Sequential activation of microcircuits underlying somatosensory-evoked potentials in rat neocortex.

Evoked cortical field potentials are widely used in neurophysiological studies into cortical functioning, but insight in the underlying neural mechanisms is severely hampered by ambiguities in the interpretation of the field potentials. The present study aimed at identifying the precise relationships between the primary evoked cortical field potential (the positive-negative [P1-N1]response) and the temporal and spatial sequence in which different local cortical micro-circuits are recruited. We electrically stimulated the median nerve and recorded field potentials using a 12-channel depth probe in somatosensory cortex of ketamine anesthetized rats. Current source density analysis was used and a grand average was constructed based on all individual animals taking into account individual differences in cortical layering. Manipulation of stimulus strength, selective averaging of single trial responses, and double-pulse stimulation, were used to help disentangle overlapping dipoles and to determine the sequence of neuronal events. We discriminated three phases in the generation of the P1-N1 wave. In the first phase, specific thalamic afferents depolarize both layer III and layer V pyramidal cells. In the second phase, superficial pyramidal cells are depolarized via supragranular intracortical projections. In the third phase, population spikes are generated in layer Vb pyramidal cells, associated with a distinct fast (approximately 1 ms) sink/source configuration. Axon-collaterals of layer Vb pyramidal cells produce an enhanced activation of the supragranular pyramidal cells in layer I-II, which generates N1.

Anticipatory attention to verbal and non-verbal stimuli is reflected in a modality-specific SPN.

The time estimation paradigm allows the recording of anticipatory attention for an upcoming stimulus unconfounded by any anticipatory motor activity. Three seconds after a warning signal (WS) subjects have to press a button. A button press within a time window from 2,850 ms to 3,150 ms after the WS is considered 'correct', a movement prior to 2,850 ms after the WS is labelled 'too early' and a movement after 3,150 ms is labelled 'too late'. Two seconds after the button press a Knowledge of Results (KR) stimulus is presented, informing the subject about the correctness of the response. Stimulus Preceding Negativity (SPN) is a slow wave which is recorded prior to the presentation of the KR stimulus. The SPN has a right hemisphere preponderance and is based upon activity in a network in which prefrontal cortex, the insula Reili and the parietal cortex are crucial. In the present study we asked two questions: (1) does the SPN show modality specificity and (2) does the use of verbal KR stimuli influence the right hemisphere preponderance? Auditory and visual stimuli were presented, in a verbal mode and in a non-verbal mode. SPN amplitudes prior to visual stimuli were larger over the visual cortex than prior to auditory stimuli. SPN amplitudes prior to auditory stimuli were larger over the frontal areas than prior to visual stimuli. The use of verbal stimuli did not influence the right hemisphere preponderance. We concluded that apart from the supramodal effect of KR stimuli in general, there is (first) a modality-specific activation of the relevant sensory cortical areas. The supramodal network underlying the attention for and the use of KR information is activated either from different sensory areas or from language processing cortical areas.

Wait and see.

Anticipatory behavior is aimed at goals that can be reached in the near future. Underlying this behavior are neurophysiological processes, which realize a setting of brain structures involved in the future perception, information processing and action. Anticipatory behavior is accompanied by slow brain potentials, which are generated in the cerebral cortex. They are known as the readiness potential (RP), the contingent negative variation (CNV) and the stimulus preceding negativity (SPN). The RP reflects the timing of a future voluntary movement. The CNV reflects the preparation of a signaled movement and the simultaneous anticipatory attention for the imperative stimulus. The SPN reflects partly the anticipatory attention for the upcoming stimulus. Although these slow potentials are generated in the cortex, the paper shows that a subcortical input from basal ganglia, and in the case of the RP also from the cerebellum, is a necessary condition for their emergence. Slow cortical potentials are the result of concerted activity in a number of cerebral networks, in which the thalamus forms a crucial node. It is suggested that the reticular nucleus of the thalamus plays a pivotal role in anticipatory attention.

Anticipatory attention: an event-related desynchronization approach.

This paper addresses the question of whether anticipatory attention--i.e. attention directed towards an upcoming stimulus in order to facilitate its processing--is realized at the neurophysiological level by a pre-stimulus desynchronization of the sensory cortex corresponding to the modality of the anticipated stimulus, reflecting the opening of a thalamocortical gate in the relevant sensory modality. It is argued that a technique called Event-Related Desynchronization (ERD) of rhythmic 10-Hz activity is well suited to study the thalamocortical processes that are thought to mediate anticipatory attention. In a series of experiments, ERD was computed on EEG and MEG data, recorded while subjects performed a time estimation task and were informed about the quality of their time estimation by stimuli providing Knowledge of Results (KR). The modality of the KR stimuli (auditory, visual, or somatosensory) was manipulated both within and between experiments. The results indicate to varying degrees that preceding the presentation of the KR stimuli, ERD is present over the sensory cortex, which corresponds to the modality of the KR stimulus. The general pattern of results supports the notion that a thalamocortical gating mechanism forms the neurophysiological basis of anticipatory attention. Furthermore, the results support the notion that Event-Related Potential (ERP) and ERD measures reflect fundamentally different neurophysiological processes.

A psychophysiological analysis of inhibitory motor control in the stop-signal paradigm.

We examined two potential inhibitory mechanisms for stopping a motor response. Participants performed a standard visual two-choice task in which visual stop signals and no-go signals were presented on a small proportion of the trials. Psychophysiological measures were taken during task performance to examine the time course of response activation and inhibition. The results were consistent with a horse race model previously proposed to account for data obtained using a stop-signal paradigm. The pattern of psychophysiological responses was similar on stop-signal and no-go trials suggesting that the same mechanism may initiate inhibitory control in both situations. We found a distinct frontal brain wave suggesting that inhibitory motor control is instigated from the frontal cortex. The results are best explained in terms of a single, centrally located inhibition mechanism. Results are discussed in terms of current neurophysiological knowledge.

Event-related desynchronization during anticipatory attention for an upcoming stimulus: a comparative EEG/MEG study.

OBJECTIVES: Our neurophysiological model of anticipatory behaviour (e.g. Acta Psychol 101 (1999) 213; Bastiaansen et al., 1999a) predicts an activation of (primary) sensory cortex during anticipatory attention for an upcoming stimulus. In this paper we attempt to demonstrate this by means of event-related desynchronization (ERD). METHODS: Five subjects performed a time estimation task, and were informed about the quality of their time estimation by either visual or auditory stimuli providing Knowledge of Results (KR). EEG and MEG were recorded in separate sessions, and ERD was computed in the 8-10 and 10-12 Hz frequency bands for both datasets. RESULTS: Both in the EEG and the MEG we found an occipitally maximal ERD preceding the visual KR for all subjects. Preceding the auditory KR, no ERD was present in the EEG, whereas in the MEG we found an ERD over the temporal cortex in two of the 5 subjects. These subjects were also found to have higher levels of absolute power over temporal recording sites in the MEG than the other subjects, which we consider to be an indication of the presence of a 'tau' rhythm (e.g. Neurosci Lett 222 (1997) 111). CONCLUSIONS: It is concluded that the results are in line with the predictions of our neurophysiological model.

Visual feedback about time estimation is related to a right hemisphere activation measured by PET.

In previous EEG experiments we have presented a time estimation task to our subjects, who had to press a button with either the left or right index finger 3 s after an auditory warning stimulus (WS). Two seconds later a visual Knowledge of Results (KR) stimulus was presented on a screen in front, informing them about whether the movement had been made in the correct time window (a vertical line), whether it was too early (a minus sign) or too late (a plus sign). The potential distribution underlying the anticipatory attention for the KR stimulus suggested a right hemisphere network in which the prefrontal cortex, the insula Reili and the parietal cortex were involved. In the present positron emission tomography (PET) activation study we aimed to further localize the exact positions of these regions, using the same paradigm. Two conditions were compared in which the WS had to be followed by a button press with the left index finger. In experimental condition A, subjects received true information about their performance, while in condition B false information was given, utilizing the same stimuli, but randomly, thus without any relation to the actual performance. In both conditions identical stimuli were presented and identical movements were made. Therefore we applied statistical parameter mapping (SPM) for comparison of condition A with B in order to identify regional increases in perfusion related to the anticipation and use of the KR. We found in line with our predictions a right hemisphere activation of (1) BA45, (2) the junction of the posterior insula with the temporal transverse gyrus and (3) the posterior part of the parietal cortex. This activation pattern was accompanied by a better performance due to KR. A second, though not predicted, effect was the increase in correct responses during the last two sessions compared to the first two sessions, independent of KR. This learning effect was accompanied by an activation of BA46 and the supplementary motor area (SMA), again in the right hemisphere. Summarizing, two different prefrontal areas in the right hemisphere were activated: a more ventral area, related to the use of external stimuli providing feedback about a past performance, in order to produce movements in time, and another mid-dorsal one, related to temporal programming on the basis of internal cues.

An ERP correlate of metrical stress in spoken word recognition.

Rhythmic properties of spoken language such as metrical stress, that is, the alternation of strong and weak syllables, are important in speech recognition of stress-timed languages such as Dutch and English. Nineteen subjects listened passively to or discriminated actively between sequences of bisyllabic Dutch words, which started with either a weak or a strong syllable. Weak-initial words, which constitute 12% of the Dutch lexicon, evoked more negativity than strong-initial words in the interval between P2 and N400 components of the auditory event-related potential. This negativity was denoted as N325. The N325 was larger during stress discrimination than during passive listening. N325 was also larger when a weak-initial word followed a sequence of strong-initial words than when it followed words with the same stress pattern. The latter difference was larger for listeners who performed well on stress discrimination. It was concluded that the N325 is probably a manifestation of the extraction of metrical stress from the acoustic signal and its transformation into task requirements.

Neural aspects of anticipatory behavior.

Anticipatory behavior reveals itself in the perceptual domain and in the motor domain. Expectant attention and motor preparation are characterized by selection, aimed at an amelioration of the signal-to-noise ratio in the information to be processed. The functional similarity of anticipatory attention and motor preparation is reflected in the underlying anatomical substrate. The prefrontal cortex, involved in a number of different networks, organizes anticipatory behavior in a top-down way by activating cortico-cortical loops and thalamo-cortical loops to sensory and motor areas. The sensory areas are set to receive the impinging stimulus presentation, the motor areas are set to implement and execute the different motor programs. Thalamic nuclei are also activated from the prefrontal cortex, especially the large association nuclei, the dorsomedial nucleus and the pulvinar. In different models of selective attention the reticular nucleus of the thalamus has a special role in the distribution of the inhibitory control upon the information processing in the "relay" nuclei. It is hypothesized that it has the same pivotal position in motor preparation. Although the anatomical relations do not allow a direct test of the proposed hypothesis, the available psychophysiological evidence does not contradict it.

Event-related desynchronization related to the anticipation of a stimulus providing knowledge of results.

In the present paper, event-related desynchronization (ERD) in the alpha and beta frequency bands is quantified in order to investigate the processes related to the anticipation of a knowledge of results (KR) stimulus. In a time estimation task, 10 subjects were instructed to press a button 4 s after the presentation of an auditory stimulus. Two seconds after the response they received auditory or visual feedback on the timing of their response. Preceding the button press, a centrally maximal ERD is found. Preceding the visual KR stimulus, an ERD is present that has an occipital maximum. Contrary to expectation, preceding the auditory KR stimulus there are no signs of a modality-specific ERD. Results are related to a thalamo-cortical gating model which predicts a correspondence between negative slow potentials and ERD during motor preparation and stimulus anticipation.

Event-related potential measures of information processing during general anesthesia.

To investigate the incidence and manner of auditory information processing during a state of presumed unconsciousness event-related brain potentials (ERPs) were studied in 41 patients undergoing cardiac surgery with propofol/alfentanil anesthesia. The ERPs were recorded during auditory oddball tasks administered before and within several periods of the operation. Mean nasopharyngeal temperature and anesthetic concentrations were determined for each intraoperative ERP recording epoch. During anesthesia ERP waves could still be observed up to 500 ms after stimulus onset indicating that auditory information processing was not suppressed completely by the administered anesthetic agents. Relative to the preoperative recordings, the P1-N1-P2 complex was delayed and more positive going during anesthesia. Comparable changes in ERP morphology have been observed during Stage II-IV sleep, suggesting parallels in the mechanisms underlying early auditory processing in both states of reduced arousal level, possibly related to a selective reduction of a non-specific activity. N1 and P2 peak amplitudes were found to be larger for the deviant tones compared to the standard tones. These amplitude differences most likely reflect automatic detection of stimulus deviance, although it cannot be excluded entirely that they were due to differences in refractoriness. Anesthetic concentrations and nasopharyngeal temperature were found to be of minor significance for ERP control. It is suggested that ERPs could serve as intraoperative reference measures, providing the earliest evidence for auditory processing. This characteristic is important for validation of signals and techniques that are proposed to improve conventional monitoring of anesthesia with respect to detecting unintended awareness.

The crossroads of anticipatory attention and motor preparation. I.

This series of papers suggests that processes underlying anticipatory attention and motor preparation share a common control mechanism. Both functions are of utmost importance for an optimal adaptation to our environment. While the posterior part of the cortex is aimed at the analysis of incoming information, both from the outer world and our own body, the anterior part is involved in action and reaction. If we know when in the near future we will be confronted with relevant information, and how this has to be responded upon, modality specific sensoric and motoric information channels have to be open in order to guarantee an adequate response. The anterior and posterior parts of the cortex are activated from the thalamus and the information transmission is influenced via the reticular nucleus (RN) of the thalamus. The RN itself is under a double control: excitatory from the prefrontal cortex and inhibitory from the neostriatum. It is suggested that selection in anticipatory attention and motor preparation is realized in a comparable way via the RN. In Part I the relevant anatomical structures are described. In Part II the model for anticipatory attention and motor preparation is presented. In Part III the supporting neuropsychological evidence is presented, after which psychophysiological experiments will be discussed having lead to the formulation of the model.

The crossroads of anticipatory attention and motor preparation.

Samenvatting In een drietal artikelen wordt de overeenkomst besproken tussen Processen die ten grondslag liggen aan twee verschillende functies: anticipatoire attentie en motorische preparatie. Beide functies zijn van belang voor een optimale afstemming op onze omgeving. Het achterste gedeelte van de hersenschors houdt zieh voornamelijk bezig met binnenkomende informatie uit de buiten-wereld en het eigen lichaam, terwijl het voorste deel essentieel is voor onze acties en readies. Als bekend is wanneer wij met relevante informatie worden geconfronteerd en vermoed kan worden hoe daarop moet worden gereageerd, worden modali-teitsspeeifieke sensorische en motorische kanalen geopend om zo een snelle en adequate reactie te garanderen. In deze tekst wordt gepostuleerd dat de achterste en voorste cortexhelften op vergelijkbare wijze vanuit de thalamus worden geactiveerd als onder-deel van anticipatoire attentie en motorische preparatie. Cruciaal is de inhiberende invloed die de nucleus reticularis (NR) uitoefent op de onderliggende thalamuskernen. De NR staat onder een dubbele controle: exciterend vanuit de prefrontale cortex en inhiberend vanuit het neostriatum. Aangegeven wordt hoe selectie in de informatieverwerking via deze balans kan worden gerealiseerd, zowel in het sensorische als het motorische domein. Nadat in Deel I een overzicht is gegeven van de relevante anatomische structuren, wordt hier het model voor anticipatoire attentie en motorische preparatie gepresenteerd. In het volgende artikel wordt neuropsychologische evidentie gepresenteerd, waarna het psychofysiologisch onderzoek aan de orde komt dat heeft geleid tot de formule ring van het model.

The differential effects of extremity and movement side on the scalp distribution of the readiness potential (RP) and the stimulus-preceding negativity (SPN).

In a time estimation task subjects had to press a button 3 s after the presentation of a warning stimulus. Two seconds after the movement they were informed about their performance by a knowledge of results (KR) stimulus. Preceding the movement a readiness potential (RP) and prior to the presentation of the KR stimulus a stimulus-preceding negativity (SPN) was recorded. Movement side (left/right) and extremity (hand/foot) were varied within subjects to demonstrate that the RP but not the SPN is affected by such manipulations. The scalp distribution of the late part of the RP was affected by movement side and extremity. Yet it exhibited the expected lateral asymmetry only preceding a movement of the left hand or of the right foot. The scalp distribution of the SPN was not affected by extremity. The size of the right hemisphere preponderance of the SPN depended on movement side following a finger flexion, but not following a plantar flexion of the foot. The experimental design was intended to avoid the temporal overlap between movement-related and stimulus-related activity. Yet it is argued that both results of this experiment can best be explained by such an overlap.

Midlatency auditory evoked potentials as indicators of perceptual processing during general anaesthesia.

We tested the hypothesis that midlatency auditory evoked potentials (MLAEP) can predict the occurrence of long latency AEP components (LLAEP), which are taken as evidence for perceptual processing. Forty-one patients undergoing cardiac surgery were anaesthetized with propofol and alfentanil. During several periods of surgery we recorded LLAEP. Peak-to-peak amplitude measures were used to determine if a particular LLAEP recording trace contained a recognizable waveform. Both before and after each LLAEP recording epoch, MLAEP and the spontaneous electroencephalogram (EEG) were recorded. Peak latencies and amplitudes of brainstem peak V and midlatency peaks Na, Pa, Nb, Pb and Nc, characteristic frequencies from the spontaneous EEG, mean arterial pressure (MAP) and nasopharyngeal temperature (7) were compared between recording epochs with and without clear LLAEP waveforms. These variables were also used in a discriminant analysis to predict the occurrence of an LLAEP waveform. Pa and Nb latencies were significantly shorter both before and after recording epochs in which an LLAEP waveform occurred, compared with epochs in which no LLAEP waveform occurred. Using a combination of up to six EEG, MLAEP, MAP and T measures, it was possible to predict the occurrence or absence of an LLAEP waveform with a sensitivity of 89% and specificity of 86%. We conclude that MLAEP components provide information on the possibility of perceptual processing during general anaesthesia, and thus may be relevant for monitoring depth of anaesthesia.

Anticipatory EMG responses of pericranial muscles in relation to heart rate during a warned simple reaction time task.

Obrist's cardiac-somatic coupling hypothesis predicts a widespread inhibition of heart rate and task-irrelevant muscle activity during expectancy situations. This hypothesis was tested by measuring heart rate and pericranial electromyographic (EMG) activity during a warned simple reaction time task with visual or auditory reaction signals and hand or foot responses. In each of three groups of 24 participants, EMG activity of three different facial, masticatory, or neck muscles was recorded. During the warning interval preceding the presentation of the reaction signal, masticatory and lower facial muscles predominantly showed a gradual inhibition in activity concomitant with heart rate deceleration. In contrast, two upper facial muscles showed increasing activity. Pericranial EMG responses were little affected by reaction signal modality and were independent of responding limb. Greater heart rate deceleration was associated with greater inhibition and weaker facilitation of EMG responses. The results suggest a functional role of inhibitory EMG responses in increasing the perceptual sensitivity to expected signals.

A psychophysiological investigation of the selection and the use of partial stimulus information in response choice.

Two alternative explanations were examined for why selective response activation sometimes starts before stimulus identification is complete (e.g., J. O. Miller & S. A. Hackley, 1992) and sometimes starts only after stimulus identification is complete (e.g., R. De Jong, M. Wierda, G. Mulder, & L. J. M. Mulder, 1988). Distinct psychophysiological methods related to stimulus identification and response selection provided evidence suggesting that partial stimulus information is identified but is or is not used before the stimulus is identified more fully, depending on task requirements. This result (a) suggests strategic adaptation of task performance, (b) is inconsistent with particular discrete and continuous models of information processing, and (c) shows the existence of a central selection mechanism that can prevent the automatic activation of responses associated with preliminary available stimulus information.

Event-related potentials as indirect measures of recognition memory.

Event-related potentials (ERPs) were recorded during an auditory word-recognition task to determine whether they can be used as indirect measures of recognition memory, defined as the ability to differentiate learned from unlearned material when no overt recognition response from the subject is required. A modified version of the two-choice reaction time task developed by Allen, Iacono and Danielson (Allen et al., 1992) was used. In three recognition tasks, administered on two consecutive days, subjects were instructed to indicate recognition of recently learned words. These words were presented along with unlearned words and along with previously learned words which both required a non-recognition response. Recently learned target words as well as previously learned nontarget words elicited a centro-parietal positivity around 500-1000 ms post-stimulus. The size and onset of this late positivity (P300) were affected by the requirement of an overt recognition response. The results suggest that ERPs are sensitive to differences between learned and unlearned words, to some extent independently of the behavioral response. ERPs may therefore be used as indirect measures of recognition memory. In addition, because the present results held for stimuli presented in the auditory modality and because recognition indices were still observed after a one-day interval between learning and testing, this procedure might prove useful in various applications when the integrity of memory is in question.