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Abstract Background Early auditory experiences are a prerequisite for speech and language acquisition. In healthy children, phoneme discrimination abilities improve for native and degrade for unfamiliar, socially irrelevant phoneme contrasts between 6 and 12 months of age as the brain tunes itself to, and specializes in the native spoken language. This process is known as perceptual narrowing, and has been found to predict normal native language acquisition. Prematurely born infants are known to be at an elevated risk for later language problems, but it remains unclear whether these problems relate to early perceptual narrowing. To address this question, we investigated early neurophysiological phoneme discrimination abilities and later language skills in prematurely born infants and in healthy, full-term infants. Results Our follow-up study shows for the first time that perceptual narrowing for non-native phoneme contrasts found in the healthy controls at 12 months was not observed in very prematurely born infants. An electric mismatch response of the brain indicated that whereas full-term infants gradually lost their ability to discriminate non-native phonemes from 6 to 12 months of age, prematurely born infants kept on this ability. Language performance tested at the age of 2 years showed a significant delay in the prematurely born group. Moreover, those infants who did not become specialized in native phonemes at the age of one year, performed worse in the communicative language test (MacArthur Communicative Development Inventories) at the age of two years. Thus, decline in sensitivity to non-native phonemes served as a predictor for further language development. Conclusion Our data suggest that detrimental effects of prematurity on language skills are based on the low degree of specialization to native language early in development. Moreover, delayed or atypical perceptual narrowing was associated with slower language acquisition. The results hence suggest that language problems related to prematurity may partially originate already from this early tuning stage of language acquisition.

N1 of event-related potentials (ERPs) is augmented in amplitude in ∼50-150 ms by occasional changes (deviants) in the physical features of a sound repeated at intervals of from ∼400 ms to seconds (standard). The release-from-refractoriness hypothesis links the N1 augmentation to a deviant-feature-specific neural population that is fresh to fully respond as opposed to a standard-feature-specific neural population that is unresponsive due to its post-response refractoriness. The present work explored this hypothesis in the context of ERP studies, behavioral habituation studies and studies on stimulus-specific adaptation (SSA). The idea of hundreds of milliseconds neural population-level refractoriness was observed to be founded upon negative N1 evidence (no observable effect of dishabituating stimuli on N1 to standards - the null hypothesis retained) and merely supported by positive N1 evidence (null hypotheses rejected). This idea was also found to be directly challenged by positive N1 evidence. No conclusive network- or single-neuron-level evidence was found for the refractoriness. Therefore, the validity of the release-from-refractoriness hypothesis of N1 to guide psychophysiological research needs reassessment., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Spectrotemporally complex sounds carry important information for acoustic communication. Among the important features of these sounds is the temporal duration. An event-related potential called mismatch negativity indexes auditory change detection in humans. An analogous response (mismatch response) has been found to duration changes in speech sounds in rats but not yet in mice. We addressed whether mice show this response, and, if elicited, whether this response is functionally analogous to mismatch negativity or whether adaptation-based models suffice to explain them. Auditory-evoked potentials were epidurally recorded above the mice auditory cortex. The differential response to the changes in a repeated human speech sound /a/ was elicited 53-259 ms post-change (oddball condition). The differential response was observable to the largest duration change (from 200 to 110 ms). Any smaller (from 200 to 120-180 ms at 10 ms steps) duration changes did elicit an observable response. The response to the largest duration change did not robustly differ in amplitude from the response to the change-inducing sound presented without its repetitive background (equiprobable condition). The findings suggest that adaptation may suffice to explain responses to duration changes in spectrotemporally complex sounds in anaesthetized mice. The results pave way for development of a variety of murine models of acoustic communication., (© 2019 Federation of European Neuroscience Societies and John Wiley & Sons Ltd.)

Volume and number are addressed separately rather than combined in perception research. Yet, our everyday problems often involve summed continuous volumes of countable solid objects with partial depth cues (e.g., food items). The participants were presented with a set of black-and-white silhouettes of spheres that independently varied in numerosity (from 1 to 6) and total volume (2, 4, 6, or 8), and an adjacent silhouette of a partially filled cylinder. They judged how much the silhouetted sphere(s) in the set would raise the level of the cylinder content if the spheres were immersed into that content. Higher total volumes and numerosities of the spheres were judged slower and underestimated. Lower total volumes and numerosities were judged faster and overestimated. These effects strongly reflected the total silhouette area of the spheres in a set. The discontinuous effect of numerosity on judgment accuracy and speed suggested separate judgment modes below and above Numerosity 3.

The brain hosts a primitive number sense to non-symbolically represent numerosities of objects or events. Small exact numerosities (~4 or less) can be individuated in parallel. In contrast, large numerosities (more than ~4) can only be approximated. However, whether small numerosities can be approximated without their parallel individuation remains unclear. Parallel individuation is suggested to be an attentive process and numerical approximation an automatic process. We, therefore, tested whether small numerosities can be represented preattentively. We recorded adult humans׳ event-related potentials (ERPs) and behavioral responses to 300-ms sequences of six tones (each of either 440 Hz or 660 Hz in frequency). Mostly, a sequence was of 3 tones of each frequency. Occasionally (P=0.1), the numerosities were 4 and 2 (minor changes) or 5 and 1 (major changes). Mismatch negativity (MMN), but no later attention-related positive-polarity ERPs, was observed to the major but not to the minor changes during a visual non-numerical task. In a following attentive task, behavioral responses even to major changes resulted in a very low hit rates (0.11 for major and 0.023 for minor changes) and yet an above-zero false-alarm rate (0.052). The findings support a view that small numerosities of objects can be automatically approximated independently of their attentive individuation., (Copyright © 2015 Elsevier B.V. All rights reserved.)

The human brain can automatically detect auditory changes, as indexed by the mismatch negativity of event-related potentials. The mechanisms that underlie this response are poorly understood. We recorded primary auditory cortical and hippocampal (dentate gyrus, CA1) local-field potentials to serial tones in urethane-anesthetized rats. In an oddball condition, a rare (deviant) tone (p=0.11) randomly replaced a repeated (standard) tone. The deviant tone was either lower (2200, 2700, 3200, 3700Hz) or higher (4300, 4800, 5300, 5800Hz) in frequency than the standard tone (4000Hz). In an equiprobability control condition, all nine tones were presented at random (p=0.11). Differential responses to deviant tones relative to the standard tone were found in the auditory cortex and the dentate gyrus but not in CA1. Only in the dentate gyrus, the responses were found to be standard- (i.e., oddball condition-) specific. In the auditory cortex, the sound frequencies themselves sufficed to explain their generation. These findings tentatively suggest dissociation among non-contextual afferent, contextual afferent and auditory change detection processes. Most importantly, they remind us about the importance of strict control of physical sound features in mismatch negativity studies in animals., (Copyright © 2015 Elsevier B.V. All rights reserved.)

Human infants are able to detect changes in grammatical rules in a speech sound stream. Here, we tested whether rats have a comparable ability by using an electrophysiological measure that has been shown to reflect higher order auditory cognition even before it becomes manifested in behavioral level. Urethane-anesthetized rats were presented with a stream of sequences consisting of three pseudowords carried out at a fast pace. Frequently presented "standard" sequences had 16 variants which all had the same structure. They were occasionally replaced by acoustically novel "deviant" sequences of two different types: structurally consistent and inconsistent sequences. Two stimulus conditions were presented for separate animal groups. In one stimulus condition, the standard and the pattern-obeying deviant sequences had an AAB structure, while the pattern-violating deviant sequences had an ABB structure. In the other stimulus condition, these assignments were reversed. During the stimulus presentation, local-field potentials were recorded from the dura, above the auditory cortex. Two temporally separate differential brain responses to the deviant sequences reflected the detection of the deviant speech sound sequences. The first response was elicited by both types of deviant sequences and reflected most probably their acoustical novelty. The second response was elicited specifically by the structurally inconsistent deviant sequences (pattern-violating deviant sequences), suggesting that rats were able to detect changes in the pattern of three-syllabic speech sound sequence (i.e., location of the reduplication of an element in the sequence). Since all the deviant sound sequences were constructed of novel items, our findings indicate that, similarly to the human brain, the rat brain has the ability to automatically generalize extracted structural information to new items.

It is not known whether animals can, similarly to humans, categorize auditory objects based on an abstract rule in combining their physical features. We recorded local-field potentials from the dura above the primary auditory cortex in urethane-anesthetized rats presented with sound series occasionally violating a rule (e.g., "the higher the frequency, the weaker the intensity"). In a separate control condition, the same frequency and intensity levels were applied in the sound objects, but they obeyed no rule. Responses found selectively to the violations of the rule suggest that an abstract rule was represented in the rat brain, enabling auditory categorization., (Copyright © 2014 Society for Psychophysiological Research.)

Eyeblinks, whether reflexive or voluntary, play an important role in protecting our vision. When viewing pictures, reflexive eyeblinks are known to be modulated by the emotional state induced thereby. More specifically, the hedonic valence (unpleasantness-pleasantness) induced by the picture has been shown to have a linear relationship with the amplitude of a startle blink elicited during picture viewing. This effect has been attributed to congruence between an ongoing state and task demands: an unpleasant emotional state is assumed to bias our attention towards potentially harmful stimuli, such as startle tones. However, recent research suggests that the valence-specific modulation may not be limited to the sensory parts of the reflexive pathway related to startle responses. Here, we examined the effect of emotional picture viewing on voluntary (in response to a written command) eyeblinks in adult humans. Emotional modulation of startle blinks was also evaluated. We found that when viewing unpleasant pictures, the amplitude of reflexive eyeblinks was augmented, but the amplitude of voluntary eyeblinks was unaffected. Nevertheless, the response latencies of voluntary eyeblinks were found to be delayed during the viewing of pleasant and unpleasant relative to neutral pictures. We conclude that these results support the theory that emotional experience augments sensory processing specific to potentially harmful stimuli. Further, the emotional state seems not to exert an effect on voluntarily elicited motor activity.

Any change in the invariant aspects of the auditory environment is of potential importance. The human brain preattentively or automatically detects such changes. The mismatch negativity (MMN) of event-related potentials (ERPs) reflects this initial stage of auditory change detection. The origin of MMN is held to be cortical. The hippocampus is associated with a later generated P3a of ERPs reflecting involuntarily attention switches towards auditory changes that are high in magnitude. The evidence for this cortico-hippocampal dichotomy is scarce, however. To shed further light on this issue, auditory cortical and hippocampal-system (CA1, dentate gyrus, subiculum) local-field potentials were recorded in urethane-anesthetized rats. A rare tone in duration (deviant) was interspersed with a repeated tone (standard). Two standard-to-standard (SSI) and standard-to-deviant (SDI) intervals (200 ms vs. 500 ms) were applied in different combinations to vary the observability of responses resembling MMN (mismatch responses). Mismatch responses were observed at 51.5-89 ms with the 500-ms SSI coupled with the 200-ms SDI but not with the three remaining combinations. Most importantly, the responses appeared in both the auditory-cortical and hippocampal locations. The findings suggest that the hippocampus may play a role in (cortical) manifestation of MMN.

Change blindness is a failure of reporting major changes across consecutive images if separated, e.g., by a brief blank interval. Successful change detection across interrupts requires focal attention to the changes. However, findings of implicit detection of visual changes during change blindness have raised the question of whether the implicit mode is necessary for development of the explicit mode. To this end, we recorded the visual mismatch negativity (vMMN) of the event-related potentials (ERPs) of the brain, an index of implicit pre-attentive visual change detection, in adult humans performing an oddball-variant of change blindness flicker task. Images of 500 ms in duration were presented repeatedly in continuous sequences, alternating with a blank interval (either 100 ms or 500 ms in duration throughout a stimulus sequence). Occasionally (P = 0.2), a change (referring to color changes, omissions, or additions of objects or their parts in the image) was present. The participants attempted to explicitly (via voluntary button press) detect the occasional change. With both interval durations, it took 10-15 change presentations in average for the participants to eventually detect the changes explicitly in a sequence, the 500 ms interval only requiring a slightly longer exposure to the series than the 100 ms one. Nevertheless, prior to this point of explicit detectability, the implicit detection of the changes vMMN could only be observed with the 100 ms intervals. These findings of explicit change detection developing with and without implicit change detection may suggest that the two modes of change detection recruit independent neural mechanisms.

Understanding speech is based on neural representations of individual speech sounds. In humans, such representations are capable of supporting an automatic and memory-based mechanism for auditory change detection, as reflected by the mismatch negativity (MMN) of event-related potentials. There are also findings of neural representations of speech sounds in animals, but it is not known whether these representations can support the change detection mechanism analogous to that underlying the MMN in humans. To this end, we presented synthesized spoken syllables to urethane-anesthetized rats while local field potentials were epidurally recorded above their primary auditory cortex. In an oddball condition, a deviant stimulus /ga/ or /ba/ (probability 1:12 for each) was rarely and randomly interspersed between frequently presented standard stimulus /da/ (probability 10:12). In an equiprobable condition, 12 syllables, including /da/, /ga/, and /ba/, were presented in a random order (probability 1:12 for each). We found evoked responses of higher amplitude to the deviant /ba/, albeit not to /ga/, relative to the standard /da/ in the oddball condition. Furthermore, the responses to /ba/ were higher in amplitude in the oddball condition than in the equiprobable condition. The findings suggest that anesthetized rat's brain can form representations of human speech sounds, and that these representations can support the memory-based change detection mechanism analogous to that underlying the MMN in humans. Our findings show a striking parallel in speech processing between humans and rodents and may thus pave the way for feasible animal models of memory-based change detection.

Any occasional changes in the acoustic environment are of potential importance for survival. In humans, the preattentive detection of such changes generates the mismatch negativity (MMN) component of event-related brain potentials. MMN is elicited to rare changes ('deviants') in a series of otherwise regularly repeating stimuli ('standards'). Deviant stimuli are detected on the basis of a neural comparison process between the input from the current stimulus and the sensory memory trace of the standard stimuli. It is, however, unclear to what extent animals show a similar comparison process in response to auditory changes. To resolve this issue, epidural potentials were recorded above the primary auditory cortex of urethane-anesthetized rats. In an oddball condition, tone frequency was used to differentiate deviants interspersed randomly among a standard tone. Mismatch responses were observed at 60-100 ms after stimulus onset for frequency increases of 5% and 12.5% but not for similarly descending deviants. The response diminished when the silent inter-stimulus interval was increased from 375 ms to 600 ms for +5% deviants and from 600 ms to 1000 ms for +12.5% deviants. In comparison to the oddball condition the response also diminished in a control condition in which no repetitive standards were presented (equiprobable condition). These findings suggest that the rat mismatch response is similar to the human MMN and indicate that anesthetized rats provide a valuable model for studies of central auditory processing.

In this article, we present a new view on the nature of cognitive processes suggesting that there is a common core, viz., automatic sensory-cognitive processes that form the basis for higher-order cognitive processes. It has been shown that automatic sensory-cognitive processes are shared by humans and various other species and occur at different developmental stages and even in different states of consciousness. This evidence, based on the automatic electrophysiological change-detection response mismatch negativity (MMN), its magnetoencephalographic equivalent MMNm, and behavioral data, indicates that in audition surprisingly complex processes occur automatically and mainly in the sensory-specific cortical regions. These processes include, e.g. stimulus anticipation and extrapolation, sequential stimulus-rule extraction, and pattern and pitch-interval encoding. Furthermore, these complex perceptual-cognitive processes, first found in waking adults, occur similarly even in sleeping newborns, anesthetized animals, and deeply sedated adult humans, suggesting that they form the common perceptual-cognitive core of cognitive processes in general. Although the present evidence originates mainly from the auditory modality, it is likely that analogous evidence could be obtained from other sensory modalities when measures corresponding to those used in the study of the auditory modality become available.

Affective states are known to modulate reflexive actions. Aversive states potentiate defensive reflexes while appetitive states diminish them. The present study examined whether the same holds for associatively learned defensive eyeblinks to mild, initially neutral auditory stimuli. First, delay eyeblink conditioning was applied to human participants while they viewed emotionally neutral images. Next, the conditioned eyeblink responses (CRs) of the participants were tested during the viewing of unpleasant, neutral, or pleasant images. The most vigorous CRs were found during the unpleasant images, although they did not differ between neutral and pleasant images. The results add to the motivational priming hypothesis by demonstrating its partial applicability to associatively learned defensive behaviour.

Humans are able to attentively discriminate number from 6 months of age. However, the age of the emergence of this ability at the pre-attentive stage of processing remains unclear. Event-related potentials (ERPs) were recorded in newborn human infants aged from 3 to 5 days. At 500-ms intervals, the infants were passively exposed to 200-ms sequences of four tones. Each tone could be either 1000 or 1500 Hz in frequency. In most sequences (standards), the ratio of the tones of one frequency to those of the other frequency in a sequence was 2 : 2. In the remaining sequences (deviants, P = 0.1), this ratio was either 3 : 1 or 4 : 0. The mismatch response of ERPs could not be found for 3 : 1 deviants, but it was a robust finding for 4 : 0 deviants, showing the neurophysiological ability of the infants to register the larger deviant-standard difference. The findings suggest very early sensitivity to auditory numerical information in infancy.

It remains unclear whether the mismatch negativity of event-related potentials (ERPs) in vision resembles its auditory counterpart in terms of memory relatedness. We recorded ERPs to visual bars in adult humans engaged in an auditory task. In one condition, a bar ('standard') repeated at 400- or 1100-ms non-stimulated intervals was rarely (P = 0.1) replaced by another bar of a different orientation ('deviant'). In the other condition (400-ms intervals), the occurrences of the standards were replaced by 10 (P = 0.1 each) bars of different orientations, including that of the deviant ('control-deviant'). Deviants shifted ERPs towards negative polarity relative to standards in occipital electrodes and towards positive polarity in frontal electrodes at 185-205 ms post-stimulus but only when 400-ms non-stimulated intervals were applied. Furthermore, the shift existed even relative to ERPs to control-deviants. The findings suggest that, as in audition, vision supports the detection of voluntarily unattended changes per se within the constraints of sensory memory. The findings also pave the way for the future exploration of both intact and impaired memory-based visual processing and memory capacity.

The human brain can automatically quantify objects and events. The mismatch negativity (MMN) of the event-related potential (ERP) was recently found to reflect one such ability in the auditory modality. The present study aimed to further validate the finding. ERPs were recorded in adult humans who passively listened to a series of 300-ms sequences of tones. The sequences occurred at 300-ms silent intervals. They comprised a total of six tones that each was either 440 or 660 Hz in frequency. MMN was found for a rare 4:2 ratio interspersed with a frequent 3:3 ratio between the tones of the different frequencies in a sequence, suggesting the ability of the human brain to preattentively discriminate numbers of sounds of specific frequencies at least at a 3:2 ratio.

In the human brain, auditory sensory memory has been extensively studied using a well-defined component of event-related potential named the mismatch negativity (MMN). The MMN is generated in the auditory and frontal cortices in response to deviant stimuli. In monkeys, cortical N-methyl-d-aspartate (NMDA) receptors have a central role in the generation of the MMN. MMN-like responses have also been recorded in other animals, including rats. The present study aimed at determining whether the MMN-like response in rats depends on an intact NMDA-receptor system. We recorded auditory evoked responses during an oddball paradigm epidurally in anesthetized rats that had received intraperitoneal injections of saline or an NMDA-receptor antagonist MK-801. An MMN-like response was recorded in the oddball paradigm in saline-treated rats. Further, this response was dose-dependently blocked by MK-801. These results suggest that the MMN-like response in rats depends on an intact NMDA-receptor system.

Two-tone pitch-comparison tasks typically comprise several successive pairs of successive tones separated by silent intervals. The serial occurrence of such pairs has been associated with degraded task performance, but the nature of this association is not fully understood. Human adult participants were presented with successive pairs of successive tones. The latter, to-be-compared tone of a pair could differ from the former, to-be-remembered tone of 1046.5 Hz by no more than +/-15 Hz (25 cents). The direction of this difference was easier to identify when it was opposite to that of the preceding pair than when being the same. Merely responding accordingly (irrespectively of whether the response was correct or not) was found not to account for this finding. Our study demonstrates proactive interference in a two-tone pitch comparison task as the difficulty to remember when the first tone of the present pair occurred relative to the last tone of the immediately preceding pair.

The human brain can automatically detect changes even in repeated melodic contours of spectrally varying sounds. However, it is unclear whether this ability is specific to humans. We recorded event-related potentials (ERPs) in urethane-anaesthetized Wistar rats presented with rare pairs of tones ('deviants') interspersed with frequently repeated ones ('standards'). The frequency of the tones varied nonsystematically across their pairs so that deviants stood out from standards only in the melodic ordering (ascending or descending) of the tones of a pair. We found that the absolute amplitude of the ERP was significantly higher to deviants than standards between 106 and 136 ms from the onset of the deviance, suggesting that the ability to automatically detect changes in higher-order invariant attributes that emerge from consecutive sounds is not specific to humans.

It is unclear whether the ability of the brain to discriminate rare from frequently repeated combinations of sound features is limited to the normal sleep/wake cycle. We recorded epidural auditory event-related potentials in urethane-anesthetized rats presented with rare tones ('deviants') interspersed with frequently repeated ones ('standards'). Deviants differed from standards either in frequency alone or in frequency combined with intensity. In both cases, deviants elicited event-related potentials exceeding in amplitude event-related potentials to standards between 76 and 108 ms from the stimulus onset, suggesting the independence of the underlying integrative and memory-based change detection mechanisms of the brain from the normal sleep/wake cycle. The relations of these event-related potentials to mismatch negativity and N1 in humans are addressed.

Nictitating membrane movement and multiple-unit activity in the somatosensory cortex were recorded from rabbits during paired (N=6) and unpaired (N=5) presentations of a tone conditioned stimulus (CS) and an airpuff unconditioned stimulus (US). A behavioural conditioned response (CR) to the CS and an accompanying neural response in the somatosensory cortex developed only in the paired group. Inactivation of the cerebellar interpositus nucleus abolished both the acquired CR and the accompanying neural response. However, the CS facilitated both behavioural and neural responses to the US during the inactivation. Thus, the absence of the CR could not be accounted for by the general inability of the CS to alter the behaviour constituting the CR or the activity of the somatosensory cortex. These findings suggest that the efferent copy of the signal related to the eyeblink CR is projected from the cerebellum to the cerebral cortical areas of the US modality.

Whereas sensory memory in humans has been found to store auditory information for a longer time than visual information, it is unclear whether this is the case also in other species. We recorded hippocampal event-related potentials (ERPs) in awake rabbits exposed to occasional changes in a repeated 50-ms acoustic (1000 versus 2000 Hz) and visual (vertical versus horizontal orientation) stimulus. Three intervals (500, 1500, or 3000 ms) between stimulus repetitions were applied. Whereas acoustic changes significantly affected ERPs with the repetition intervals of 500 and 1500 ms, visual changes did so only with the repetition interval of 500 ms. Our finding, thus, suggests a similarity in sensory processing abilities between human and non-human mammals.

Natural environments are usually composed of multiple sources for sounds. The sounds might physically differ from one another only as feature conjunctions, and several of them might occur repeatedly in the short term. Nevertheless, the detection of rare sounds requires the identification of the repeated ones. Adults have some limited ability to effortlessly identify repeated sounds in such acoustically complex environments, but the developmental onset of this finite ability is unknown. Sleeping newborn infants were presented with a repeated tone carrying six frequent (P = 0.15 each) and six rare (P approximately 0.017 each) conjunctions of its frequency, intensity and duration. Event-related potentials recorded from the infants' scalp were found to shift in amplitude towards positive polarity selectively in response to rare conjunctions. This finding suggests that humans are relatively hard-wired to preattentively identify repeated auditory feature conjunctions even when such conjunctions occur rarely among other similar ones.

Event-related potentials (ERPs) to visual stimuli were recorded from the scalp of eight adult humans performing a task in which they counted vowels from a heard story. In the oddball condition, a repeated (standard) light bar of 50 ms in duration was rarely (P = 0.1) replaced by a (deviant) one differing in orientation from the standard. In the control condition, standards were simply omitted from the series and only (alone-) deviants retained. In both conditions, visual stimuli were asynchronous with auditory-task-relevant stimuli. ERPs to deviants significantly differed in amplitude from those to standards in the midline electrodes centrally, parietally and occipitally at 160-200 ms from stimulus onset. Occipitally, such a difference was absent between ERPs to alone-deviants and those to standards. The occipital differential ERPs to deviants, which thus could be found only when standards were present in the series, are discussed in the context of the mismatch negativity (MMN).

The human brain is known to preattentively trace repeated sounds as holistic entities. It is not clear, however, whether the same holds true if these sounds are rare among other repeated sounds. Adult humans passively listened to a repeated tone with frequent (standard) and rare (deviant) conjunctions of its three features. Six equiprobable variants per conjunction type were assigned from a space built from these features so that the standard variants (P=0.15 each) were not inseparably traceable by means of their linear alignment in this space. Differential scalp-recorded event-related potentials to deviants indicate that the standard variants were traced as repeated wholes despite their preperceptual distinctiveness and resulting rarity among one another.

To process a stimulus as a holistic entity, the human brain must be able to conjoin its different features. Previous evidence suggests that this ability emerges during the first months of life, implying its considerable dependence on postnatal development. We recorded human newborn (1-3 days of age) electrical brain responses to frequently occurring (standard) sounds and to rarely occurring (deviant) sounds in a series. Responses to deviants differed from those to standards despite the fact that only the combination of sound frequency and intensity could be used as a cue for discriminating between these sound types. Our finding suggests that the human brain is ready for auditory feature binding very soon after birth.

Behavioral responses and neural responses in the somatosensory cortex were recorded in nine rabbits during the unpaired and paired treatments of classical eyeblink conditioning with a tone conditioned stimulus (CS) and an airpuff unconditioned stimulus. During the unpaired treatment, neither the behavioral nor neural responses to the CS were observed. During the paired treatment, behavioral conditioned response (CR), accompanied by neural activity, was developed. In well-trained animals occasional failures to elicit the CR were accompanied by an absence of neural responses. Nevertheless, the CS modified the behavioral unconditioned response in paired trials, implying that the CR-failures could not reflect the inability of the CS to modulate the pathways triggering the behavior constituting the CR. Thus, a close link between CR elicitation and somatosensory cortical neural response was established. Our finding suggests that this neural activity to a tone CS during classical eyeblink conditioning reflects an efferent copy of the procedural memory trace.

In eyeblink conditioning in the rabbit (Oryctolagus cuniculus), not only is a conditioned response (CR) acquired, but also the original reflex is modified as a function of training. In Experiment 1, by comparing unconditioned responses in unpaired and paired groups, 3 types of reflex facilitation were distinguished. One type was linked to exposure to the unconditioned stimuli (USs) and/or experimental setting. The 2nd type was related to the formation of the memory trace for conditioned eyeblink. The 3rd type was linked to the conditioned stimulus immediately preceding the US in the paired group. In Experiment 2, reversible inactivation of the interpositus nucleus (IPN) abolished the CR and reduced the CR-related reflex facilitation, indicating that the latter depends on the plasticity of the IPN.

Subjects compared pitches of a standard tone and a comparison tone separated by 1,300-3,000 msec and responded according to whether the comparison tone sounded higher or lower in pitch than the standard tone. Three interfering tones at 300-msec intervals were presented before each pair of tones. Their pitch range varied, being either below or above the pitch of the standard tone; in some of the trials, their pitches were identical to the pitch of the standard tone (no interference). The highest error rate in performance was found when the interfering tones and the comparison tone deviated in the same direction in pitch from the standard tone. In turn, their deviations in the opposite directions resulted in the lowest error rate. This effect was not found to be dependent on whether the interfering tones were randomly ordered or monotonically ordered, together with the standard tone, into melodically ascending/descending sequences. An intermediate error rate in performance was found when the interfering tones and the standard tone were identical. The results support earlier hypotheses, presented in the context of retroactive interference, by demonstrating proactive interference of a tone sequence at the level of representations of individual tones.

Event-related potentials (ERPs) to changes in the visual environment were recorded in rabbits. In the oddball condition, infrequently presented (deviant) stimuli occurred in a series of frequently presented (standard) stimuli. In the deviant-alone condition, standards were omitted. ERPs to oddball-deviants differed from those to standards in all recording sites (cerebellar cortex, visual cortex, dentate gyrus). No corresponding differences were found between ERPs to deviants in the oddball condition and those in the deviant-alone condition. However, because ERPs to deviants in the deviant-alone condition and those to standards did not differ either, ERPs to stimulus changes in the oddball condition seemed to be dependent on the presence of standards, thus representing an analogue to mismatch negativity (MMN) in humans.

Direction and the frequency of spontaneous head movements during the ITIs following forward and backward paired trials were compared to an acquisition of a conditioned orienting (alpha) response directed to the side of the tone source. The head movements were analyzed from video recordings using classification of head turns to preferred and to nonpreferred directions. The results showed a significant increase in the alpha responses during the forward paired conditioning to the preferred direction and rapid extinction during the subsequent backward conditioning sessions. Spontaneous head movements during the ITIs increased to the same preferred direction as the conditioned alpha responses. The results of this experiment suggest that the response initially elicited by the CS can later appear as "spontaneous," instrumental behavior, the form and the nature of which is determined by the characteristics of the conditioned alpha response developing as a result of classical conditioning.