bluuebunny/crossref_metadata_2025 huggingface dataset:
Two Dynamically Distinct Inhibitory Networks in Layer 4 of the Neocortex
Authors: Beierlein Michael , Gibson Jay R. , Connors Barry W.
Published: May 2006

Normal operations of the neocortex depend critically on several types of inhibitory interneurons, but the specific function of each type is unknown. One possibility is that interneurons are differentially engaged by patterns of activity that vary in frequency and timing. To explore this, we studied the strength and short-term dynamics of chemical synapses interconnecting local excitatory neurons (regular-spiking, or RS, cells) with two types of inhibitory interneurons: fast-spiking (FS) cells, and low-threshold spiking (LTS) cells of layer 4 in the rat barrel cortex. We also tested two other pathways onto the interneurons: thalamocortical connections and recurrent collaterals from corticothalamic projection neurons of layer 6. The excitatory and inhibitory synapses interconnecting RS cells and FS cells were highly reliable in response to single stimuli and displayed strong short-term depression. In contrast, excitatory and inhibitory synapses interconnecting the RS and LTS cells were less reliable when initially activated. Excitatory synapses from RS cells onto LTS cells showed dramatic short-term facilitation, whereas inhibitory synapses made by LTS cells onto RS cells facilitated modestly or slightly depressed. Thalamocortical inputs strongly excited both RS and FS cells but rarely and only weakly contacted LTS cells. Both types of interneurons were strongly excited by facilitating synapses from axon collaterals of corticothalamic neurons. We conclude that there are two parallel but dynamically distinct systems of synaptic inhibition in layer 4 of neocortex, each defined by its intrinsic spiking properties, the short-term plasticity of its chemical synapses, and (as shown previously) an exclusive set of electrical synapses. Because of their unique dynamic properties, each inhibitory network will be recruited by different temporal patterns of cortical activity.
Responses of Thalamic Neurons to Input From the Male Genitalia
Authors: Hubscher Charles H. , Johnson Richard D.
Published: May 2006

There have been relatively few electrophysiological studies, in any species, describing the supraspinal processing of inputs from the male genital tract. The thalamus was the focus of the present study. In 11 urethan-anesthetized male rats, subregions of the thalamus were surveyed for neuronal responses to the search stimulus, bilateral electrical stimulation of the dorsal nerve of the penis (DNP). A total of 133 DNP-responsive neurons were found and further tested for degree of somatovisceral convergence from other peripheral structures. Histological reconstruction of the recording sites revealed that the penile-responsive neurons were distributed among various thalamic subregions. These thalamic subregions included the medial-dorsal nuclei and ventral and lateral thalamic subregions (majority of neurons responsive to both tactile and pinch stimulation of the penis) as well as intralaminar, posterior and reticular subregions (majority responsive to pinch only). Taken together, the data demonstrate the existence of thalamic neurons with inputs from the male genitalia with widespread somatovisceral convergence. These neurons likely contribute to the neural circuitries underlying various aspects of penile sensation associated with reproductive and nociceptive events.
Song System Auditory Responses Are Stable and Highly Tuned During Sedation, Rapidly Modulated and Unselective During Wakefulness, and Suppressed By Arousal
Authors: Cardin Jessica A. , Schmidt Marc F.
Published: May 2006

We used auditory responsiveness in the avian song system to investigate the complex relationship between behavioral state and sensory processing in a high-order sensorimotor brain area. We present evidence from recordings in awake, anesthetized, and sleeping male zebra finches ( Taeniopygia guttata) that auditory responsiveness in nucleus HVc is profoundly affected by changes in behavioral state. In anesthetized and sleeping birds, auditory responses were characterized by an increase in firing rate that was selective for the bird's own song (BOS) and highly stable over time. In contrast, HVc responses during wakefulness were extremely variable and transitioned between undetectable and robust levels over short intervals. Surprisingly, auditory responses in awake birds were not selective for the BOS stimulus. The variability of HVc auditory responses in awake birds suggests that, as in mammals, wakefulness is not a uniform behavioral state. Rather, auditory responsiveness likely is continually influenced by variables such as arousal state. We therefore developed several experimental paradigms in which we could manipulate arousal levels during auditory stimulus presentation. In all cases, arousal suppressed HVc auditory responses. This effect was specific to the song system, as auditory responses in Field L, a primary auditory area that is a source of auditory input to HVc, were unaffected. While arousal acts as a negative regulator of HVc auditory responsiveness, the presence and variability of the responses observed in awake, alert birds suggests that other mechanisms, such as attention, may enhance auditory responsiveness. The interplay between behavioral state and sensory processing may regulate song system responsiveness according to the bird's behavioral and social context.
Kinematic Rules for Upper and Lower Arm Contributions to Grasp Orientation
Authors: Marotta J. J. , Medendorp W. P. , Crawford J. D.
Published: May 2006

The purpose of the current study was to investigate the contribution of upper and lower arm torsion to grasp orientation during a reaching and grasping movement. In particular, we examined how the visuomotor system deals with the conflicting demands of coordinating upper and lower arm torsion and maintaining Donders' Law of the upper arm (a behavioral restriction of the axes of arm rotation to a two-dimensional “surface”). In experiment 1, subjects reached out and grasped a target block that was presented in one of 19 orientations (5° clockwise increments from horizontal to vertical) at one position in a vertical presentation board. In experiment 2, target blocks were presented in one of three orientations (horizontal, three-quarter, and vertical) at nine different positions in the presentation board. If reach and grasp commands control the proximal and distal arms separately, then one would only expect the lower arm to contribute to grasp orientations and that Donders' Law would hold for the upper arm—independent of grasp orientations. Instead, as the required grasp orientation increased from horizontal to vertical, there was a significant clockwise torsional rotation in the upper arm, which accounted for 9% of the final vertical grasp orientation, and the lower arm, which accounted for 42%. A linear relationship existed between the torsional rotations of the upper and lower arm, which indicates that the components of the arm rotate in coordination with one another. The location-dependent aspects of upper and lower arm torsion remained invariant, however, yielding consistently shaped Donders' “surfaces” (with different torsional offsets) for different grasp orientations. These observations suggest that the entire arm-hand system contributes to grasp orientation, and therefore, the reach/grasp distinction is not directly reflected in proximal-distal kinematics but is better reflected in the distinction between these coordinated orienting rules and the location-dependent kinematic rules for the upper arm that result in Donders' Law for one given grasp orientation.
Stimulus Specificity and Temporal Dynamics of Working Memory for Visual Motion
Authors: Pasternak Tatiana , Zaksas Daniel
Published: May 2006

When asked to compare two moving stimuli separated by a delay, observers must not only identify stimulus direction but also store it in memory. We examined the properties of this storage mechanism in two macaque monkeys by sequentially presenting two random-dot stimuli, sample and test, in opposite hemifields and introducing a random-motion mask during the delay. The mask interfered with performance only at the precise location of the test, 100–200 ms after the start of the delay, and when its size and speed matched those of the remembered sample. This selective interference suggests that the representation of the motion stimulus in memory preserves its direction, speed, and size and is most fragile shortly after the completion of the encoding phase of the task. This precise preservation of sensory attributes of the motion stimulus suggests that the neural mechanisms involved in the processing of visual motion may also be involved in its storage.
Axotomy-Induced Expression of Calcium-Activated Chloride Current in Subpopulations of Mouse Dorsal Root Ganglion Neurons
Authors: André Sylvain , Boukhaddaoui Hassan , Campo Brice , Al-Jumaily Mohammed , Mayeux Véronique , Greuet Denis , Valmier Jean , Scamps Frédérique
Published: May 2006

Whole cell patch-clamp recordings of calcium-activated chloride current [ ICl(Ca)] were made from adult sensory neurons of naive and axotomized mouse L4–L6 lumbar dorsal root ganglia after 1 day of culture in vitro. A basal ICl(Ca) was specifically expressed in a subset of naive medium-diameter neurons (30–40 μm). Prior nerve injury, induced by sciatic nerve transection 5 days before experiments, increased both ICl(Ca) amplitude and its expression in medium-diameter neurons. Moreover, nerve injury also induced ICl(Ca) expression in a new subpopulation of neurons, the large-diameter neurons (40–50 μm). Small-diameter neurons (inferior to 30 μm) never expressed ICl(Ca). Regulated ICl(Ca) expression was strongly correlated with injury-induced regenerative growth of sensory neurons in vitro and nerve regeneration in vivo. Cell culture on a substrate not permissive for growth, d,l-polyornithine, prevented both elongation growth and ICl(Ca) expression in axotomized neurons. Regenerative growth and the induction of ICl(Ca) expression take place 2 days after injury, peak after 5 days of conditioning in vivo, slowly declining thereafter to control values. The selective expression of ICl(Ca) within medium- and large-diameter neurons conditioned for rapid, efficient growth suggests that these channels play a specific role in postinjury behavior of sensory neuron subpopulations such as neuropathic pain and/or axonal regeneration.
Medial Lateral Extent of Thermal and Pain Sensations Evoked By Microstimulation in Somatic Sensory Nuclei of Human Thalamus
Authors: Ohara Shinji , Lenz Fred A.
Published: May 2006

We explored the region of human thalamic somatic sensory nucleus (ventral caudal, Vc) with threshold microstimulation during stereotactic procedures for the treatment of tremor (124 thalami, 116 patients). Warm sensations were evoked more frequently in the posterior region than in the core. Proportion of sites where microstimulation evoked cool and pain sensations was not different between the core and the posterior region. In the core, sites where both thermal and pain sensations were evoked were distributed similarly in the medial two planes and the lateral plane. In the posterior region, however, warm sensations were evoked more frequently in the lateral plane (10.8%) than in the medial planes (3.9%). No mediolateral difference was found for sites where pain and cool sensations were evoked. The presence of sites where stimulation evoked taste or where receptive and projected fields were located on the pharynx were used as landmarks of a plane located as medial as the posterior part of the ventral medial nucleus (VMpo). Microstimulation in this plane evoked cool, warm, and pain sensations. The results suggest that thermal and pain sensations are processed in the region of Vc as far medial as VMpo. Thermal and pain sensations seem to be mediated by neural elements in a region likely including the core of Vc, VMpo, and other nuclei posterior and inferior to Vc.
Temporal Regulation of Light-Induced Extracellular Signal-Regulated Kinase Activation in the Suprachiasmatic Nucleus
Authors: Butcher Greg Q. , Lee Boyoung , Obrietan Karl
Published: May 2006

Signaling via the p42/p44 mitogen activated protein kinase (MAPK) pathway has been implicated as an intermediate event coupling light to entrainment of the mammalian circadian clock located in the suprachiasmatic nucleus (SCN). To examine how photic input dynamically regulates the activation state of the MAPK pathway, we monitored extracellular signal-regulated kinase (ERK) activation using different light stimulus paradigms. Compared with control animals not exposed to light, a 15 min light exposure during the early night triggered a marked increase in ERK activation and the translocation of ERK from the cytosol to the nucleus. ERK activation peaked 15 min after light onset, then returned to near basal levels within ∼45 min. The MAPK pathway could be reactivated multiple times by light pulses spaced 45 min apart, indicating that the MAPK cascade rapidly resets and resolves individual light pulses into discrete signaling events. Under conditions of constant light (120 min), the time course for ERK activation, nuclear translocation, and inactivation was similar to the time course observed after a 15-min light treatment. The parallels between the ERK inactivation profiles elicited by a 15 and a 120 min light exposure suggest that SCN cells contain a MAPK pathway signal-termination mechanism that limits the duration of pathway activation. This concept was supported by the observation that the small G protein Ras, a regulator of the MAPK pathway, remained in the active, GTP-bound, state under conditions of constant light (120-min duration), indicating that photic information was relayed to the SCN and that SCN cells maintained their responsiveness for the duration of the light treatment. The SCN expressed both nuclear MAPK phosphatases (MKP-1 and MKP-2) and the cytosolic MAPK phosphatase Mkp-3, thus providing mechanisms by which light-induced ERK activation is terminated. Collectively, these observations provide important new information regarding the regulation of the MAPK cascade, a signaling intermediate that couples light to resetting of the SCN clock.
Receptive Field Properties of Human Periodontal Afferents Responding to Loading of Premolar and Molar Teeth
Authors: Johnsen Skjalg E. , Trulsson Mats
Published: May 2006

Impulses in 45 single mechanoreceptive afferents were recorded from the human inferior alveolar nerve with permucosally inserted tungsten microelectrodes. All afferents responded to mechanical stimulation of one or more premolar or molar teeth and most likely innervated their periodontal ligaments. For each afferent, isolated “ramp-and-hold” shaped force profiles of similar magnitudes (252 ± 24 mN; mean ± SD) were applied to the lower first premolar, the second premolar, and the first molar on the recording side. The tooth loads were applied in six directions: lingual, facial, mesial, and distal in the horizontal plane and up and down in the vertical direction of the tooth. The afferents response during the static phase of the stimulus was analyzed. All afferents were slowly adapting, discharging continuously in response to static forces in at least one stimulation direction. Twenty-nine afferents (64%) were spontaneously active, exhibiting an ongoing discharge in the absence of external stimulation. Stimulation of a single tooth was found to excite each afferent most strongly. The most sensitive tooth (MST) was the first premolar for 23, the second premolar for 13, and the first molar for 9 afferents. About half of the afferent population also responded to loading of one or two more teeth. The response profiles of these afferents indicated that the multiple-teeth receptive fields were due to mechanical coupling between the teeth rather than branching of single afferents to innervate several teeth. The afferent responses to loading the mesial and distal halves of the first molars were very similar. Thus both intensive and directional aspects of the afferent response when loading one side of the tooth was preserved to a great extent when loading the other side. When loading the MST, the afferents typically showed excitatory responses in two to four of the six stimulation directions, i.e., the afferents were broadly tuned to direction of tooth loading. In the horizontal plane, the afferent populations at the premolar teeth expressed no clear directional preferences. The afferents at the molar, however, showed a strong directional bias in the distal-lingual direction. In the vertical plane, there was a preference for downward-directed forces with a gradually decreasing sensitivity distally along the dental arch. The present results demonstrate that human periodontal afferents supplying anterior and posterior teeth differ in their capacity to signal horizontal and vertical forces, respectively.
Eyelid Movements: Behavioral Studies of Blinking in Humans Under Different Stimulus Conditions
Authors: VanderWerf Frans , Brassinga Peter , Reits Dik , Aramideh Majid , Ongerboer de Visser Bram
Published: May 2006

The kinematics and neurophysiological aspects of eyelid movements were examined during spontaneous, voluntary, air puff, and electrically induced blinking in healthy human subjects, using the direct magnetic search coil technique simultaneously with electromyographic recording of the orbicularis oculi muscles (OO-EMG). For OO-EMG recordings, surface electrodes were attached to the lower eyelids. To measure the vertical lid displacement, a search coil with a diameter of 3 mm was placed 1 mm from the rim on the upper eyelid on a marked position. Blink registrations were performed from the zero position and from 28 randomly chosen positions. Blinks elicited by electrical stimulation of the supraorbital nerve had shortest duration and were least variable. In contrast, spontaneous blinks had longer duration and greater variability. Blinks induced by air puff had a slightly longer duration and similar variability as electrically induced blinks. There was a correlation between the maximal down phase amplitude and the integrated OO-EMG. Blink duration and maximal down phase amplitude were affected by eye position. Eyes positioned 30° above horizontal displayed the shortest down phase duration and the largest maximal down phase amplitude and velocity. At 30° below horizontal, blinks had the longest total duration, the longest down phase duration, and the lowest maximal down phase amplitude and velocity. The simultaneously recorded integrated OO-EMG was largest in the 30° downward position. In four subjects, the average blinking data showed a linear relation between eye position and OO-EMG, maximal down phase amplitude, and maximal downward velocity.
Efficacy and Short-Term Plasticity at GABAergic Synapses Between Purkinje and Cerebellar Nuclei Neurons
Authors: Pedroarena Christine M. , Schwarz Cornelius
Published: May 2006

Although the entire output of the cerebellar cortex is conveyed to the deep cerebellar nuclei neurons (DCNs) via the GABAergic synapses established by Purkinje cells (PCs), very little is known about the strength and dynamic properties of PC-DCN connections. Here we show that activation of PC-DCN unitary connections induced large conductance changes (11.7 nS) in DCNs recorded in whole cell patch configuration in acute slices, suggesting that activity of single PCs might significantly affect the output of its target neurons. Based on the large unitary quantal content (18) inferred from calculations of PC-DCN quantal size (0.65 nS) and the near absence of failures in synaptic transmission during control conditions, we conclude that PC-DCN connections are highly multi-sited. The analysis of dynamic properties of PC-DCN synapses demonstrated remarkable paired pulse depression (PPD), maximal at short intervals (paired pulse ratio of 0.15 at 7-ms interval). We provide evidence that PPD is presynaptic in origin and release-independent. In addition, multiple pulse stimulation revealed that PC-DCN synapses exhibited larger sensitivity to dynamic than to steady signals. We postulate that the, otherwise paradoxical, combination of marked short-term depression with strong multi-sited connections is optimal to transfer dynamic information at unitary level by performing spatial average of release probability across the numerous release sites. This feature could enable these synapses to encode presynaptic time-varying signals of single PCs as moment-to-moment changes in synaptic strength, a capacity well suited to the postulated role of cerebellum in control of temporal aspects of motor or cognitive behaviors.
Retinogeniculate Synaptic Properties Controlling Spike Number and Timing in Relay Neurons
Authors: Blitz Dawn M. , Regehr Wade G.
Published: May 2006

Retinal ganglion cells (RGC) transmit visual signals to thalamocortical relay neurons in the lateral geniculate nucleus via retinogeniculate synapses. Relay neuron spike patterns do not simply reflect those of RGCs, but the mechanisms controlling this transformation are not well understood. We therefore examined synaptic properties controlling the strength and precision of relay neuron firing in mouse (p28–33) brain slices using physiological stimulation patterns and a combination of current clamp and dynamic clamp. In tonic mode (-55 mV), activation of single RGC inputs elicited stereotyped responses in a given neuron. In contrast, responses in different neurons varied from unreliable, to faithfully following, to a gain in the number of spikes. Dynamic clamp experiments indicated these different responses primarily reflected variability in the amplitudes of the N-methyl-d-aspartate (NMDA) and AMPA components. Each of these components played a distinct role in transmission. The AMPA component evoked a single precisely timed, short-latency spike per stimulus, but efficacy decreased during repetitive stimulation due to desensitization and depression. The NMDA component elicited longer-latency spikes and multiple spikes per stimulus and became more effective during repetitive stimuli that led to NMDA current summation. We found that in burst mode (–75 mV), where low-threshold calcium spikes are activated, AMPA and NMDA components and synaptic plasticity influenced spike number, but no combination enabled relay cells to faithfully follow the stimulus. Thus the characteristics of AMPA and NMDA currents, the ratio of these currents and use-dependent plasticity interact to shape how RGC activity is conveyed to relay neurons.
Electrophysiological Classes of Cat Primary Visual Cortical Neurons In Vivo as Revealed by Quantitative Analyses
Authors: Nowak Lionel G. , Azouz Rony , Sanchez-Vives Maria V. , Gray Charles M. , McCormick David A.
Published: May 2006

To facilitate the characterization of cortical neuronal function, the responses of cells in cat area 17 to intracellular injection of current pulses were quantitatively analyzed. A variety of response variables were used to separate the cells into subtypes using cluster analysis. Four main classes of neurons could be clearly distinguished: regular spiking (RS), fast spiking (FS), intrinsic bursting (IB), and chattering (CH). Each of these contained significant subclasses. RS neurons were characterized by trains of action potentials that exhibited spike frequency adaptation. Morphologically, these cells were spiny stellate cells in layer 4 and pyramidal cells in layers 2, 3, 5, and 6. FS neurons had short-duration action potentials (<0.5 ms at half height), little or no spike frequency adaptation, and a steep relationship between injected current intensity and spike discharge frequency. Morphologically, these cells were sparsely spiny or aspiny nonpyramidal cells. IB neurons typically generated a low frequency (<425 Hz) burst of spikes at the beginning of a depolarizing current pulse followed by a tonic train of action potentials for the remainder of the pulse. These cells were observed in all cortical layers, but were most abundant in layer 5. Finally, CH neurons generated repetitive, high-frequency (350–700 Hz) bursts of short-duration (<0.55 ms) action potentials. Morphologically, these cells were layer 2–4 (mainly layer 3) pyramidal or spiny stellate neurons. These results indicate that firing properties do not form a continuum and that cortical neurons are members of distinct electrophysiological classes and subclasses.
Rhythmic Whisking by Rat: Retraction as Well as Protraction of the Vibrissae Is Under Active Muscular Control
Authors: Berg Rune W. , Kleinfeld David
Published: May 2006

The rhythmic motor activity of the vibrissae that rodents use for the tactile localization of objects provides a model system for understanding patterned motor activity in mammals. The muscles that drive this whisking are only partially fixed relative to bony attachments and thus shift their position along with the movement. As a means to characterize the pattern of muscular dynamics during different patterns of whisking, we recorded electromyogram (EMG) activity from the muscles that propel individual follicles, as well as EMG activity from a muscle group that moves the mystacial pad. The dominant pattern of whisking in our behavioral paradigm, referred to as exploratory whisking, consisted of large amplitude sweeps in the frequency range of 5–15 Hz. The frequency remained remarkably constant within a bout of whisking but changed values between bouts. The extrinsic musculature, which shifts the surface of the pad backwards, was found to be activated in approximate antiphase to that of the intrinsic muscles, which rotate individual vibrissae forward. Thus retraction of the vibrissae was driven by a backward shift in the attachment point of the follicles to the mystacial pad. In a less frequent pattern of whisking, referred to as foveal whisking, the vibrissae are thrust forward and palpate objects with low-amplitude movements that are in the higher frequency range of 15–25 Hz. Protraction of the vibrissae remains driven by the intrinsic muscles, while retraction in this pattern is largely passive. Interestingly, a mechanical argument suggests that activation of the extrinsic muscles during foveal whisking is not expected to affect the angle of the vibrissae. As a means to establish if the phasic control of the intrinsic versus extrinsic muscles depended on sensory feedback, we characterized whisking before and after bilateral transections of the infraorbital branch of the trigeminal sensory nerve. The loss of sensory feedback had no net effect on the antiphase relation between activation of the intrinsic versus extrinsic muscles over the full frequency range for exploratory whisking. These data point to the existence of a dual-phase central pattern generator that drives the vibrissae.
Activity in the Supplementary Motor Area Related to Learning and Performance During a Sequential Visuomotor Task
Authors: Lee Daeyeol , Quessy Stephan
Published: May 2006

Monkeys were trained in a serial reaction time task to produce hand movements according to changing locations of visual targets. In most trials, targets followed the same sequence repeatedly, whereas in other trials targets were presented in random locations or switched unpredictably between two alternative sequences. Single-unit activity was recorded from the caudal supplementary motor area (SMA-proper). Based on the activity associated with random movement sequences, effects of hand position and movement direction were evaluated. Activity was influenced by the hand position in ∼60% of the neurons, and the movement direction influenced the activity of 51% of the neurons. In addition, 37 and 71% of SMA neurons displayed nonstationarity in their activity across successive movements within a given trial and across trials, respectively. Such nonstationarity in the ongoing neural activity and the effects of performance-related variables were evaluated using a regression model and separated from learning-related activity changes. About a third of SMA neurons displayed gradual changes in neural activity related to experience with a movement sequence across trials. Furthermore, about a quarter of SMA neurons showed similar changes within individual trials. When the individual movements included in the frequently repeated movement sequences were introduced unexpectedly, learning-related changes in neural activity were reduced, indicating that many SMA neurons changed their activity in relation to the learning of particular movement sequences. These results suggest that the pattern of neural activity in the cortical network involved in the control of movement sequences can be modified continuously by experience.
Spectral Shape Sensitivity Contributes to the Azimuth Tuning of Neurons in the Cat's Inferior Colliculus
Authors: Poirier Pierre , Samson Frank K. , Imig Thomas J.
Published: May 2006

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.
A Learning and Memory Area in the Octopus Brain Manifests a Vertebrate-Like Long-Term Potentiation
Authors: Hochner Binyamin , Brown Euan R. , Langella Marina , Shomrat Tal , Fiorito Graziano
Published: May 2006

Cellular mechanisms underlying learning and memory were investigated in the octopus using a brain slice preparation of the vertical lobe, an area of the octopus brain involved in learning and memory. Field potential recordings revealed long-term potentiation (LTP) of glutamatergic synaptic field potentials similar to that in vertebrates. These findings suggest that convergent evolution has led to the selection of similar activity-dependent synaptic processes that mediate complex forms of learning and memory in vertebrates and invertebrates.
Excitatory and Inhibitory Intensity Tuning in Auditory Cortex: Evidence for Multiple Inhibitory Mechanisms
Authors: Sutter M. L. , Loftus W. C.
Published: May 2006

The intensity tuning of excitatory and suppressive domain frequency response areas was investigated in 230 cat primary auditory cortical and 92 posterior auditory field neurons. Suppressive domains were explored using simultaneous 2-tone stimulation with one tone at the best excitatory frequency. The intensity tuning of excitatory and suppressive domains was negatively correlated, supporting the hypothesis that inhibitory sidebands are related to excitatory domain intensity tuning. To further test this hypothesis, we compared the slopes of the edges of suppressive bands to the intensity tuning of excitatory domains. Edges of suppressive bands next to excitatory domains had slopes significantly more slanted toward the excitatory area in neurons with intensity-tuned excitatory domains. This relationship was not observed for suppressive band edges not next to the excitatory domain (e.g., the lower edge of lower suppressive bands). This indicates that intensity tuning ultimately observed in the excitatory domain results from overlapping excitatory and inhibitory inputs. In combination with results using forward masking, our results suggest that there are separate early and late sources of inhibition contributing to cortical frequency response areas, and only the early-stage inhibition contributes to excitatory domain intensity tuning.
Effects of Attention and Emotion on Repetition Priming and Their Modulation by Cholinergic Enhancement
Authors: Bentley Paul , Vuilleumier Patrik , Thiel Christiane M. , Driver Jon , Dolan Raymond J.
Published: May 2006

We examined whether behavioral and neural effects of repeating faces are modulated by independent factors of selective attention, emotion, and cholinergic enhancement, during functional MRI. Face repetition occurred either between task-relevant (spatially attended) or task-irrelevant (unattended) stimuli; faces could be fearful or neutral; subjects received either placebo or physostigmine. Under placebo, a reaction time advantage occurred with repetition (i.e., priming) that did not differ between levels of attention, but was attenuated with emotion. Inferior temporo-occipital cortex demonstrated repetition decreases to both attended and unattended faces, and showed either equivalent or greater repetition decreases with emotional compared with neutral faces. By contrast, repetition decreases were attenuated for emotional relative to neutral faces in lateral orbitofrontal cortex. These results distinguish automatic repetition effects in sensory cortical regions from repetition effects modulated by emotion in orbitofrontal cortex, which parallel behavioral effects. Under physostigmine, unlike placebo, behavioral repetition effects were seen selectively for attended faces only, whereas emotional faces no longer impaired priming. Physostigmine enhanced repetition decreases in inferior occipital cortex selectively for attended faces, and reversed the emotional interaction with repetition in lateral orbitofrontal cortex. Thus we show that cholinergic enhancement both augments a neural signature of priming and modulates the effects of attention and emotion on behavioral and neural consequences of repetition.
Recognition by Top-Down and Bottom-Up Processing in Cortex: The Control of Selective Attention
Authors: Graboi Dan , Lisman John
Published: May 2006

Visual recognition is achieved by a hierarchy of bidirectionally connected cortical areas. The entry of signals into higher areas involves the serial sampling of information within a movable window of attention. Here we explore how the cortex can move this window and integrate the sampled information. To make this concrete, we modeled the process of visual word recognition by hierarchical cortical areas representing features, letters, and words. At the start of the recognition process, nodes representing all contextually possible words are active. Simple connectivity rules allow a parallel top-down (T-D) computation of the relative probability of each feature at each location given the set of active words. This information is then used to guide the window of attention to information-rich features (e.g., a feature that is present in the visual image but has lowest probability). Bottom-up processing of this feature excludes words that do not contain it and leads to T-D recomputation of feature probabilities. Recognition occurs after several such cycles when all but one word has been excluded. We show that when 950 words are stored in long-term memory, recognition occurs after an average of 4.9 cycles. Because covert attention can be moved every 20–30 ms, word recognition could be as fast as determined experimentally (<200 ms of cortical processing). This model accounts for the findings that recognition time depends logarithmically on set size, recognition time is reduced when context reduces the number of possible targets, the time to classify a nonword decreases when its approximation to English decreases, and in high level cortex, the firing of neurons tuned to an object increases progressively as its recognition occurs. More generally the model provides a physiologically plausible view of how bi-directional signal flow in cortex guides attention to produce efficient recognition.
Substructure of Direction-Selective Receptive Fields in Macaque V1
Authors: Livingstone Margaret S. , Conway Bevil R.
Published: May 2006

We used two-dimensional (2-D) sparse noise to map simultaneous and sequential two-spot interactions in simple and complex direction-selective cells in macaque V1. Sequential-interaction maps for both simple and complex cells showed preferred-direction facilitation and null-direction suppression for same-contrast stimulus sequences and the reverse for inverting-contrast sequences, although the magnitudes of the interactions were weaker for the simple cells. Contrast-sign selectivity in complex cells indicates that direction-selective interactions in these cells must occur in antecedent simple cells or in simple-cell-like dendritic compartments. Our maps suggest that direction selectivity, and on andoff segregation perpendicular to the orientation axis, can occur prior to receptive-field elongation along the orientation axis. 2-D interaction maps for some complex cells showed elongated alternating facilitatory and suppressive interactions as predicted if their inputs were orientation-selective simple cells. The negative interactions, however, were less elongated than the positive interactions, and there was an inflection at the origin in the positive interactions, so the interactions were chevron-shaped rather than band-like. Other complex cells showed only two round interaction regions, one negative and one positive. Several explanations for the map shapes are considered, including the possibility that directional interactions are generated directly from unoriented inputs.
TRPM8 mRNA Is Expressed in a Subset of Cold-Responsive Trigeminal Neurons From Rat
Authors: Nealen Michele L. , Gold Michael S. , Thut Paul D. , Caterina Michael J.
Published: May 2006

Recent electrophysiological studies of cultured dorsal root and trigeminal ganglion neurons have suggested that multiple ionic mechanisms underlie the peripheral detection of cold temperatures. Several candidate “cold receptors,” all of them ion channel proteins, have been implicated in this process. One of the most promising candidates is TRPM8, a nonselective cationic channel expressed in a subpopulation of sensory neurons that is activated both by decreases in temperature and the cooling compound menthol. However, evidence for the expression of TRPM8 in functionally defined cold-sensitive neurons has been lacking. Here, we combine fluorometric calcium imaging of cultured rat trigeminal neurons with single-cell RT-PCR to demonstrate that there are distinct subpopulations of cold responsive neurons and that TRPM8 likely contributes to cold transduction in one of them. TRPM8 is preferentially expressed within a subset of rapidly responsive, low-threshold (approximately 30°C), cold-sensitive neurons. A distinct class of slowly responsive cold-sensitive neurons that is activated at lower temperatures (approximately 20°C) generally lacks detectable TRPM8 mRNA. Together with previous findings, our data support the notion that cold responsive neurons are functionally heterogeneous.
Functional Coupling of Motor Units Is Modulated During Walking in Human Subjects
Authors: Halliday D. M. , Conway B. A. , Christensen L.O.D. , Hansen N. L. , Petersen N. P. , Nielsen J. B.
Published: May 2006

Time- and frequency-domain analysis of the coupling between pairs of electromyograms (EMG) recorded from leg muscles was investigated during walking in healthy human subjects. For two independent surface EMG signals from the tibialis anterior (TA) muscle, coupling estimated from coherence measurements was observed at frequencies ≤50 Hz, with identifiable peaks occurring in two frequency bands ranging approximately from 8 to 15 and 15 to 20 Hz. The coherence between TA recordings was greatest toward the end of swing, reduced in early swing, and largely absent in midswing. In time-domain estimates constructed from paired TA EMG recordings, a short-lasting central peak indicative of motor-unit synchronization was observed. This feature of motor-unit coupling was also reduced in mid swing. In paired recordings made among triceps surae, quadriceps, and hamstring muscles, a similar pattern of correlation to that for paired TA recordings was observed. However, no significant coupling was observed in recordings for which one EMG recording was made from an ankle flexor/extensor muscle and the other from a knee extensor/flexor muscle. These results demonstrate that for TA a modulation exists in the functional coupling of motor units recruited during swing. The data also indicate that human motoneurons belonging to different muscles are only weakly coupled during walking. This absence of widespread short-term synchronization between the activities of muscles of the leg may provide a basis for the highly adaptive nature of human gait patterns.
Cellular and Network Mechanisms of Slow Oscillatory Activity (<1 Hz) and Wave Propagations in a Cortical Network Model
Authors: Compte Albert , Sanchez-Vives Maria V. , McCormick David A. , Wang Xiao-Jing
Published: May 2006

Slow oscillatory activity (<1 Hz) is observed in vivo in the cortex during slow-wave sleep or under anesthesia and in vitro when the bath solution is chosen to more closely mimic cerebrospinal fluid. Here we present a biophysical network model for the slow oscillations observed in vitro that reproduces the single neuron behaviors and collective network firing patterns in control as well as under pharmacological manipulations. The membrane potential of a neuron oscillates slowly (at <1 Hz) between a down state and an up state; the up state is maintained by strong recurrent excitation balanced by inhibition, and the transition to the down state is due to a slow adaptation current (Na+-dependent K+ current). Consistent with in vivo data, the input resistance of a model neuron, on average, is the largest at the end of the down state and the smallest during the initial phase of the up state. An activity wave is initiated by spontaneous spike discharges in a minority of neurons, and propagates across the network at a speed of 3–8 mm/s in control and 20–50 mm/s with inhibition block. Our work suggests that long-range excitatory patchy connections contribute significantly to this wave propagation. Finally, we show with this model that various known physiological effects of neuromodulation can switch the network to tonic firing, thus simulating a transition to the waking state.
Spatial Organization of Receptive Fields of V1 Neurons of Alert Monkeys: Comparison With Responses to Gratings
Authors: Kagan Igor , Gur Moshe , Snodderly D. Max
Published: May 2006

We studied the spatial organization of receptive fields and the responses to gratings of neurons in parafoveal V1 of alert monkeys. Activating regions (ARs) of 228 cells were mapped with increment and decrement bars while compensating for fixational eye movements. For cells with two or more ARs, the overlap between ARs responsive to increments (INC) and ARs responsive to decrements (DEC) was characterized by a quantitative overlap index (OI). The distribution of overlap indices was bimodal. The larger group (78% of cells) was composed of complex cells with strongly overlapping ARs (OI ≥ 0.5). The smaller group (14%) was composed of simple cells with minimal spatial overlap of ARs (OI ≤ 0.3). Simple cells were preferentially located in layers dominated by the magnocellular pathway. A third group of neurons, the monocontrast cells (8%), responded only to one sign of contrast and had more sustained responses to flashed stimuli than other cells. One hundred fourteen neurons were also studied with drifting sinusoidal gratings of various spatial frequencies and window widths. For complex cells, the relative modulation (RM, the ratio of the 1st harmonic to the mean firing rate), ranged from 0.6 ± 0.4 to 1.1 ± 0.5 (mean ± SD), depending on the stimulus conditions and the mode of correction for eye movements. RM was not correlated with the degree of overlap of ARs, indicating that the spatial organization of receptive fields cannot reliably be predicted from RM values. In fact, a subset of complex cells had RM > 1, the traditional criterion for identifying simple cells. However, unlike simple cells, even those complex cells with high RM could exhibit diverse nonlinear responses when the spatial frequency or window size was changed. Furthermore, the responses of complex cells to counterphase gratings were predominantly nonlinear even harmonics. These results show that RM is not a robust test of linearity. Our results indicate that complex cells are the most frequently encountered neurons in primate V1, and their behavior needs to receive more emphasis in models of visual function.
Canceling of Pursuit and Saccadic Eye Movements in Humans and Monkeys
Authors: Kornylo Krista , Dill Natalie , Saenz Melissa , Krauzlis Richard J.
Published: May 2006

The countermanding paradigm provides a useful tool for examining the mechanisms responsible for canceling eye movements. The key feature of this paradigm is that, on a minority of trials, a stop signal is introduced some time after the appearance of the target, indicating that the subject should cancel the incipient eye movement. If the delay in giving the stop signal is too long, subjects fail to cancel the eye movement to the target stimulus. By modeling this performance as a race between a go process triggered by the appearance of the target and a stop process triggered by the appearance of the stop signal, it is possible to estimate the processing interval associated with canceling the movement. We have now used this paradigm to analyze the canceling of pursuit and saccades. For pursuit, we obtained consistent estimates of the stop process regardless of our technique or assumptions—it took 50–60 ms to cancel pursuit in both humans and monkeys. For saccades, we found different values depending on our assumptions. When we assumed that saccade preparation was under inhibitory control up until movement onset, we found that saccades took longer to cancel (humans: ∼110, monkeys: ∼80 ms) than pursuit. However, when we assumed that saccade preparation includes a final “ballistic” interval not under inhibitory control, we found that the same rapid stop process that accounted for our pursuit results could also account for the canceling of saccades. We favor this second interpretation because canceling pursuit or saccades amounts to maintaining a state of fixation, and it is more parsimonious to assume that this involves a single inhibitory process associated with the fixation system, rather than two separate inhibitory processes depending on which type of eye movement will not be made. From our behavioral data, we estimate that this ballistic interval has a duration of 9–25 ms in monkeys, consistent with the known physiology of the final motor pathways for saccades, although we obtained longer values in humans (28–60 ms). Finally, we examined the effect of trial sequence during the countermanding task and found that pursuit and saccade latencies tended to be longer if the previous trial contained a stop signal than if it did not; these increases occurred regardless of whether the preceding trial was associated with the same or different type of eye movement. Together, these results suggest that a common inhibitory mechanism regulates the initiation of pursuit and saccades.