharge rate, and a systematic reduction in response latency and temporal jitter as a function of stimulus intensity for all fibres examined. Short-duration current pulses elicited a highly synchronous response (latency < 0.7 ms), with a less well synchronized response sometimes present (0.7-1.1 ms). There were, however, a number of significant differences between responses from normal and deafened cochleae. Electrophonic activity was only present in recordings from the normal animal, while mean threshold, dynamic range and latency of the direct electrical response varied with cochlear pathology. Differences in the ability of fibres to follow high stimulation rates were also observed; while neurons from the normal cochlea were capable of 100% entrainment at high rates (600-800 pulses per second (pps)), fibres recorded from deafened animals were often not capable of such entrainment at rates above 400 pps. Finally, a number of fibres in deafened animals showed evidence of 'bursting', in which responses rapidly alternated between high entrainment and periods of complete inactivity. This bursting pattern was presumably associated with degenerating auditory nerve fibres, since it was not recorded from the normal animal. The present study has shown that the pathological response of the cochlea following a sensorineural hearing loss can lead to a number of significant changes in the patterns of neural activity evoked via electrical stimulation. Knowledge of the extent of these changes have important implications for the clinical application of cochlear implants.     

Behavioral responsiveness of adrenalectomized, hypophysectomized, and intact rats to electric shock.

Tested whether the altered rates of acquisition and extinction of avoidance behavior in adrenalectomized and hypophysectomized rats are associated with abnormal responsiveness to electric shock. The electrical threshold for flinch, jump, and vocalization behaviors in adrenalectomized and hypophysectomized Ss (N = 95) was measured in 2 experiments. Adrenalectomized Ss had higher thresholds for flinch and jump responses than hypophysectomized Ss, and also a higher flinch threshold than weight-matched controls. Hypophysectomized Ss had normal thresholds for all 3 behaviors. The difference in threshold for the flinch response between adrenalectomized and hypophysectomized or normal Ss was not explained by differences in body weight, although heavy Ss responded less to electric shock than light Ss. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Comparison of acoustic reflex and behavioral thresholds as a function of stimulus frequency and duration

Threshold-duration functions for the acoustic reflex and threshold-duration functions based upon behavioral measures were obtained on the same group of subjects and compared. The mean temporal integration for the acoustic reflex threshold appears to be comparable to that for the behavioral threshold for stimuli of 500, 1000, and 2000 Hz, but marked individual differences exist in temporal integration of the acoustic reflex. At 4000 Hz, the mean change in the reflex threshold as a function of stimulus duration is significantly greater than the change in the behavioral threshold under the same conditions, suggesting that in the elicitation of the acoustic reflex the auditory system processes energy less efficiently at 4000 Hz than it does at lower frequencies.     

Dynamics of conditioned reflex reorganization in the human visual system

The dymamics of absolute and relative changes in the brightness thresholds of letters recognition was studied in 21 subjects, under conditions of dark adaptation and molecular observation, before and after pairing of one of the letters with electrocutaneous stimulation of the index finger during one session. Isolated electrocutaneous stimulation was applied on the 10th or 35th day after the session. It has been found that after pairings the relative threshold of recognition of the letter previously paired with electrical stimuli (i.e. the threshold in relation to recognition thresholds for the other letters) becomes for three to three and a half hours significantly lower than the initial one. Then there sharply sets in a phase of threshold elevation. Both in the case of double and multiple tests, this phase persists for not less than 35 days. Following an isolated electrical stimulation, a momentary significant lowering of the threshold sets in only for the letter which was paired with it in the first sessions.     

Gender differences in rowing performance and power with aging

Gender differences in experimental pain sensitivity have been widely investigated, and the results generally indicate that females exhibit greater sensitivity to noxious stimuli than males. However, results using thermal pain procedures have been inconsistent, with some studies reporting greater responses among females and other studies reporting no gender differences. The present study investigated gender differences in thermal pain perception using several different psychophysical procedures. Twenty-seven females and 22 males underwent thermal testing, including: determination of thermal pain threshold and tolerance, a thermal discrimination procedure, real-time magnitude estimates of heat pulses, and temporal summation of thermal pain. The results indicated lower thermal pain threshold and tolerance and greater temporal summation of thermal pain among females, but no gender differences in thermal discrimination or real-time magnitude estimates of discrete heat pulses. These findings suggest that gender differences in thermal pain perception may be more robust for sustained, temporally dynamic thermal stimuli with a strong C-fiber component.