Hemispheric specialization in dogs for processing different acoustic stimuli

Considerable experimental evidence shows that functional cerebral asymmetries are widespread in animals. Activity of the right cerebral hemisphere has been associated with responses to novel stimuli and the expression of intense emotions, such as aggression, escape behaviour and fear. The left hemisphere uses learned patterns and responds to familiar stimuli. Although such lateralization has been studied mainly for visual responses, there is evidence in primates that auditory perception is lateralized and that vocal communication depends on differential processing by the hemispheres. The aim of the present work was to investigate whether dogs use different hemispheres to process different acoustic stimuli by presenting them with playbacks of a thunderstorm and their species-typical vocalizations. The results revealed that dogs usually process their species-typical vocalizations using the left hemisphere and the thunderstorm sounds using the right hemisphere. Nevertheless, conspecific vocalizations are not always processed by the left hemisphere, since the right hemisphere is used for processing vocalizations when they elicit intense emotion, including fear. These findings suggest that the specialisation of the left hemisphere for intraspecific communication is more ancient that previously thought, and so is specialisation of the right hemisphere for intense emotions.     

Hemispheric asymetry of the evoked electrical activity of the cerebral cortex to letter and non-verbal stimuli]

Average evoked potentials (AEP) were recorded in practically healthy subjects to "meaningless" figures and letters, presented to different halves of the visual field. Analysis of the amplitudes of AEP late components to verbal and non-verbal stimuli reveals hemispheric asymmetry. A higher amplitude of the late positive evoked response (P300) to a "direct" stimulation both by verbal and non-verbal stimuli (in the contralateral field of vision) is recorded in the left hemisphere than in the right one. Similar stimulation of the right hemisphere does not reveal sucha difference. In the left hemisphere the P300 wave is of a clearly greater amplitude to a "direct" stimulation (contralateral visual field) than to an "indirect" one (ipsilateral visual field), regardless of the nature of the stimulus. No such difference is observed in the right hemisphere. The magnitude of the late negative wave (component N200) to non-verbal stimuli is greater in the right hemisphere both in response to "direct" and "indirect" stimulations. No intrahemispheric difference has been found in the amplitude of late evoked responses of the cerebral cortex to verbal and non-verbal stimuli.     

Instrumental heterogeneous responses to 2 successive complex stimuli with a common initial stimulus in the dog

Formation and achievement of heterogenous instrumental reflexes to two consecutive complex stimuli was studied in dogs. Under the action of conditioned complex stimulus tone--Pause--tone, the dogs may be trained not to have a motor response to the first tone presentation, but perform alimentary instrumental reaction only to its repeated administration. Introducing into the experiment not only alimentary but also defensive complex stimulus stone-pause--light resulted in a change in animal's reactions in such a sequence: at first alimentary instrumental reaction was disinhibited under the action of the preparatory stimulus and during the pause, then bieffector responses appeared, further on in most of the dogs motor defensive reactions mainly took place. Trigger stimuli evoked the adequate instrumental reaction.     

Dogs discriminate between barks: The effect of context and identity of the caller

In the present study we explored whether dogs (Canis familiaris) are able to discriminate between conspecific barks emitted in different contexts recorded either from the same or different individuals. Playback experiments were conducted with dogs using barks as stimuli in a habituation–dishabituation paradigm. Barks were recorded in two contexts (stranger at the fence and when the dog was left alone) from different individuals. We found that dogs distinguished between barks emitted in these two contexts and were also able to discriminate between different individuals which were barking in the same context. These findings suggest that dog bark may carry context- and individual-specific information for the conspecifics.     

Effect of masking on interhemispheric and bilateral asymmetry of long-latency auditory evoked potentials

Considering the effects of noise on the amplitude of long-latent auditory evoked potentials, it is concluded that masking interference diminishes the right hemisphere dominance in the processing of the non-speech acoustic information. The decrease is observed both with the monotic and dichotic delivery of signals and noise. With monotic presentation masking interference also decreases the extent of the lateralization of monaurally delivered information predominantly to the contralateral hemisphere.     

Selectivity of visual cortical neurons of rabbits to magnitude of moving stimuli

Unit responses of the rabbit visual cortex were investigated in relation to size of visual stimuli moving in their receptive field. With an increase in size of the stimulus in a direction perpendicular to the direction of movement ("width" of the stimulus) an initial increase in the intensity of the unit response through spatial summation of excitory effects is followed by a decrease through lateral inhibition. This inhibition is observed between zones of the receptive field which behave as activating when tested by a stimulus of small size. Each neuron has its own "preferred" size of stimuli evoking its maximal activation. No direct correlation is found between the "preferred" stimulus size and the size of the receptive field. With a change in stimulus size in the direction of movement ("length" of the stimulus) the responses to stimuli of optimal size may be potentiated through mutual facilitation of the effects evoked by the leading and trailing edges of the stimulus and weakened in response to stimuli of large size. The selective behavior of the neurons with respect to stimulus size is intensified in the case of coordinated changes in their length and width. It is postulated that the series of neurons responding to stimuli of different "preferred" dimensions may constitute a system classifying stimuli by their size.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 4, No. 6, pp. 636–644, November–December, 1972.     

Sex differences in the cerebral lateralization of a cichlid fish when detouring to view emotionally conditioned stimuli

The lateralization of emotion has been described in a variety of animals. The right hemisphere has been implicated in the processing of negative emotions while positive emotions are processed in the left. Most animal studies of this phenomenon to date have used intrinsically emotionally arousing stimuli and there are few examples of lateralized responses to learned emotional triggers. It is known that males and females may demonstrate different patterns of lateralization, and that these sex differences may interact with other variables. We investigated the lateralized response of male and female convict cichlids to emotionally conditioned stimuli. One stimulus was given an appetitive (positive emotional valence) association by pairing with food. The other stimulus was given an aversive (negative emotional valence) association by pairing with a chemical alarm signal. We found that males tend to be more strongly lateralized to aversive stimuli while females are more strongly lateralized when responding to appetitive stimuli.     

Dissociable influences of auditory object vs. spatial attention on visual system oscillatory activity

Given that both auditory and visual systems have anatomically separate object identification ("what") and spatial ("where") pathways, it is of interest whether attention-driven cross-sensory modulations occur separately within these feature domains. Here, we investigated how auditory "what" vs. "where" attention tasks modulate activity in visual pathways using cortically constrained source estimates of magnetoencephalograpic (MEG) oscillatory activity. In the absence of visual stimuli or tasks, subjects were presented with a sequence of auditory-stimulus pairs and instructed to selectively attend to phonetic ("what") vs. spatial ("where") aspects of these sounds, or to listen passively. To investigate sustained modulatory effects, oscillatory power was estimated from time periods between sound-pair presentations. In comparison to attention to sound locations, phonetic auditory attention was associated with stronger alpha (7-13 Hz) power in several visual areas (primary visual cortex; lingual, fusiform, and inferior temporal gyri, lateral occipital cortex), as well as in higher-order visual/multisensory areas including lateral/medial parietal and retrosplenial cortices. Region-of-interest (ROI) analyses of dynamic changes, from which the sustained effects had been removed, suggested further power increases during Attend Phoneme vs. Location centered at the alpha range 400-600 ms after the onset of second sound of each stimulus pair. These results suggest distinct modulations of visual system oscillatory activity during auditory attention to sound object identity ("what") vs. sound location ("where"). The alpha modulations could be interpreted to reflect enhanced crossmodal inhibition of feature-specific visual pathways and adjacent audiovisual association areas during "what" vs. "where" auditory attention.     

The effects of neck flexion on cerebral potentials evoked by visual,auditory and somatosensory stimuli and focal brain blood flow in related sensory cortices

Background

A flexed neck posture leads to non-specific activation of the brain. Sensory evoked cerebral potentials and focal brain blood flow have been used to evaluate the activation of the sensory cortex. We investigated the effects of a flexed neck posture on the cerebral potentials evoked by visual, auditory and somatosensory stimuli and focal brain blood flow in the related sensory cortices.

Methods

Twelve healthy young adults received right visual hemi-field, binaural auditory and left median nerve stimuli while sitting with the neck in a resting and flexed (20° flexion) position. Sensory evoked potentials were recorded from the right occipital region, Cz in accordance with the international 10–20 system, and 2 cm posterior from C4, during visual, auditory and somatosensory stimulations. The oxidative-hemoglobin concentration was measured in the respective sensory cortex using near-infrared spectroscopy.

Results

Latencies of the late component of all sensory evoked potentials significantly shortened, and the amplitude of auditory evoked potentials increased when the neck was in a flexed position. Oxidative-hemoglobin concentrations in the left and right visual cortices were higher during visual stimulation in the flexed neck position. The left visual cortex is responsible for receiving the visual information. In addition, oxidative-hemoglobin concentrations in the bilateral auditory cortex during auditory stimulation, and in the right somatosensory cortex during somatosensory stimulation, were higher in the flexed neck position.

Conclusions

Visual, auditory and somatosensory pathways were activated by neck flexion. The sensory cortices were selectively activated, reflecting the modalities in sensory projection to the cerebral cortex and inter-hemispheric connections.     

Eye and head movements to visual and auditory targets

Coordinated eye-head movements evoked by the presentation of visual, auditory and combined audio-visual targets were studied in 24 human subjects. At 60 deg located targets latencies of eye and head movements were shorter for auditory than for visual stimuli. Latencies were shorter for bisensory than for monosensory targets. The eye and head latencies were differently influenced by the modality of the stimulus when the eccentricity of the target was changed, but not by the variation of the stimulus duration. The different responses of the eye and the head depending on target modality and target eccentricity can be partially attributed to perceptual and central processing mechanisms, and are important to answer the question about the initial event in coordinated eye-head orientation.     

Lateralized behaviour of a non-social cichlid fish (Amatitlania nigrofasciata) in a social and a non-social environment

Cerebral lateralization, the partitioning of cognitive function preferentially into one hemisphere of the brain, is a trait ubiquitous among vertebrates. Some species exhibit population level lateralization, where the pattern of cerebral lateralization is the same for most members of that species; however, other species show only individual level lateralization, where each member of the species has a unique pattern of lateralized brain function. The pattern of cerebral lateralization within a population and an individual has been shown to differ based on the stimulus being processed. It has been hypothesized that sociality within a species, such as shoaling behaviour in fish, may have led to the development and persistence of population level lateralization. Here we assessed cerebral lateralization in convict cichlids (Amatitlania nigrofasciata), a species that does not shoal as adults but that shoals briefly as juveniles. We show that both male and female convict cichlids display population level lateralization when in a solitary environment but only females show population level lateralization when in a perceived social environment. We also show that the pattern of lateralization differs between these two tasks and that strength of lateralization in one task is not predictive of strength of lateralization in the other task.     

Lateralized visual behavior in bottlenose dolphins (Tursiops truncatus) performing audio-visual tasks: the right visual field advantage

Analyzing cerebral asymmetries in various species helps in understanding brain organization. The left and right sides of the brain (lateralization) are involved in different cognitive and sensory functions. This study focuses on dolphin visual lateralization as expressed by spontaneous eye preference when performing a complex cognitive task; we examine lateralization when processing different visual stimuli displayed on an underwater touch-screen (two-dimensional figures, three-dimensional figures and dolphin/human video sequences). Three female bottlenose dolphins (Tursiops truncatus) were submitted to a 2-, 3- or 4-, choice visual/auditory discrimination problem, without any food reward: the subjects had to correctly match visual and acoustic stimuli together. In order to visualize and to touch the underwater target, the dolphins had to come close to the touch-screen and to position themselves using monocular vision (left or right eye) and/or binocular naso-ventral vision. The results showed an ability to associate simple visual forms and auditory information using an underwater touch-screen. Moreover, the subjects showed a spontaneous tendency to use monocular vision. Contrary to previous findings, our results did not clearly demonstrate right eye preference in spontaneous choice. However, the individuals' scores of correct answers were correlated with right eye vision, demonstrating the advantage of this visual field in visual information processing and suggesting a left hemispheric dominance. We also demonstrated that the nature of the presented visual stimulus does not seem to have any influence on the animals' monocular vision choice.     

Functional aspects of evoked alpha and theta responses in humans and cats

A systems theoretical approach was used to compare possible functional roles of theta (4--7 Hz) and alpha (8--15 Hz) response components of brain evoked potentials. These response components were described earlier by Ba\c sar (1980). We recorded EEG and evoked potentials (EPs) from occipital scalp locations in 11 subjects. We used auditory and visual stimuli as inadequate and adequate stimuli, respectively (``cross-modality' measurements). The combined EEG-EP epochs were analysed in frequency domain with fast Fourier transform and adaptive digital filters. Alpha (8--15 Hz) response components turned out to be dependent on whether the stimulus was adequate or not (median amplitude with inadequate vs. adequate stimulation: vs. ). Theta (4--7 Hz) response components were less dependent on stimulus modality (inadequate vs. adequate stimulation: vs. ). In EP recordings the occipital alpha response almost disappeared in the first 250 ms following auditory stimulation. Comparable behaviour was observed in similar experiments with recordings from the cat visual cortex (area 17) and with occipital magnetoencephalographic recordings. Taking into account the above-mentioned previous reports on intracranial recordings in primary sensory areas of the cat brain and preliminary results of magnetoencephalographic measurements, we propose the following hypothesis: alpha responses in a time window of about 250 ms after stimulation might predominantly reflect primary sensory processing whereas the theta responses in the first 250 ms after stimulation might be more involved in supra-modality -- or cross-modality -- associative-cognitive processing. Received: 25 February 1994 / Accepted in revised form: 5 August 1994     

Opponent inhibition: a developmental model of layer 4 of the neocortical circuit.

We model the development of the functional circuit of layer 4 (the input-recipient layer) of cat primary visual cortex. The observed thalamocortical and intracortical circuitry codevelop under Hebb-like synaptic plasticity. Hebbian development yields opponent inhibition: inhibition evoked by stimuli anticorrelated with those that excite a cell. Strong opponent inhibition enables recognition of stimulus orientation in a manner invariant to stimulus contrast. These principles may apply to cortex more generally: Hebb-like plasticity can guide layer 4 of any piece of cortex to create opposition between anticorrelated stimulus pairs, and this enables recognition of specific stimulus patterns in a manner invariant to stimulus magnitude. Properties that are invariant across a cortical column are predicted to be those shared by opponent stimulus pairs; this contrasts with the common idea that a column represents cells with similar response properties.     

Does emotional state influence motor lateralization in California sea lions (<Emphasis Type="Italic">Zalophus californianus</Emphasis>)?

Motor lateralization is a behavioural asymmetry between the left and the right side of an individual due to hemispheric specialization. The right hemisphere controls the left side of the body and the left hemisphere the right side. The right hemisphere processes negative emotions such as fear and frustration, and on the contrary, the left hemisphere processes positive emotions such as happiness. This study, conducted at Parc Asterix Delphinarium (Plailly, France), tested the influence of supposedly positive, negative and neutral emotional situations on four California sea lions’ (Zalophus californianus) motor lateralization while performing a known exercise, here climbing on a stool. Latency between the caretakers’ command and the animals’ response was recorded. The results showed an interindividual variability in the effect of the supposed emotional situations on their motor lateralization and their response latency. The nature and the strength of this effect require deeper investigation by further studies, on a larger number of individuals and contexts.     

Positive potentials ot the human brain in the latent presaccadic period under conditions of stimulation of the leading and nonleading eye

We used backward averaging method to study fast positive presaccadic EEG-potentials under conditions of the monocular stimulation of the leading and nonleading eye. Two schemes of the visual stimulus presentation ("no gap" and "overlap") were used. In the "no gap" condition, potential P1 dominated in the hemispere ipsilateral to a saccade direction. In the "overlap" condition, when the gaze was fixed at the central point, foci of this potential were localized in the sagittal derivations or in the same sites as in the "no gap" conditions. Irrespective on the stimulation scheme, the P2 foci were localized in the hemisphere contralateral to a saccade direction. We assume that the fast positive potentials involve both activation and inhibition processes in visuomotor structures and can be also associated with cognitive presaccadic processes (such as fixation disengage, attention lateralization and a preliminary extraction of motor programs from memory).     

Modification of median nerve somatic evoked potentials by prior median nerve,peroneal nerve,and auditory stimulation

In a recovery function design, changes were measured in the somatic evoked potentials (SEP) to right median nerve (RMN) shocks preceded by stimulation of: the same nerve (RMN-RMN); the left median nerve having primary input to the homologous sensory area in the contralateral hemisphere (LMN-RMN); the right peroneal nerve having primary input to a different region of the same hemisphere (RPN-RMN); and the auditory nerve with primary input to a different sensory modality (AUD-RMN). Eight inter-stimulus intervals ranged from zero (simultaneous) to 2.5 sec. It was assumed that the degree of interaction between evoked potentials would be related to the degree to which common neural structures are activated or modulated in response to the stimuli. Results were: (a) the primary somatosensory response N20-P30 was little influenced by other somatic or auditory stimulation, interaction occurring predominantly in the RMN-RMN condition; (b) with increasing latency, components showed increasing interaction across modalities; (c) preceding homolateral stimulation (RPN-RMN) showed no greater interaction than preceding contralateral stimulation (LMN-RMN); (d) N55-P100 differed from the primary somatosensory response N20-P30 by showing greater interaction with other somatic stimuli; and (e) N140-P190 showed similarly shaped recovery functions across stimulus pairs but significant differences in magnitude of interaction. These results show that components with similar wave form and topographical characteristics can have different neurophysiological properties.     

Mismatch negativity (MMN) for sequences of auditory and visual stimuli: evidence for a mechanism specific to the auditory modality

ERPs to sequences of standard and deviant sinusoidal 100 msec tone pips, high-contrast sinusoidal gratings and to their simultaneously presented combinations were recorded. Mismatch negativity (MMN), an ERP component elicited by deviant stimuli, was estimated for the different stimulus sequences in order to find out whether it reflects modality-specific processes or non-specific attentive phenomena. In addition to the auditory modality, we studied whether the mismatch response could be evoked by a deviant visual stimulus in a visual sequence or by a deviant stimulus in either modality. The results show that only auditory stimuli produced the mismatch response, suggesting that MMN is not a manifestation of a general attentional mechanism but is probably specific to the auditory modality.     

Interweaving and overlapping of evoked potentials

The refractory effect of one stimulus upon the response to a closely following stimulus in a different modality is much less than upon the response to a stimulus in the same modality. It is therefore far more efficient to record responses to stimuli in different modalities concurrently than to record each one separately. We evaluated 2 techniques for concurrent recording. Interweaving involves recording the response to one stimulus in the intervals between recording responses to other stimuli. Overlapping occurs when two or more responses are at times being simulateneously recorded. Interweaving and overlapping reduced the time required to record auditory brain-stem responses, short-latency somatosensory evoked potentials and pattern-reversal visual evoked potentials by a factor of 3 over the time required to record each response separately. Overlapping caused no significant change in the evoked potentials. Depending upon the actual timing schedule, interweaving may distort the evoked potentials if later parts of the response to one stimulus override the evoked potential to a following stimulus. Filtering and randomization of stimulus timing may attenuate the effects of these overriding potentials.     

Dynamics of cortical evoked activity during human learning to discriminate microintervals of time using feedback

The dynamics of the cortical evoked activity in the process of learning of time microintervals (10, 60 and 180 ms) discrimination was studied in healthy adults. Feedback stimulus visually informing of the real correlations of the differentiated pauses facilitates the discrimination. The factor of the visual field does not affect the estimation of brief time intervals. At correct identifications, the P300 wave is recorded with a higher amplitude, than at errors. In the trial following the "nonconfirming" feedback stimulus, the standard and test stimuli evoke in the left hemisphere a greater P300 wave, than in the trial after the "confirming" stimulus. Feedback influence is retained in the long-term memory.