Spectral determination of responses to species-specific calls in the dorsal nucleus of the lateral lemniscus

This study evaluated how neurons in the dorsal nucleus of the lateral lemniscus (DNLL) in Mexican free-tailed bats respond to both tone bursts and species-specific calls. Up to 20 calls were presented to each neuron, of which 18 were social communication and 2 were echolocation calls. We also measured excitatory response regions (ERRs): the range of tone burst frequencies that evoked discharges at a fixed intensity. Neurons were unselective for one or another call in that each neuron responded to any call so long as the call had energy that encroached on its ERR. Additionally, responses were evoked by the same set of calls, and with similar spike counts, when they were presented normally or reversed. By convolving activity in the ERRs with the spectrogram of each call, we showed that responses to tones accurately predicted discharge patterns evoked by species-specific calls. DNLL cells are remarkably homogeneous in that neurons having similar BFs responded to each of the species-specific calls with similar response profiles. The homogeneity was further illustrated by the ability to accurately predict the response profiles of a particular DNLL cell to species-specific calls from the ERR of another similarly tuned DNLL cell. Thus DNLL neurons tuned to the same or similar frequencies responded to species-specific calls with latencies and temporal discharge patterns that were so similar as to be virtually interchangeable. What this suggests is that DNLL responses evoked by complex sounds can be largely explained by a simple summation of the excitation in each neuron's ERR. Finally, superimposing the spectrograms of each call on the responses evoked by that call revealed that the DNLL population response re-creates both the spectral and the temporal features of each signal.     

Differing roles of inhibition in hierarchical processing of species-specific calls in auditory brainstem nuclei

Here we report on response properties and the roles of inhibition in three brain stem nuclei of Mexican-free tailed bats: the inferior colliculus (IC), the dorsal nucleus of the lateral lemniscus (DNLL) and the intermediate nucleus of the lateral lemniscus (INLL). In each nucleus, we documented the response properties evoked by both tonal and species-specific signals and evaluated the same features when inhibition was blocked. There are three main findings. First, DNLL cells have little or no surround inhibition and are unselective for communication calls, in that they responded to approximately 97% of the calls that were presented. Second, most INLL neurons are characterized by wide tuning curves and are unselective for species-specific calls. The third finding is that the IC population is strikingly different from the neuronal populations in the INLL and DNLL. Where DNLL and INLL neurons are unselective and respond to most or all of the calls in the suite we presented, most IC cells are selective for calls and, on average, responded to approximately 50% of the calls we presented. Additionally, the selectivity for calls in the majority of IC cells, as well as their tuning and other response properties, are strongly shaped by inhibitory innervation. Thus we show that inhibition plays only limited roles in the DNLL and INLL but dominates in the IC, where the various patterns of inhibition sculpt a wide variety of emergent response properties from the backdrop of more expansive and far less specific excitatory innervation.     

Serotonin 1B receptor modulates frequency response curves and spectral integration in the inferior colliculus by reducing GABAergic inhibition

The selectivity of sensory neurons for stimuli is often shaped by a balance between excitatory and inhibitory inputs, making this balance an effective target for regulation. In the inferior colliculus (IC), an auditory midbrain nucleus, the amplitude and selectivity of frequency response curves are altered by the neuromodulator serotonin, but the changes in excitatory-inhibitory balance that mediate this plasticity are not well understood. Previous findings suggest that the presynaptic 5-HT1B receptor may act to decrease the release of GABA onto IC neurons. Here, in vivo extracellular recording and iontophoresis of the selective 5-HT1B agonist CP93129 were used to characterize inhibition within and surrounding frequency response curves using two-tone protocols to indirectly measure inhibition as a decrease in spikes relative to an excitatory tone alone. The 5-HT1B agonist attenuated such two-tone spike reduction in a varied pattern among neurons, suggesting that the function of 5-HT1B modulation also varies. The hypothesis that the 5-HT1B receptor reduces inhibition was tested by comparing the effects of CP93129 and the GABA(A) antagonists bicuculline and gabazine in the same neurons. The effects of GABA(A) antagonists on spike count, tuning bandwidth, two-tone ratio, and temporal response characteristics mimicked those of CP93129 across the neuron population. GABA(A) antagonists also blocked or reduced the facilitation of evoked responses by CP93129. These results are all consistent with the reduction of GABA(A)-mediated inhibition by 5-HT1B receptors in the IC, resulting in an increase in the level of evoked responses in some neurons, and a decrease in spectral selectivity in others.     

Whole cell recordings of intrinsic properties and sound-evoked responses from the inferior colliculus

Response features of inferior colliculus (IC) neurons to both current injections and tone bursts were studied with in vivo whole cell recordings in awake Mexican free-tailed bats. Of 160 cells recorded, 95% displayed one of three general types of discharge patterns in response to the injection of positive current: 1) sustained discharges; 2) adapting discharges; and 3) onset-bursting discharges. Sustained neurons were the most common type (N=78), followed by onset-bursting (N=57). The least common type was adapting (N=17). In 90 neurons the profiles of synaptic and discharge activity evoked by tones of different frequencies at 50 dB SPL were recorded. Three major tone-evoked response profiles were obtained; 1) neurons dominated by excitation (N=32) in which tones evoked excitatory post-synaptic potentials (EPSPs) or EPSPs with discharges over a range of frequencies with little or no evidence of inhibitory post-synaptic potentials (IPSPs) evoked by frequencies that flanked the excitation; 2) neurons that had an excitatory frequency region in which discharges were evoked that was flanked by frequencies that evoked predominantly IPSPs (N=26); 3) neurons in which all frequencies evoked IPSPs with little or no depolarizations (N=32). The question we asked is whether IC cells that express a particular profile of PSPs and discharges to acoustic stimulation also have the same current-evoked response profile. We show that, with one exception, the intrinsic features of an IC neuron are not correlated with the pattern of its synaptic innervation; the two features are unrelated in the majority of IC cells. The exception is a subtype of inhibitory dominated cell where most frequencies evoked IPSPs to both the onset and to the offset of the tone bursts. In those cells injected current steps always evoked an onset-bursting response.     

Basolateral amygdala responds robustly to social calls: spiking characteristics of single unit activity

Vocalizations emitted within a social context can trigger call-specific changes in the emotional and physiological/autonomic state of the receiver. The amygdala is implicated in mediating these changes, but its role in call perception remains relatively unexplored. We examined call and pitch selectivity of single neurons within the basolateral amygdala (BLA) by recording spiking activity in response to 5 pitch variants of each of 14 species-specific calls presented to awake, head-restrained mustached bats, Pteronotus parnellii. A response-wise analysis across neurons revealed seven types of temporal response patterns based on the timing and duration of spiking. Roughly half of the responses to different call types were significantly affected by changes in call pitch. A neuron-wise analysis revealed that ～ 12% (8/69) of the neurons preferred the same pitch across all call types. Ninety-three percent (93/100) of neurons were excited by at least one call type and 76% exhibited either complete or transient suppression to one or more call types. The majority of neurons preferred fewer than half of the 14 different simple-syllabic calls. A call-wise analysis of spiking activity revealed that call types signaling either threat or fear most consistently evoked increases in the spike rate. In contrast, calls emitted during appeasement tended to evoke spike suppression. Our data suggest that BLA neurons participate in the processing of multiple call types and exhibit a rich variety of temporal response patterns that are neither neuron nor call specific.     

Representation of species-specific vocalizations in the inferior colliculus of the guinea pig

The responses of individual neurons to 4 typical guinea pig vocalization calls (purr, chutter, chirp, and whistle) were recorded in the inferior colliculus (IC) of anesthetized guinea pigs. All calls elicited a response in about 80% of units. Unit selectivity for individual calls was low, given that a majority of neurons (55% of 124 units) responded to all vocalizations and only a small portion of neurons (3%) responded to only one call or did not respond to any of the calls (3%). In 15% of units, the response to one call was > or =25% stronger than the response to any other sound (tone, noise, and other calls); these neurons were selective for chirp or whistle, and no unit preferred chutter or purr. Neuronal activity provided information about the spectrotemporal patterns of the calls. Peristimulus time histograms (PSTHs) reflected the energy of the near-characteristic frequency band, and the population PSTH reliably matched the sound envelope for calls characterized by one or more short impulses (chirp, purr, and chutter) but did not exactly fit the envelope for whistle--a slow-modulated and relatively long call. Calculations based on firing rates indicated the approximate positions of the main spectral peaks but did not always reflect their relative magnitude. The time-reversed version of whistle elicited on average a weaker response than did the natural whistle (by 24%), but there were neurons with a significantly stronger response to the natural ("forward-selective," 30%) as well as to the time-reversed whistle ("reverse-selective," 15%). This study does not prove the existence of units selectively responding to animal calls, but it provides evidence for the encoding of the spectrotemporal acoustic patterns of vocalizations by IC units.     

Receptive field structure varies with layer in the primary visual cortex

Here we ask whether visual response pattern varies with position in the cortical microcircuit by comparing the structure of receptive fields recorded from the different layers of the cat's primary visual cortex. We used whole-cell recording in vivo to show the spatial distribution of visually evoked excitatory and inhibitory inputs and to stain individual neurons. We quantified the distribution of 'On' and 'Off' responses and the presence of spatially opponent excitation and inhibition within the receptive field. The thalamorecipient layers (4 and upper 6) were dominated by simple cells, as defined by two criteria: they had separated On and Off subregions, and they had push-pull responses (in a given subregion, stimuli of the opposite contrast evoked responses of the opposite sign). Other types of response profile correlated with laminar location as well. Thus, connections unique to each visual cortical layer are likely to serve distinct functions.     

Spatiotemporal effects of microstimulation in rat neocortex: a parametric study using multielectrode recordings

Using microstimulation to imprint meaningful activity patterns into intrinsically highly interconnected neuronal substrates is hampered by activation of fibers of passage leading to a spatiotemporal "blur" of activity. The focus of the present study was to characterize the shape of this blur in the neocortex to arrive at an estimate of the resolution with which signals can be transmitted by multielectrode stimulation. The horizontal spread of significant unit activity evoked by near-threshold focal electrical stimulation (charge transfer 0.8–4.8 nC) and multielectrode recording in the face representation of the primary somatosensory cortex of ketamine anesthetized rats was determined to be about 1,350 µm. The evoked activity inside this range consisted in a sequence of fast excitatory response followed by an inhibition lasting >100 ms. These 2 responses could not be separated by varying the intensity of stimulation while a slow excitatory rebound after the inhibitory response was restricted to higher stimulus intensities (>2.4 nC). Stimulation frequencies of 20 and 40 Hz evoked repetitive excitatory response standing out against a continuous background of inhibition. At 5- and 10-Hz stimulation, the inhibitory response showed a complex interaction pattern attributed to highly sublinear superposition of individual inhibitory responses. The present data help to elucidate the neuronal underpinnings of behavioral effects of microstimulation. Furthermore, they provide essential information to determine spatiotemporal constraints for purposeful multielectrode stimulation in the neocortex.     

Regulation of cerebello-cortical transmission in the rat ventromedial thalamic nucleus

Summary On the basis of antidromic stimulation we have identified two distinct neuronal populations in the rat ventromedial thalamic nucleus. The largest population (96%) are thalamo-cortical relay cells which project via the internal capsule to the cerebral cortex. The smaller population of cells (4%) project caudally to the reticular formation and superior colliculus. These two cell types could be distinguished further on the basis of their patterns of spontaneous discharge. Relay cells fluctuate between two activity patterns (i) a rhythmic pattern characterized by periods of high-frequency bursting, and (ii) a more tonic discharge pattern of single spikes. The caudally projecting cells had a characteristic fast, regular type of spontaneous firing. Brachium conjunctivum stimulation evokes two distinct responses in thalamic relay cells, (i) a short-latency single spike, (ii) a longer latency, rhythmic response of 2–3 spikes. Both excitatory responses are followed by a period of cell quiescence. The type of response is dependent upon the cell's firing pattern. The short-latency response occurs during tonic, single-spike activity whilst the longer latency response occurs during highfrequency bursting activity. The short-latency response can be altered to the long latency response by increasing the level of anaesthesia or by applying a conditioning shock to known inhibitory pathways. Conversely the long latency response can be altered to the short-latency response by decreasing anaesthesia or by stimulation of the reticular formation. It is argued that both response types are evoked monosynaptically by activation of the same cerebello-thalamic fibres but that different ionic conductances which are active at different levels of membrane polarization are responsible for the two response patterns. Efficient time-locked cerebellothalamo-cortical transmission occurs only during tonic single-spike activity, when cerebellar stimulation evokes a short-latency response. Such transmission is allowed or disallowed by the fine balance between converging excitatory and inhibitory afferents. In addition to a monosynaptic excitatory input from the cerebellar nuclei, relay cells received converging synaptic inputs from the substantia nigra, cerebral cortex, reticular formation and superior colliculus. Due to the anatomical arrangement in the rat it proved impossible to assess the role of the pallidum. The population of caudally projecting cells also received several converging synaptic inputs, but unlike those influencing relay cells, these inputs were all excitatory. We have obtained no clear physiological evidence for the occurrence of local interneurones within the ventromedial nucleus. However, a powerful recurrent inhibitory circuit is activated following antidromic activation of relay cells. The interneurones responsible for this inhibition appear to lie in the thalamic nucleus reticularis.Abbreviations ACh Acetylcholine - ACG Autocorrelollogram - BC Brachium Conjunctivum - EEG Electroencephalogram - GABA Gamma aminobutyric acid - GP Globus Pallidus - IPSP Inhibitory post-synaptic potential - IC Internal capsule - ISIH Interspike interval histogram - MRF Mesencephalic reticular formation - PSTH Post stimulus time histogram - St Striatum - SN Substantia Nigra - SC Superior Colliculus - VM Ventromedial thalamic nucleus     

Spectral shape sensitivity contributes to the azimuth tuning of neurons in the cat's inferior colliculus

We recorded high-best-frequency single-unit responses to free-field noise bursts that varied in intensity and azimuth to determine whether inferior colliculus (IC) neurons derive directionality from monaural spectral-shape. Sixty-nine percent of the sample was directional (much more responsive at some azimuths than others). One hundred twenty-nine directional units were recorded under monaural conditions (unilateral ear plugging). Binaural directional (BD) cells showed weak monaural directionality. Monaural directional (MD) cells showed strong monaural directionality, i.e., were much more responsive at some directions than others. Some MD cells were sensitive to both monaural and binaural directional cues. MD cells were monaurally nondirectional in response to tone bursts that lack direction-dependent variation in spectral shape. MD cells were unresponsive to noise bursts at certain azimuths even at high intensities showing that particular spectral shapes inhibit their responses. Two-tone inhibition was stronger where MD cells were unresponsive to noise stimulation than at directions where they were responsive. According to the side-band inhibition model, MD cells derive monaural directionality by comparing energy in excitatory and inhibitory frequency domains and thus should have stronger inhibitory side-bands than BD cells. MD and BD cells showed differences in breadth of excitatory frequency domains, strength of nonmonotonic level tuning, and responsiveness to tones and noise that were consistent with this prediction. Comparison of these data with previous findings shows that strength of spectral inhibition increases greatly between the level of the cochlear nucleus and the IC, and there is relatively little change in strength of spectral inhibition among the IC, auditory thalamus, and cortex.     

The effectiveness of bicuculline as an antagonist of GABA and visually evoked inhibition in the cat''s striate cortex.

1. The iontophoretic application of the alkaloid bicuculline to neurones in area 17 of the cat's visual cortex effectively antagonized the inhibitory action of iontophoretically applied GABA in fifty-four out of sixty-two neurones examined. It had little or no effect on the inhibitory action of iontophoretically applied glycine. 2. At the stage that the iontophoretic application of bicuculline blocked the inhibitory action of GABA it also reduced or blocked visually evoked inhibitory influences acting on forty-three of the fifty-four cells. This effect on visually evoked inhibition was not reproduced by simply raising the neural spontaneous activity with iontophoretically applied glutamate. 3. For those seven neurones where the iontophoresis of bicuculline failed to block the inhibitory action of iontophoretically applied GABA it also failed to produce any change in visually evoked inhibition. 4. In all cases where a visually evoked inhibition of a cells resting discharge was reduced by the iontophoretic application of bicuculline, the inhibitory response was replaced by an excitatory response. The application of bicuculline also revealed excitatory responses to certain of the visual stimuli that previously appeared to exert neither inhibitory nor excitatory effects on a cell, and often where cells normally exhibited small excitatory responses it produced large increases in the magnitude of the evoked response. 5. These results indicate that the normal responses of the neurones examined in the present work, to the particular visual stimuli used, reflect an interaction between simultaneously evoked excitatory and inhibitory inputs. It is suggested that the iontophoretic application of bicuculline by blocking or reducing the inhibitory input moves the balance between the inputs in favour of the excitatory input. 6. The present results support the view that GABA is an inhibitory transmitter in the visual cortex.     

Facial electromyographic reactions and autonomic activity to auditory stimuli

This study explored whether high- and low-intensity auditory stimuli evoke different facial electromyographic reactions and autonomic responses. Subjects were repeatedly exposed to 95-dB and 75-dB tones (1000 Hz, 40 ms rise and fall times) while their facial electromyograms from the corrugator and zygomatic muscle regions, heart rate, skin conductance responses, skin conductance half recovery time and ratings were measured. The 95-dB tone evoked a "negative" reaction with increased corrugator activity and an autonomic response pattern that carried aspects of a defense reaction, that is, slowly habituating skin conductance responses with retarded recovery rate and an initial tendency to heart rate acceleration. Furthermore, the 95-dB tone was rated as unpleasant. The 75-dB tone elicited an orienting response indicated by a distinct heart rate deceleration and fast habituating skin conductance responses with a relatively short recovery time. Thus, the present study demonstrated that the facial electromyographic technique is sensitive to simple environmental stimuli such as auditory stimuli and that the facial response is consistent with the autonomic response patterns and the experience of the stimuli.     

Spatiotemporal dynamics of excitation in rat insular cortex: intrinsic corticocortical circuit regulates caudal-rostro excitatory propagation from the insular to frontal cortex

The insular cortex (IC), composing unique anatomical connections, receives multi-modal sensory inputs including visceral, gustatory and somatosensory information from sensory thalamic nuclei. Axonal projections from the limbic structures, which have a profound influence on induction of epileptic activity, also converge onto the IC. However, functional connectivity underlying the physiological and pathological roles characteristic to the IC still remains unclear. The present study sought to elucidate the spatiotemporal dynamics of excitatory propagation and their cellular mechanisms in the IC using optical recording in urethane-anesthetized rats. Repetitive electrical stimulations of the IC at 50 Hz demonstrated characteristic patterns of excitatory propagation depending on the stimulation sites. Stimulation of the granular zone of the IC (GI) and other surrounding cortices such as the motor/primary sensory/secondary sensory cortices evoked round-shaped excitatory propagations, which often extended over the borders of adjacent areas, whereas excitation of the agranular and dysgranular zones in the IC (AI and DI, respectively) spread along the rostrocaudal axis parallel to the rhinal fissure. Stimulation of AI/DI often evoked excitation in the dorsolateral orbital cortex, which exhibited spatially discontinuous topography of excitatory propagation in the IC. Pharmacological manipulations using 6,7-dinitroquinoxaline-2,3(1H,4H)-dione (DNQX), a non-NMDA receptor antagonist, D-2-amino-5-phosphonovaleric acid (D-APV), an NMDA receptor antagonist, and bicuculline methiodide, a GABA_A receptor antagonist, indicate that excitatory propagation was primarily regulated by non-NMDA and GABA_A receptors. Microinjection of lidocaine or incision of the supragranular layers of the rostrocaudally middle part of excitatory regions suppressed excitation in the remote regions from the stimulation site, suggesting that the excitatory propagation in the IC is largely mediated by cortical local circuits. These features of excitatory propagation in the AI/DI, that is the propagation along the rostrocaudal axis with less propagation in the ventro-dorsal direction, may play an important role for transmitting neural excitation arising from the limbic structures to the frontal and orbital cortices.     

Physiological response properties of neurons in the superior paraolivary nucleus of the rat

The superior paraolivary nucleus (SPON) is a prominent nucleus of the superior olivary complex. In rats, this nucleus is composed of a morphologically homogeneous population of GABAergic neurons that receive excitatory input from the contralateral cochlear nucleus and inhibitory input from the ipsilateral medial nucleus of the trapezoid body. SPON neurons provide a dense projection to the ipsilateral inferior colliculus and are thereby capable of exerting profound modulatory influence on collicular neurons. Despite recent interest in the structural and connectional features of SPON, little is presently known concerning the physiological response properties of this cell group or its functional role in auditory processing. We utilized extracellular, in vivo recording methods to study responses of SPON neurons to broad band noise, pure tone, and amplitude-modulated pure tone stimuli. Localization of recording sites within the SPON provides evidence for a medial (high frequency) to lateral (low frequency) tonotopic representation of frequencies within the nucleus. Best frequencies of SPON neurons spanned the audible range of the rat and receptive fields were narrow with V-shaped regions near threshold. Nearly all SPON neurons responded at the offset of broad band noise and pure tone stimuli. The vast majority of SPON neurons displayed very low rates of spontaneous activity and only responded to stimuli presented to the contralateral ear, although a small population showed binaural facilitation. Most SPON neurons also generated spike activity that was synchronized to sinusoidally amplitude-modulated tones. Taken together, these data suggest that SPON neurons may serve to encode temporal features of complex sounds, such as those contained in species-specific vocalizations.     

Responses of primate spinomesencephalic tract cells to intradermal capsaicin

The responses of 32 spinomesencephalic tract cells to intradermal capsaicin were examined in anesthetized monkeys. Wide dynamic range (n=20) and nociceptive specific (n=6) cells showed two types of excitatory responses to intradermal injection of capsaicin. The first excitatory response shown by the majority of wide dynamic range (n=13) and nociceptive specific (n=4) cells was consistent with their sensitization by capsaicin. The cells showed an acute and prolonged increase in ongoing activity with capsaicin injection. Responses to mechanical stimuli were substantially increased after capsaicin and an expansion of receptive field was often observed. The responses of the same cells to excitatory amino acid agonists applied locally by iontophoresis also increased. All cells showing sensitization were antidromically activated from periaqueductal gray regions dorsal to the sulcus limitans. Electrical stimulation at these sites did not affect the ongoing or evoked discharges of these cells.

The second excitatory response of wide dynamic range (n=5) and nociceptive specific (n=1) cells was a novel pattern not consistent with sensitization. These cells nevertheless showed an acute and prolonged increase in background activity after capsaicin injection. Yet, there was no change or a decrease in responses to cutaneous stimuli, no evidence for change in receptive field size and no increase in responses to locally released excitatory amino acids. These cells projected to regions in the periaqueductal gray ventral to the sulcus limitans. Electrical stimulation at these sites produced a decrease in spontaneous activity of the same cell. Low threshold mesencephalic-projecting neurons (n=6) showed a single inhibitory pattern (n=4) of responses to capsaicin. The injection produced an acute decrease in spontaneous activity that was sustained for at least 30 min after injection. The responses to cutaneous stimuli and to excitatory amino acids were also substantially reduced. Low threshold cells were found that projected to both dorsal-lateral and ventral-lateral regions of the periaqueductal gray. In summary, three patterns of responses shown by primate spinomesencephalic tract cells to intradermal capsaicin appear dependent on the functional regions of the periaqueductal gray to which they project. These results suggest that inputs of spinomesencephalic tract neurons to the periaqueductal gray may evoke important components of the systemic response to the neurogenic hyperalgesia produced by intradermal capsaicin.     

Membrane and action potential responses evoked by excitatory amino acids acting at N-methyl-D-aspartate receptors and non-N-methyl-D-aspartate receptors in the rat thalamus in vivo.

The membrane potential responses and firing patterns of rat thalamic neurons evoked by iontophoretically applied excitatory amino acids were recorded in vivo. All excitatory amino acids, including N-methyl-D,L-aspartate, evoked a membrane depolarization and a repetitive, regular pattern of action potential firing in the thalamus. Both non-nociceptive and nociceptive thalamic neurons responded to all agonists tested. Iontophoretic application of magnesium ions selectively antagonized responses to N-methyl-D,L-aspartate but did not convert the repetitive firing pattern into a burst firing pattern. In contrast, in the hippocampus, N-methyl-D,L-aspartate evoked a burst pattern of action potential firing associated with rhythmic depolarizing membrane potential shifts, similar to those seen by other workers in the hippocampus and in other brain regions. These findings are discussed in relation to the possibility that the regular firing pattern of spikes evoked by excitatory amino acids in the thalamus is primarily determined by the intrinsic membrane properties of thalamic neurons.     

Membrane properties and the neuro-effector transmission of smooth muscle cells in the canine internal anal sphincter.

The most distal part of the circular muscle layer functions as the internal anal sphincter, which constitutes a high pressure zone at rest, but maintains a relaxed state during defecation. To elucidate such sphincter mechanisms of the smooth muscle cells, the circular muscle layer in the canine anal canal was examined within 2 cm from the anal verge. Both the mechanical and intracellular electrical activities were recorded simultaneously. The examined region could be divided into three different regions according to the pattern of spontaneous activity and innervation and consisted of an upper region (20-15 mm from the anal verge), a transitional region (15-5 mm from the anal verge) and a lower region (within 5 mm from the anal verge), respectively. The spontaneous membrane activity was characterized by ongoing slow potential changes and each potential change was associated with a phasic contraction in the three regions. The mean frequencies of spontaneous electrical activity were 6.8, 15.9, and 24.1 c/min in the upper, transitional and lower regions, respectively. In the transitional and lower region, muscle tone generation was observed. Transmural field stimulation (0.4 msec in pulse duration) evoked membrane depolarization and contractions in the lower region. The application of an alfa adrenergic blocking agent completely suppressed the generation of excitatory responses, leaving a long lasting hyperpolarization associated with relaxation. In the transitional and upper region, stimulation consistently evoked membrane hyperpolarization with relaxation. The characteristics of this hyperpolarization response varied among the three regions. The total duration of hyperpolarization increased distally, while the time to peak hyperpolarization became decreases in a reverse direction. These regional differences in the characteristics of spontaneous membrane activity and innervation indicate that the transitional and lower region might therefore function as the internal anal sphincter.     

Timing of sound-evoked potentials and spike responses in the inferior colliculus of awake bats

Neurons in the inferior colliculus (IC), one of the major integrative centers of the auditory system, process acoustic information converging from almost all nuclei of the auditory brain stem. During this integration, excitatory and inhibitory inputs arrive to auditory neurons at different time delays. Result of this integration determines timing of IC neuron firing. In the mammalian IC, the range of the first spike latencies is very large (5-50 ms). At present, a contribution of excitatory and inhibitory inputs in controlling neurons' firing in the IC is still under debate. In the present study we assess the role of excitation and inhibition in determining first spike response latency in the IC. Postsynaptic responses were recorded to pure tones presented at neuron's characteristic frequency or to downward frequency modulated sweeps in awake bats. There are three main results emerging from the present study: (1) the most common response pattern in the IC is hyperpolarization followed by depolarization followed by hyperpolarization, (2) latencies of depolarizing or hyperpolarizing responses to tonal stimuli are short (3-7 ms) whereas the first spike latencies may vary to a great extent (4-26 ms) from one neuron to another, and (3) high threshold hyperpolarization preceded long latency spikes in IC neurons exhibiting paradoxical latency shift. Our data also show that the onset hyperpolarizing potentials in the IC have very small jitter (<100 mus) across repeated stimulus presentations. The results of this study suggest that inhibition, arriving earlier than excitation, may play a role as a mechanism for delaying the first spike latency in IC neurons.     

Functional organization within the medullary reticular formation of the intact unanesthetized cat. III. Microstimulation during locomotion.

1. This article presents the results from stimulation in 21 loci within the medullary reticular formation (MRF; between 0.5 and 2.5 mm from the midline) and in 5 loci in the medial longitudinal fasciculus (MLF) of four intact, unanesthetized cats during locomotion. Stimulus trains (11 pulses, 0.2-ms duration, 330 Hz, stimulus strength 35 microA) were applied at those loci in each track at which the most widespread effects in each of the four limbs were obtained with the cat at rest. Electromyograms were recorded from flexor and extensor muscles of each limb. 2. As previously reported, stimulation with the cat at rest generally evoked brief, short-latency, twitch responses in both flexor and extensor muscles of more than one limb. In contrast, stimulation during locomotion evoked a more complex pattern of activity in which responses were normally evoked in one or other of the muscle pairs and incorporated into the locomotor pattern. 3. In the majority of sites, the stimulation evoked excitatory responses in the flexor muscles of each of the four limbs during that period of the step cycle in which each respective muscle was naturally active; stimulation in the stance phase of locomotion, although less effective, was also capable of producing responses in these muscles. All three ipsilateral extensor muscles studied [long and lateral heads of triceps and vastus lateralis (Tri, TriL, and VL, respectively)] were normally inhibited during their phase of muscle activity, although excitatory responses were occasionally seen. Responses in the contralateral (co) Tri were invariably excitatory and were largest during the period of muscle activity, whereas responses during the period of activity of the coVL were mixed, with both excitatory and inhibitory responses being seen from any one locus. 4. Excitatory responses were normally largest when stimulation was applied during the time that the muscle was active during the locomotor cycle. Responses evoked at times when the muscle was inactive were sometimes larger than those evoked with the animal at rest; such responses were most commonly seen in the hindlimb flexors and in the coVL. 5. In both flexors and extensors of each of the four limbs, the latency of the responses was greatest when the cat was at rest and least for stimuli given during the period of activity of the respective muscle. Average latencies during the period of muscle activity ranged from a minimum of 9.0 +/- 2.6 (SD) ms for inhibitory responses in the ipsilateral Tri and TriL to a maximum of 17.1 +/- 3.0 ms for the responses evoked in the ipsilateral semitendinosus.(ABSTRACT TRUNCATED AT 400 WORDS)     

Prevalence of stereotypical responses to mistuned complex tones in the inferior colliculus

The human auditory system has an exceptional ability to separate competing sounds, but the neural mechanisms that underlie this ability are not understood. Responses of inferior colliculus (IC) neurons to "mistuned" complex tones were measured to investigate possible neural mechanisms for spectral segregation. A mistuned tone is a harmonic complex tone in which the frequency of one component has been changed; that component may be heard as a separate sound source, suggesting that the mistuned tone engages the same mechanisms that contribute to the segregation of natural sounds. In this study, the harmonic tone consisted of eight harmonics of 250 Hz; in the mistuned tone, the frequency of the fourth harmonic was increased by 12% (120 Hz). The mistuned tone elicited a stereotypical discharge pattern, consisting of peaks separated by about 8 ms and a response envelope modulated with a period of 100 ms, which bore little resemblance to the discharge pattern elicited by the harmonic tone or to the stimulus waveform. Similar discharge patterns were elicited from many neurons with a range of characteristic frequencies, especially from neurons that exhibited short-latency sustained responses to pure tones. In contrast, transient and long-latency neurons usually did not exhibit the stereotypical discharge pattern. The discharge pattern was generally stable when the stimulus level or component phase was varied; the major effect of these manipulations was to shift the phase of the response envelope. Simulation of IC responses with a computational model suggested that off-frequency inhibition could produce discharge patterns with these characteristics.