Mode of Operation of Ampullae of Lorenzini of the Skate, Raja

Ampullae of Lorenzini are sensitive electroreceptors. Applied potentials affect receptor cells which transmit synaptically to afferent fibers. Cathodal stimuli in the ampullary lumen sometimes evoke all-or-none "receptor spikes," which are negative-going recorded in the lumen, but more frequently they evoke graded damped oscillations. Cathodal stimuli evoke nerve discharge, usually at stimulus strengths subthreshold for obvious receptor oscillations or spikes. Anodal stimuli decrease any ongoing spontaneous nerve activity. Cathodal stimuli evoke long-lasting depolarizations (generator or postsynaptic potentials) in afferent fibers. Superimposed antidromic spikes are reduced in amplitude, suggesting that the postsynaptic potentials are generated similarly to other excitatory postsynaptic potentials. Anodal stimuli evoke hyperpolarizations of nerves in preparations with tonic activity and in occasional silent preparations; presumably tonic release of excitatory transmitter is decreased. These data are explicable as follows: lumenal faces of receptor cells are tonically (but asynchronously) active generating depolarizing responses. Cathodal stimuli increase this activity, thereby leading to increased depolarization of and increased release of transmitter from serosal faces, which are inexcitable. Anodal stimuli act oppositely. Receptor spikes result from synchronized receptor cell activity. Since cathodal stimuli act directly to hyperpolarize serosal faces, strong cathodal stimuli overcome depolarizing effects of lumenal face activity and are inhibitory. Conversely, strong anodal stimuli depolarize serosal faces, thereby causing release of transmitter, and are excitatory. These properties explain several anomalous features of responses of ampullae of Lorenzini.     

Primitive nervous systems: Peripheral habituation in decerebrate polyclad flatworms

The response to a vibration stimulus recorded from the cords of the ventral submuscular plexus of the polyclad flatworm, Notoplana acticola, consists of a burst of action potentials. The response can be abolished by the application of MgCl₂ to the sea water bathing the preparation. With repeated application of the stimulus, decreasing numbers of action potentials can be measured. This waning responsiveness can be dishabituated by applying a more intense vibration stimulus or with electrical shocks applied directly to the ventral nerve plexus. With electrical stimuli a number of shocks have to be applied before the response can be dishabituated. Changes in responsiveness can be measured simultaneously in a number of sites in the plexus even after the nerves between recording sites have been severed. With different interstimulus intervals the extent of habituation changes. As interstimulus intervals increase from 1 to 5 sec, there appears to be a decrease in responsiveness which recovers when interstimulus intervals become longer than 5 sec.     

Mollusk neuron habituation following extra- and intracellular administration of an ACTH fragment

By means of intracellular recording of the mollusc neuronal activity a study was made of the influence of ACTH4-7--a fragment of the stress hormone,--on elaboration of habituation to tactile stimuli. ACTH was administered immediately after the elaboration of the series I of habituation (25 to 30 stimuli) by means of either perfusion or extra- or intracellular microionophoresis. This was followed by the II-V habituation series (with 15 min intervals between them). Perfusion and extracellular (not intracellular) ACTH4-7 administration produced an acceleration of habituation in the series II, increase of action potentials (AP) thresholds in the response to the first tactile stimulus of series II and cessation of further increase of the AP thresholds in the course of elaboration. Without ACTH, these phenomena were observed beginning with habituation series IV. The ACTH action while influencing the memory is apparently directed to the external side of the excitable neuronal membrane; ACTH seems to accelerate the consolidation process. Any of the methods of ACTH administration produced also an enhancement of EPSPs and an increase in the number of APs in the responses to stimuli, without lowering the thresholds of APs generation. This testifies toi enhanced chemosensitivity of the membrane and may underlie the influence of ACTH on the level of wakefulness.     

In vitro classical conditioning of a gill withdrawal reflex in Aplysia: neural correlates and possible neural mechanisms

An in vitro preparation consisting of the siphon, mantle, gill, and abdominal ganglion undergoes classical conditioning when a weak tactile stimulus (CS) applied to the siphon is paired with a strong tactile stimulus to the gill (UCS). When the stimuli are paired, the CS comes to evoke a gill withdrawal reflex (GWR) which increases in amplitude with training. Only when the stimuli are paired in a classical conditioning paradigm does the CS come to evoke a GWR. With classical conditioning training there is an alteration in the synaptic efficacy between central sensory neurons and central gill motor neurons. Moreover, these changes can be observed in sensory neurons not activated by the CS. The changes observed, as evidence by the number of action potentials evoked in the gill motor neuron do not completely parallel the observed behavioral changes. It is suggested that in addition to changes in the synaptic efficacy at the sensory-motor neuron synapse, other changes in neuronal activity occur at other loci which lead to the observed behavioral changes.     

Primitive nervous systems: electrophysiology of inhibitory events in flatworm nerve cords.

Evoked potentials from the major longitudinal nerve cords of Notoplana acticola are potentiated if the preparation is decerebrated or if certain nerves in the ventral submuscular plexus are severed. Concomitant with an increase in amplitude of the response (in some preparations over fourfold) is a decrease in latency of the response and a decrease in the threshold stimulus intensity needed to evoke activity. Evoked activity can also be depressed with moderate increases in stimulus intensity and is induced some distance from the recording site. The depressant effects can be lifted if the brain is bisected, if the contralateral nerve VI is severed close to the brain, and if the commissures between the two nerves VI are cut. The depressant effects of nerves V and VI are additive. Depression and the subsequent lifting of the inhibition occur in Ringer baths containing either normal sea water or equal mixtures of sea water and isotonic MgCl2. The possibility of inhibitory synapses immune to high concentrations of magnesium ions is discussed.     

Investigation of habituation reaction of neurons of the cat motor cortex

The neuronal and total surface activity of the cortical representation of the motor analyzer in the region of the posterior sigmoid gyrus of the cat brain in response to rhythmical light, sound, and electrical stimuli and their complexes was analyzed. Two groups of neurons were found, of which the first is characterized by a gradual decrease in the number of peaks in the response and by their subsequent disappearance and the second by the absence of a discharge in response to stimulation and by its development before the application of the next stimulus. The first group was comprised of neurons which do not have background activity and the second was made up of neurons with a background activity of 0.4–3.7 imp/sec. This reorganization of the activity of cortical neurons in response to rhythmical stimulation is considered to be a habituation phenomenon.A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 3, No. 3, pp. 245–251, May–June, 1971.     

Effect of sectioning the mammillothalamic tract on neurons of limbic nuclei of the thalamus

Neuronal activity of n. AV (n = 75) and n. AD (n = 55) of the thalamus was recorded extracellularly in unanaesthetized chronic rabbits after complete transection of the mammillo -thalamic tract (MTT). Elimination of this powerful afferent system produced a surprisingly small effect upon spontaneous and evoked neuronal activity. All types of responses were preserved in both nuclei, though some increase of multimodal diffuse tonic responses and decrease of more specialized phasic and complex on-effects occurred in n. AV. In both nuclei short-latency responses (less than 14 ms) to auditory stimuli disappeared. The number of units with dynamic transformations of responses during repeated stimuli application (gradual emergence and shaping of responses, as well as their habituation) decreased 2-3-fold in both nuclei. The impulse activity travelling in MTT seems to be not critical for limbic nuclei sensory reactivity but significant for plasticity of the responses.     

Stroking or Buzzing? A Comparison of Somatosensory Touch Stimuli Using 7 Tesla fMRI

Studying body representations in the brain helps us to understand how we humans relate to our own bodies. The in vivo mapping of the somatosensory cortex, where these representations are found, is greatly facilitated by the high spatial resolution and high sensitivity to brain activation available at ultra-high field. In this study, the use of different stimulus types for somatotopic mapping of the digits at ultra-high field, specifically manual stroking and mechanical stimulation, was compared in terms of sensitivity and specificity of the brain responses. Larger positive responses in digit regions of interest were found for manual stroking than for mechanical stimulation, both in terms of average and maximum t-value and in terms of number of voxels with significant responses to the tactile stimulation. Responses to manual stroking were higher throughout the entire post-central sulcus, but the difference was especially large on its posterior wall, i.e. in Brodmann area 2. During mechanical stimulation, cross-digit responses were more negative than during manual stroking, possibly caused by a faster habituation to the stimulus. These differences indicate that manual stroking is a highly suitable stimulus for fast somatotopic mapping procedures, especially if Brodmann area 2 is of interest.     

Response properties of diencephalic neurons to visual, acoustic and hydrodynamic stimulation in the goldfish, Carassius auratus

We studied the responses to sensory stimulation of three diencephalic areas, the central posterior nucleus of the dorsal thalamus, the anterior tuberal nucleus of the hypothalamus, and the preglomerular complex. Units sensitive to acoustic (500 Hz tone burst), hydrodynamic (25 Hz dipole stimulus) and visual (640 nm light flash) stimuli were found in both the central posterior and anterior tuberal nucleus. In contrast, unit responses or large robust evoked potentials confined to the preglomerular complex were not found. In the central posterior nucleus, most units were unimodal. Many units responded exclusively to visual stimulation and exhibited a variety of temporal response patterns to light stimuli. In the anterior tuberal nucleus of the hypothalamus, most units responded to more than one modality and showed a stronger response decrement to stimulus repetitions than units in the central posterior nucleus. Our data suggest that units in the central posterior nucleus are primarily involved in the unimodal processing of sensory information whereas units in the anterior tuberal nucleus of the hypothalamus may be involved in multisensory integration.     

Electrical potentials from the eye and optic nerve of Strombus: effects of electrical stimulation of the optic nerve.

1. Photic stimulation of the mature eye of Strombus can evoke in the optic nerve 'on' activity in numerous small afferent fibres and repetitive 'off' bursts of afferent impulses in a smaller number of larger fibres. 2. Synchronous invasion of the eye by electrically evoked impulses in small optic nerve fibres (apparently the 'on' afferents, antidromically activated) can evoke a burst of impulses in the larger 'off' fibres which propagate away from the eye. Invasion of the eye via one branch of optic nerve can evoke an answering burst in another branch. 3. Such electrically evoked bursts are similar to light-evoked 'off' bursts with respect to their impulse composition, their ability to be inhibited by illumination of the eye, and their susceptibility to MgCl2 anaesthesia. 4. Invasion of the eye by a train of repetitive electrically evoked impulses in the absence of photic stimulation can give rise to repetitive 'off' bursts as well as concomitant oscillatory potentials in the eye which are similar to those normally evoked by cessation of a photic stimulus. 5. The electrically evoked 'off' bursts appear to be caused by an excitatory rebound following the cessation of inhibitory synaptic input from photoreceptors which can be antidromically activated by electrical stimulation of the optic nerve. 6. The experimental results suggest that the rhythmic discharge of the 'off' fibres evoked by the cessation of a photic stimulus is mediated by the abrupt decrease of inhibitory synaptic input from the receptors.     

Characteristics of the neuronal activity of structures of the hippocampal formation (subiculum)]

Spontaneous and evoked neuronal activity of subiculum was investigated in chronic unanaesthetized rabbits. Rhythmic components of two different types were observed in the spontaneous activity of 58% of units: 4.5-6.0 cps bursts (13%) and 2.0-3.8 cps (45%). Besides their basic frequency, these types of rhythmic activity were differentiated by several additional criteria. 63% of the cells responded to applied sensory stimuli, among which auditory stimuli were the most effective. Multimodal units were slightly more numerous (59%) than unimodal ones. In 50% of the recorded units responses were of tonic type, in 31%--of phasic type, and in 19%--specific on-off responses were observed. In majority of units with repeated stimulations relatively rapid process of habituation developed after a short incremental phase. Interruption of septal afferents resulted in disappearence of 4.5-6.0 cps (theta) bursts, in reduction in the number of tonic responses and in absence of habituation. Similarity of neuronal characteristics in the field CA1 and subiculum is discussed.     

Qualitative differences in the reactions of the spinal nerves of the frog Rana temporaria during chemical stimulation of the skin

Studies have been made on total impulse activity in the skin nerves of the frog during application of acid solutions as well as during tactile stimulation. It was shown that the reactions to various stimuli differ with respect to their pattern and amplitude of integrated response curve which reflects changes in the frequency of total impulsation. In responses to test solutions and tactile stimulation, different units may be involved which specifically react to each of the stimuli. This specificity of single elements is also revealed during changes in the evoked total activity resulting from superficial skin anaesthesia. Possible nature of peripheral structures involved in these reactions is discussed.     

The effect of disconnecting the hippocampus from the septum on the activity of septal neurons]

Extracellular recording of neuronal activity was performed in the medial and lateral septal nuclei (MS and LS) in unanaesthetized rabbits after coagulation of septo-hippocampal connections. The MS neuronal activity had many pathological features. The LS activity was normal in every respect. Spontaneous activity, reactivity to sensory stimuli and main characteristics of responses to sensory stimuli were preserved in LS (and in a part of MS neurones). Sensory effects were augmented in intensity and duration, the number of neurones in LS with theta-bursts increased twofold, theta-bursts were more regular, than in control animals. These effects may be explained by an increase of ascending RF influences, which is supported by the fact of outstanding similarity between sensory and reticular effects in septal neurones after hippocampal disconnection. The number of units with inhibition of activity in response to sensory stimuli decreased, habituation of responses was absent. That means that hippocampal influences are necessary for the organization of inhibitory phenomena in the septum, and, above all, for processes of gradual habituation.     

Independence of the process of habituation from protein synthesis in the mollusk neuron

The influence of protein synthesis blocators--puromicine and gougerotine--on the dynamics of habituation to repeated tactile stimuli in mollusc's neurones was studied microionophoretically. Dynamics of habituation was judged by reduction of impulses number in neurone's response, increase of thresholds and latency of the first synaptic action potential in the response to repeated stimuli. Introduction of the substances produced the greatest effect on the dynamics of thresholds. Puromicine introduction suppressed the habituation only if it was accompanied by generation of epileptiform activity. Gougerotine improved the habituation in the animals with initially weak process. The obtained data testify to the hypothesis that protein synthesis blockade does not impede the process of habituation.     

Electrophysiology of the peripheral nerve net in the polyclad flatworm Freemania litoricola.

1. A diffuse-conducting system close to the dorsal epithelium of the polyclad flatworm Freemania litoricola is described. Tactile stimuli elicit small action potentials which can be conducted around lesions through the body wall. The potentials can occur in bursts or barrages. 2. This conducting system appears to be insensitive to Mg2+ ions. 3. Conduction velocities (0-26--71 m/sec) vary over the animal. Conduction spread in the anterior half of the animal appears to be greater than that in the posterior portion. 4. Response decrement to repeated stimulation can be recorded in the peripheral system but it is not clear if this is due to habituation or fatigue. 5. Conduction from the peripheral net to the brain occurs. Some central units appear to pick up information only, or mainly, through the anterior nerves, while other units can respond to information conducted through the network to nerves of the contralateral side. 6. Different possibilities to account for this system are discussed, and it is suggested that the animals either possess a unique Mg2+ insensitive synaptic nerve-net or else the network is electrically coupled.     

Central neuronal pathways of the cardioinhibitory reflex in the isopod crustacean Bathynomus doederleini

We have already identified central neurons for cardioinhibition and cardioacceleration in Bathynomus, an isopod crustacean. The 1st thoracic ganglion (TG1) has cardioinhibitory neurons, which we call CIs, while the 2nd and 3rd thoracic ganglia (TG2 and TG3) have cardioacceleratory neurons, which we call CA1s and CA2s. We examined neuronal pathways for cardioinhibitory reflexes in whole animal preparations, using intracellular and extracellular recording methods. Cardiac inhibition in response to a variety of external stimuli was mediated by activation of CIs and inhibition of both CAs. When preparations had the ventral nerve cord intact, CIs were activated by excitatory postsynaptic potentials and CAs were inhibited by inhibitory postsynaptic potentials in response to tactile stimuli applied to sensilla setae on appendages and afferent stimuli applied to ganglionic roots of the thoracic ganglia. However, stimulation of ganglionic nerve roots of TG2 and TG3, or tactile stimulation of the body surface, failed to evoke inhibition of CAs in preparations in which both the cerebral ganglion and TG1 had been excised. These results suggest that TG1 is an indispensable central region for the excitation of CI and for inhibition of CA neurons, induced by tactile stimuli and by stimuli applied to nerve roots of TG2 and TG3.     

Representation of moving stimuli by somatosensory neurons.

The findings obtained in neurophysiological and psychophysical investigations using tactile stimuli that move at constant velocity across the skin are reviewed. For certain neurons in the postcentral gyrus of the cerebral cortex (S-I) of macaque monkeys, direction of stimulus motion is a "trigger feature" i.e., moving tactile stimuli evoke vigorous discharge activity in these neurons only if the stimuli are moved in a particular direction across the receptive field. This directional selectivity is maximal when stimulus velocity is between 5 and 50 cm/sec, and falls off rapidly at lower or higher velocities. The capacity for human subjects to correctly identify the direction of stimulus motion on the skin exhibits a similar dependence on stimulus velocity. The similar effects of velocity on neural and psychophysical measures of directional sensitivity support the idea that direction of stimulus motion on the skin can only be recognized if the moving stimulus optimally activates the group of S-I neurons for which that directions of simulus motion is the trigger feature.     

Transfer of habituation between stimulation sites of the siphon withdrawal reflex in Aplysia californica

Habituation of the siphon withdrawal reflex (SWR) can be evoked by iterative tactile stimuli presented to one of several sites, including the siphon and gill. The SWR evoked at an arbitrary "test" site did not habituate when stimuli were presented at 20-min intervals. However, there was a large decrease in the reflex evoked at the test site when the trial was preceded by 10 repetitive stimuli (interstimuli interval = 30 s) presented to the opposite "habituation" site. Transfer of habituation occurred from gill to siphon stimulation sites, and vice versa. There was a concomitant decrease in the excitatory input evoked in the central siphon motor neurons LDS1 and LDS3. Moreover, transfer of habituation occurred after the abdominal ganglion (central nervous system) was removed. There was little change in the magnitude of the control responses or transfer of habituation after deganglionation. Since transfer of habituation between stimulation sites of the SWR was similar to that reported previously for the gill withdrawal reflex, it was suggested that a common mechanism may underlie the two behaviors.     

Configurational pattern discrimination responsible for dishabituation in common toads Bufo bufo (L.): Behavioral tests of the predictions of a neural model

Summary Recently, a neural model of visual pattern discrimination for stimulus-specific habituation was developed, based on previous behavioral studies which demonstrated that toads exhibit a dishabituation hierarchy for different worm-like stimuli. The model suggests that visual objects are represented by temporal coding and predicts that the dishabituation hierarchy changes when the stimulus/background contrast direction is reversed or the stimulus size is varied. The behavioral experiments reported in this paper were designed to test these predictions, (1) For a pair of stimuli from the contrast reversal prediction, the experimental results validated the theory. (2) For a pair of stimuli from the size reduction prediction, the experimental results failed to validate the theory. Further experiments concerning size effects suggest that configurai visual pattern discrimination in toads exhibits size invariance. (3) Inspired by the Groves-Thompson account of habituation, we found that dishabituation by a second stimulus has a separate process from habituation to a first stimulus. This paper serves as an example of a fruitful dialogue between experimentation and modeling, crucial for understanding brain functions.Abbreviations a-h worm-like stimulus patterns - AT anterior thalamus - ERF excitatory receptive field - IRF inhibitory receptive field - RF receptive field - R2 to R4 retinal ganglion cell types - vMP posterior ventromedial pallium     

Sensitization and habituation of command neurons during a defensive reflex in grape snails

All investigated neurones were classified into functional groups depending on their participation in unconditioned avoidance reflex of pneumostome closure. Habituation was produced by tactile stimuli applied with a frequency of 0.1 c/s. Spike responses of neurons participating in processing sensory information habituated gradually, while reactions of command neurones of avoidance behaviour became sensitized to the second or third stimulus of the series. Behavioral reaction was sensitized in parallel with command neurones in spite of habituation in other types of nerve cells. A conclusion is drawn that behavioral habituation is due to waning of responses in all the participating neurones, but the independnet process of sensitization is due to sensitization of command neurones. Probable mechanisms of neuronal sensitization are discussed.