Reverberation of excitation in "hippocampus-entorhinal cortex" slices in rats. Optical recording

Using RH155 voltage-sensitive dye and photodiode array for optical recording, responses to electrical stimuli were investigated in rat brain slices, which included hippocampus and entorhinal cortex. It was shown that single electrical stimulation of the entorhinal cortex, subiculum, or dentate gyrus evoked a potential consecutively spreading from the dentate gyrus to the CA3 and then CA1 hippocampal areas. When the GABAergic inhibition was partially blocked by picrotoxin, the first excitation wave was followed by additional several waves. Such secondary waves were observed in all the hippocampal areas with a constant trial-to-trial latency shift increasing in the direction from the dentate gyrus to CA3 and CA1 areas. Reverberation of activity between the hippocampus and entorhinal cortex is regarded as the most probable cause of appearance of the secondary excitation waves.     

Parametric excitation of surface waves at the boundary between a magnetized plasma and a metal

A study is made of the parametric excitation of potential surface waves propagating in a planar plasma-metal waveguide structure in a magnetic field perpendicular to the plasma-metal boundary. An external, spatially uniform, alternating electric field at the second harmonic of the excited wave is used as the source of parametric excitation. A set of equations is derived that describes the excitation of surface waves due to the onset of decay instability. Expressions for the growth rates in the linear stage of instability are obtained, and the threshold amplitudes of the external electric field above which the parametric instability can occur are found. Analytic expressions for the saturation amplitudes are derived with allowance for the self-interaction of each of the excited waves and the interaction between them. The effect of the plasma parameters and the strength of the external magnetic field on the saturation amplitude, growth rates, and the threshold amplitudes of the pump electric field are analyzed.     

Two Cortical Circuits Control Propagating Waves in Visual Cortex

Visual stimuli produce waves of activity that propagate across the visual cortex of fresh water turtles. This study used a large-scale model of the cortex to examine the roles of specific types of cortical neurons in controlling the formation, speed and duration of these waves. The waves were divided into three components: initial depolarizations, primary propagating waves and secondary waves. The maximal conductances of each receptor type postsynaptic to each population of neurons in the model was systematically varied and the speed of primary waves, durations of primary waves and total wave durations were measured. The analyses indicate that wave formation and speed are controlled principally by feedforward excitation and inhibition, while wave duration is controlled principally by recurrent excitation and feedback inhibition.     

Phantom reflexes: muscle contractions at a frequency not physically present in the input stimuli

In the motor system, the periodic stimulation of one Ia-afferent input produces reflex muscle contractions at the input frequency. However, we observed that when two Ia monosynaptic reflex-afferent inputs are involved the periodic muscle contractions may occur at a frequency physically not present in the afferent inputs even when these inputs are sub-threshold. How can the muscles respond with such phantom reflex contractions at a frequency physically absent in the sub-threshold Ia-afferent input stimuli? Here we provide an explanation for this phenomenon in the cat spinal cord, that we termed “ghost motor response”. We recorded monosynaptic reflexes in the L7 ventral root, intracellular potentials in the motoneurons, and the associated muscular contractions elicited by stimulation of the lateral and medial gastrocnemius nerves. By stimulating with periodic pulses of sub-threshold intensities and distinct frequencies of 2 and 3 Hz the lateral and medial gastrocnemius nerves, respectively, we observed monosynaptic responses and phantom reflex muscle contractions occurring at the fundamental frequency (1 Hz), which was absent in the input stimuli. Thus we observed a reflex ghost motor response at a frequency not physically present in the inputs. We additionally studied the inharmonic case for sub-threshold stimuli and observed muscular contractions occurring at much lower frequencies, which were also conspicuously absent in the inputs. This is the first experimental evidence of a phantom reflex response in the nervous system. The observed behavior was modeled by numerical simulations of a pool of neurons subjected to two different input pulses.     

Conditions for noise reduction and stable encoding of spatial structure by cortical neural networks

Cortical circuits have been proposed to encode information by forming stable spatially structured attractors. Experimentally in the primary somatosensory cortex of the monkey, temporally invariant stimuli lead to spatially structured activity patterns. The purpose of this work is to study a recurrent cortical neural network model with lateral inhibition and examine what effect additive random noise has on the networks' ability to form stable spatially structured representations of the stimulus pattern. We show numerically that this network performs edge enhancement and forms statistically stationary, spatially structured responses when the lateral inhibition is of moderate strength. We then derive analytical conditions on the connectivity matrix that ensure stochasticly stable encoding of the stimulus spatial structure by the network. For stimuli whose strength falls in the near linear region of the sigmoid, we are able to give explicit conditions on the eigenvalues of the connection matrix. Finally, we prove that a network with a connection matrix, where the total excitation and inhibition impinging upon a neural unit are nearly balanced, will yield stable spatial attractor responses. Received: 16 October 1998 / Accepted in revised form: 25 November 1999     

Vulnerability in an excitable medium: analytical and numerical studies of initiating unidirectional propagation.

Cardiac tissue can display unusual responses to certain stimulation protocols. In the wake of a conditioning wave of excitation, spiral waves can be initiated by applying stimuli timed to occur during a period of vulnerability (VP). Although vulnerability is well known in cardiac and chemical media, the determinants of the VP and its boundaries have received little theoretical and analytical study. From numerical and analytical studies of reaction-diffusion equations, we have found that 1) vulnerability is an inherent property of Beeler-Reuter and FitzHugh-Nagumo models of excitable media; 2) the duration of the vulnerable window (VW) the one-dimensional analog of the VP, is sensitive to the medium properties and the size of the stimulus field; and 3) the amplitudes of the excitatory and recovery processes modulate the duration of the VW. The analytical results reveal macroscopic behavior (vulnerability) derived from the diffusion of excitation that is not observable at the level of isolated cells or single reaction units.     

The Spatial Properties of L- and M-Cone Inputs to Electroretinograms That Reflect Different Types of Post-Receptoral Processing

We studied the spatial arrangement of L- and M-cone driven electroretinograms (ERGs) reflecting the activity of magno- and parvocellular pathways. L- and M-cone isolating sine wave stimuli were created with a four primary LED stimulator using triple silent substitution paradigms. Temporal frequencies were 8 and 12 Hz, to reflect cone opponent activity, and 30, 36 and 48 Hz to reflect luminance activity. The responses were measured for full-field stimuli and for different circular and annular stimuli. The ERG data confirm the presence of two different mechanisms at intermediate and high temporal frequencies. The responses measured at high temporal frequencies strongly depended upon spatial stimulus configuration. In the full-field conditions, the L-cone driven responses were substantially larger than the full-field M-cone driven responses and also than the L-cone driven responses with smaller stimuli. The M-cone driven responses at full-field and with 70° diameter stimuli displayed similar amplitudes. The L- and M-cone driven responses measured at 8 and 12 Hz were of similar amplitude and approximately in counter-phase. The amplitudes were constant for most stimulus configurations. The results indicate that, when the ERG reflects luminance activity, it is positively correlated with stimulus size. Beyond 35° retinal eccentricity, the retina mainly contains L-cones. Small stimuli are sufficient to obtain maximal ERGs at low temporal frequencies where the ERGs are also sensitive to cone-opponent processing.     

Stochastic Properties of Discrete Waves of the Limulus Photoreceptor

In the dark-adapted photoreceptor of the horseshoe crab, Limulus, transient discrete depolarizations of the cell membrane, discrete waves, occur in total darkness and their rate of occurrence is increased by illumination. The individual latencies of the discrete waves evoked by a light stimulus often cannot be resolved because the discrete waves overlap in time. The latency of the first discrete wave that follows a stimulus can be determined with reasonable accuracy. We propose a model which allows us to make an estimate of the distribution of the latencies of the individual light-evoked discrete waves, and to predict the latency distribution of the first discrete wave that follows a stimulus of arbitrary intensity-time course from the latency distribution of the first discrete wave that follows a brief flash of light. For low intensity stimuli, the predictions agree well with the observations. We define a response as the occurrence of one or more discrete waves following a stimulus. The distribution of the peak amplitudes of responses suggests that the peak amplitude of individual discrete waves sometimes has a bimodal distribution. The latencies of the two types of discrete waves, however, follow similar distributions. The area under the voltage-time curve of responses that follow equal energy long (1.25 sec) and short (10 msec) light stimuli follows similar distributions, and this suggests that discrete waves summate linearly.     

Changes in the phasic activity of neuron microsystems of the somatosensory cortex of the cat during extinction of activation reactions to unreinforced stimuli

In chronic experiments on cats, three-phasic responses of neuronal microsystems in the cortical somatic area I were studied during habituation of the EEG activation reactions. Repeated stimuli of different modalities were used: electrical pulses to the forepaw, sounds, direct stimulation of the mesencephalic RF. Simultaneously with the extinction of EEG activation reactions, the three-phasic responses of the multiunit activity (MUA) also became progressively extinct: the 1st phase of primary excitation--only a little, the 2nd phase (inhibitory)--greatly, as well as the 3rd phase--the phase of secondary excitation (if it existed at the beginning). The MUA responses to all stimuli show that these neuronal microsystems are polysensory. Relatively to the nonspecific activating RF macrosystem, the investigated neuronal microsystems are autonomous because their two functionally opposed response phases--the 1st excitatory and the 2nd inhibitory--occur against the monotonous excitatory background of the EEG activation. But in some way the neuronal microsystems are connected with the RF-system because of the parallel development of the extinction process.     

Antidromic vasodilation caused by A-delta afferents in the frog

In anesthetized immobilized frog we recorded changes in hind leg volume evoked by electrical stimulation of peripheral end of the sciatic nerve. The ranges of the stimulus amplitudes sufficient to induce vasodilator or vasoconstrictor reactions were estimated. In a separate set of experiments thresholds of A alpha beta, A delta and C-afferent fibers excitation were evaluated by recording waves of compound action potentials in VIII-X dorsal roots. It was found that vasodilation is elicited by the stimuli of virtually the same intensity range as the excitation of A delta afferent fibers, including low threshold one. Consequently we concluded that in frog these myelinated afferent fibers are capable of dilating the blood vessels by antidromic action. This finding is in contrast with antidromic vasodilation in mammals which is known to result mainly from the impulses of the unmyelinated afferent fibers.     

普氏蹄蝠下丘对回声定位信号恒频成分的群体加工

利用听觉诱发电位和计算机叠加平均技术研究了普氏蹄蝠下丘(Inferior colliculus,IC)500-4 000 μm 记录深度间,神经元群对3 个谐波恒频(Constant frequency,CF) 声刺激(CF1 -CF3 )的反应。结果显示,在蝙蝠回声定位信号CF 成分刺激下,其IC 的诱发电位包括2 -4 个波,在1 000 μm 以下的记录区域,3 种刺激均能诱发on-off 反应,on-反应的幅度均在3 000 μm 达最大之后减小(P < 0.001),而潜伏期则逐渐缩短(P < 0.001);CF2 能诱发大幅度的off-反应,而CF1 、CF3 诱发的off-反应幅度较小,随着记录深度的增加,CF2 (P <0. 001)的off-反应潜伏期逐渐缩短,而CF1 (P > 0. 05)和CF3 (P >0. 05)的潜伏期则无此单调性。结果表明神经元群体加工CF1 - CF3 的on-反应存在一致性变化,提示其对行为相关的声信号加工可能存在频率层间复杂的相互作用;对CF 信号加工的off-反应对主频附近声较敏感,提示其可能在恒频-调频蝙蝠的多普勒频移、捕获振翅昆虫信息或种间交流中起着某种作用。     

Linear Relations between Stimulus Amplitudes and Amplitudes of Retinal Action Potentials from the Eye of the Wolf Spider

Incremental photic stimuli have been used to elicit small amplitude retinal action potentials from light-adapted ocelli of the wolf spider, Lycosa baltimoriana (Keyserling) in order to see whether or not the amplitudes of these potentials are linearly related to the stimulus amplitudes. Sine wave variations of light intensity around a mean elicit sine wave variations in potential which contain inappreciable harmonics of the stimulus frequency and whose amplitudes are linearly related to the stimulus amplitudes. Likewise, the responses to the first two periodic Fourier components of incremental rectangular wave stimuli of variable duty cycle are directly proportional to the amplitudes of these components and have phases dependent only on the frequencies and phases of these components. Thirdly, a linear transfer function can be found which describes the amplitudes and phases of responses recorded at different frequencies of sine wave stimulation and this transfer function is sufficient to predict the responses to incremental step stimuli. Finally, it is shown that flash response amplitudes are linearly related to incremental flash intensities at all levels of adaptation. The relations of these linear responses to non-linear responses and to physiological mechanisms of the eye are discussed.     

The Coda of the Transient Response in a Sensitive Cochlea: A Computational Modeling Study

In a sensitive cochlea, the basilar membrane response to transient excitation of any kind–normal acoustic or artificial intracochlear excitation–consists of not only a primary impulse but also a coda of delayed secondary responses with varying amplitudes but similar spectral content around the characteristic frequency of the measurement location. The coda, sometimes referred to as echoes or ringing, has been described as a form of local, short term memory which may influence the ability of the auditory system to detect gaps in an acoustic stimulus such as speech. Depending on the individual cochlea, the temporal gap between the primary impulse and the following coda ranges from once to thrice the group delay of the primary impulse (the group delay of the primary impulse is on the order of a few hundred microseconds). The coda is physiologically vulnerable, disappearing when the cochlea is compromised even slightly. The multicomponent sensitive response is not yet completely understood. We use a physiologically-based, mathematical model to investigate (i) the generation of the primary impulse response and the dependence of the group delay on the various stimulation methods, (ii) the effect of spatial perturbations in the properties of mechanically sensitive ion channels on the generation and separation of delayed secondary responses. The model suggests that the presence of the secondary responses depends on the wavenumber content of a perturbation and the activity level of the cochlea. In addition, the model shows that the varying temporal gaps between adjacent coda seen in experiments depend on the individual profiles of perturbations. Implications for non-invasive cochlear diagnosis are also discussed.     

A comparison of corticospinal activation by magnetic coil and electrical stimulation of monkey motor cortex

The effects of different orientations of a Cadwell round magnetic coil (MC) were compared with each other and with surface electrical stimulation of motor cortex in monkeys anesthetized with pentobarbital or urethane. Recordings were made from within the lateral corticospinal tract, either from axonal populations or with a microelectrode from individual axons. A lateral-sagittally orientated MC directly excited corticospinal neurons at lower stimulus intensity than was required for indirect, i.e., transsynaptic excitation via inputs to corticospinal neurons. By contrast, in 2 out of 3 macaques tested, a vertex-tangential orientation could excite corticospinal neurons indirectly at lower intensities than were required for direct excitation; at higher intensities, direct excitation also occurred.The site of direct corticospinal excitation by a lateral-sagitally oriented MC was inferred by comparing the response variability and latency to MC and surface electrical stimuli. Cathodal stimuli elicited more variable corticospinal population responses and later individual axonal responses than were obtained with anodal stimuli. The variability in response is attributed to interaction between nearby, on-going synaptic bombardment and the stimulus, implying that surface cathodal stimuli directly activate corticospinal neurons at the spike trigger zone (presumably the initial segment). By contrast, the consistency and reduced latency of the corticospinal responses to surface anodal stimuli are attributed to the direct excitation of corticospinal fibers within the white matter. When the stimulus intensity is clearly above threshold, surface anodal and cathodal stimuli can activate corticospinal neurons both directly and indirectly.Direct corticospinal excitation by the MC can resemble the effects of either surface anodal or surface cathodal stimuli. We conclude that the MC can activate corticospinal neurons at the spike trigger zone or their fibers deeper in white matter. The findings in the monkey are used to interpret the effects of different MC orientations in the human.     

Cognitive auditory evoked potentials in children with special language impairment

Perception and discrimination of auditory and speech stimuli in children aged 7-9 years with either receptive (n=6) or expressive (n=5) type of special language impairment and 7 healthy age-matched controls was investigated using evoked potential technique. The measurements were performed with a 32-channel Neuroscan electroencephalographic system. Two types of stimuli were applied, pure tones (1 kHz and 2 kHz) and double syllabi consisting of one consonant and one vocal characteristic of Croatian language. The stimuli were presented in an oddball paradigm, requiring a conscious reaction for the subjects. Latencies and amplitudes of P1, N1, P2, N2, P3, N4, and SW waves were analized, as well as the reaction time and number of responses. There were found no statistically significant difference between children with special language impairment and the control group in average response time and number of responses to tone burst or double syllable. Analysis of variance of all used variables showed a statistically significant difference in P3 and Sw wave latencies after double syllable stimulation, P3 and N4 waves latencies after target stimulation, P2 and Sw wave amplitude; and in N1 wave amplitude after pure tone stimulation. Our study showed that children with speech and language disorder take longer time to perceive and discriminate between either tonal or speech auditory stimuli than children with typical speech and language development.     

Functionally independent components of early event-related potentials in a visual spatial attention task.

Spatial visual attention modulates the first negative-going deflection in the human averaged event-related potential (ERP) in response to visual target and non-target stimuli (the N1 complex). Here we demonstrate a decomposition of N1 into functionally independent subcomponents with functionally distinct relations to task and stimulus conditions. ERPs were collected from 20 subjects in response to visual target and non-target stimuli presented at five attended and non-attended screen locations. Independent component analysis, a new method for blind source separation, was trained simultaneously on 500 ms grand average responses from all 25 stimulus-attention conditions and decomposed the non-target N1 complexes into five spatially fixed, temporally independent and physiologically plausible components. Activity of an early, laterally symmetrical component pair (N1aR and N1aL) was evoked by the left and right visual field stimuli, respectively. Component N1aR peaked ca. 9 ms earlier than N1aL. Central stimuli evoked both components with the same peak latency difference, producing a bilateral scalp distribution. The amplitudes of these components were no reliably augmented by spatial attention. Stimuli in the right visual field evoked activity in a spatio-temporally overlapping bilateral component (N1b) that peaked at ca. 180 ms and was strongly enhanced by attention. Stimuli presented at unattended locations evoked a fourth component (P2a) peaking near 240 ms. A fifth component (P3f) was evoked only by targets presented in either visual field. The distinct response patterns of these components across the array of stimulus and attention conditions suggest that they reflect activity in functionally independent brain systems involved in processing attended and unattended visuospatial events.     

Orienting behaviour during aerial and underwater visual discrimination by the mink (Mustela vison schreber)

Orienting responses by mink during aerial and underwater visual discrimination tests were most frequent when the grating lines subtended angles at the eye near the visual threshold angle. Factorial analysis showed that in air and in water at ranges from 10 to 90 cm most responses occurred at 30 cm discrimination distance and more occurred to marginally supra-threshold than to marginally sub-threshold stimuli. Between media, more responses occurred in air than in water. At longer ranges the mink oriented less readily than at 30 cm but if orienting occurred better discrimination followed than if the mink did not orient.     

Gas exchange and carbon metabolism in two Prosopis species (Fabaceae) from semiarid habitats: effects of elevated CO2, N supply, and N source

Predicting future plant and ecosystem responses to elevated CO(2) also requires an understanding of the role of other factors, especially soil nitrogen. This is particularly challenging for global aridlands where total N and the relative amounts of nitrate and ammonia vary both spatially and seasonally. We measured gas exchange and primary and secondary C metabolites in seedlings of two dominant aridland shrub species (Prosopis flexuosa [S America] and P. glandulosa [N America]) grown at ambient (350 ppm) or elevated (650 ppm) CO(2) and nitrogen at two levels (low [0.8 mM] and high [8.0 mM]) and at either 1 : 1 or 3 : 1 nitrate to ammonia. Whereas elevated CO(2) increased assimilation rate, water use efficiency, and primary carbon metabolites in both species, these increases were strongly contingent upon nitrogen availability. Elevated CO(2) did not increase secondary metabolites (i.e., phenolics). For these important aridland species, the effects of elevated CO(2) are strongly influenced by nitrogen availability and to a lesser extent by the relative amounts of nitrate and ammonia supplied, which underscores the importance of both the amount and chemical composition of soil nitrogen in mediating the potential responses of seedling growth and establishment of aridland plants under future CO(2)-enriched atmospheres.     

Physiology and morphology of descending neurons in pheromone-processing olfactory pathways in the male moth Manduca sexta

1. We have characterized the responses and structure of olfactory descending neurons (DNs) that reside in the protocerebrum (PC) of the brain of male sphinx moths Manduca sexta and project toward thoracic ganglia. 2. Excitatory responses of DNs to pheromone blends were of two general types: (a) brief excitation (BE) that recovered to background in less than 1 s after the stimulus, and (b) long-lasting excitation (LLE) that outlasted the stimulus by greater than or equal to 1 s and, in many cases, as long as 30 s. Individual pheromone components were ineffective in eliciting LLE. 3. Some neurons showing LLE also exhibited state-dependent responses to pheromonal stimuli. When such neurons were in a state of low background firing, stimulation with pheromone blend elicited LLE. When they were in a state of LLE, an identical stimulus reduced firing for 5-10 s after which firing gradually increased to the initial higher level. 4. Thirteen stained DNs were reconstructed from serial sections for detailed analysis of their morphology in the brain. DNs exhibiting LLE had neurites concentrated in the lateral accessory lobes (LALs) in the protocerebrum and adjacent neuropil. Most DNs exhibiting only BE to pheromonal stimuli and other DNs showing responses only to visual or mechanosensory stimuli did not have branches in the LALs.     

Dopaminergic modulation of the P50 auditory-evoked potential in a computer model of the CA3 region of the hippocampus: its relationship to sensory gating in schizophrenia

 We modeled the neuronal circuits that may underlie a sensory-processing deficit associated with schizophrenia. Schizophrenic patients have small P50 auditory-evoked responses to click stimuli compared to normal subjects. The P50 auditory-evoked response is a positive waveform recorded in the EEG approximately 50 ms after the auditory click stimulus. In addition to relatively small amplitudes, schizophrenic patients do not gate or suppress the P50 auditory-evoked response to the second of two paired-click stimuli spaced 0.5 s apart. Neuropleptic medication, which decreases dopaminergic neuronal transmission, increases the amplitude of the P50 auditory-evoked response but does not improve gating. Normal subjects have large P50 auditory-evoked responses to click stimuli when compared to unmedicated schizophrenic patients, and they gate their response to paired click stimuli or have smaller P50 auditory-evoked response amplitudes to the second of two click stimuli spaced 0.5 s apart. Schizophrenic patients do not gate and have similar response amplitudes to both clicks. We hypothesized that the small amplitudes of unmedicated schizophrenic subjects were due to a state of occlusion whereby excessive background noise in local circuits reduced the ability of cells to respond synchronously to sensory input, thereby reducing the amplitude of the P50 waveform in the EEG. Because the P50 auditory-evoked potential amplitudes increased with neuroleptic medication, which reduces dopaminergic neuronal transmission, we hypothesized a role for dopamine in modulating the signal-to-noise (S/N) in the local circuits responsible for sensory gating. To test the hypothesis that modulation of the S/N ratio reduces sensory gating, we developed a model of the effects of dopaminergic neuronal transmission that modulates the S/N in neuronal circuits. The model uses the biologically relevant computer model of the CA3 region of the hippocampus developed in the companion paper [Moxon et al. (2003) Biol Cybern, this volume]. Modified Hebb cell assemblies represented the response of the network to the click stimulus. The results of our model showed that excessive dopaminergic input impaired the ability of cells to respond synchronously to sensory input, which reduced the amplitudes of the P50 evoked responses. Received: 3 December 2001 / Accepted: 23 October 2002 / Published online: 28 February 2003 Correspondence to: K.A. Moxon (e-mail: karen.moxon@drexel.edu, Tel.: +1-215-8951959, Fax: +1-215-8954983) Supported by USPHS, MH01245 & MH58414, MH-01121, and research grants from the Department of Veterans Affairs and the National Alliance for Research on Schizophrenia and Depression.