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Jean-Marc Edeline

Publications and source records attributed to Jean-Marc Edeline.

13 recordsLinked to original sources

Neural representations during sleep: from sensory processing to memory traces.

In the course of a day, the brain undergoes large-scale changes in functional modes, from attentive wakefulness to the deepest stage of sleep. The present paper evaluates how these state changes affect the neural bases of sensory and cognitive representations. Are organized neural representations still maintained during sleep? In other words, despite the absence of conscious awareness, do neuronal signals emitted during sleep contain information and have a functional relevance? Through a critical evaluation of the animal and human literature, neural representations at different levels of integration (from the most elementary sensory level to the most cognitive one) are reviewed. Recordings of neuronal activity in animals at presentation of neutral or significant stimuli show that some analysis of the external word remains possible during sleep, allowing recognition of behaviorally relevant stimuli. Event-related brain potentials in humans confirm the preservation of some sensory integration and discriminative capacity. Behavioral and neuroimaging studies in humans substantiate the notion that memory representations are reactivated and are reorganized during post-learning sleep; these reorganisations may account for the beneficial effects of sleep on behavioral performance. Electrophysiological results showing replay of neuronal sequences in animals are presented, and their relevance as neuronal correlates of memory reactivation is discussed. The reviewed literature provides converging evidence that structured neural representations can be activated during sleep. Which reorganizations unique to sleep benefit memory representations, and to what extent the operations still efficient in processing environmental information during sleep are similar to those underlying the non-conscious, automatic processing continually at work in wakefulness, are challenging questions open to investigation.

Animals↗

Contribution of spike timing to the information transmitted by HVC neurons.

In many species, neurons with highly selective stimulus-response properties characterize higher order sensory areas and/or sensory motor areas of the CNS. In the songbird nuclei, the responses of HVC (used as a proper name) neurons during playback of the bird's own song (BOS) are probably one of the most striking examples of selectivity for natural stimuli. We examined here to what extent spike-timing carries information about natural and time-reversed versions of the BOS. From a heterogenous population of 107 HVC neurons recorded in long-day or short-day conditions, a standard indicator of stimulus preference based on spike-count (the d' index) indicates that a limited proportion of cells can be classified as selective for the BOS (20% with a |d'| > 1). In contrast, quantifying the information conveyed by spike trains with the metric-space of J.D. Victor & K.P Purpura [(1996) J. Neurophysiol., 76, 1310-1326] indicates that 62% of the cells display significant amounts of transmitted information, among which 77% are 'temporal cells'. 'Temporal cells' correspond to cells transmitting significant amounts of information when spike-timing is considered, whereas no information, or lower amounts of transmitted information, is obtained when only spike-count is considered. Computing a correlation index between spike trains [S. Schreiber et al. (2003) Neurocomputing, 52-54,925-931] revealed that spike-timing reliability is higher for the forward than for the reverse BOS, whatever the day length and the cell type are. Cells classified as selective in terms of spike-counts (d' index) had greater amounts of transmitted information, but cells classified as non-selective (d' < 0.5) can also transmit significant amounts of information. Thus, information theory methods demonstrate that a much larger proportion of neurons than expected based on spike-count only participate in the discrimination between stimuli.

Action Potentials↗

The selectivity of canary HVC neurons for the Bird's Own Song: rate coding, temporal coding, or both?

The neuronal selectivity observed in the avian song system for the Bird's Own Song progressively emerged as an extraordinary fruitful model to investigate the neural code underlying the recognition of complex stimuli and the occurrence of learned behaviors. In adult zebra finch, neurons from the HVC (used as a proper name) show very selective auditory responses, firing more to presentation of the Bird's Own Song (BOS) than to reverse BOS or other conspecific songs. However, as adult zebra finches always produce the same stereotyped song, the presence of such highly selective neurons in birds with larger repertoire still remains an open question. Data presented here show that neurons selective for the BOS can be found in adult canary, a seasonal breeding bird which display a large repertoire. More precisely, we found that a large proportion of neurons (29/36) exhibits higher responses to presentation of the forward than to the reverse BOS, and that 22% of the cells were identified as selective on the basis of the d' value. For a cell that was extensively studied, we evaluated to what extent temporal stimulus-related structure predicts the acoustic stimulus using linear or non-linear artificial neural networks (ANN). These analyses indicated that the temporal structure contained in spike trains characterizes more accurately the stimulus than the firing rate. The limitations of applying ANN analyses to electrophysiological data are discussed and potential solutions to increase the confidence in these analysis are proposed.

Action Potentials↗

Selectivity of canary HVC neurons for the bird's own song: modulation by photoperiodic conditions.

To what extent seasonal factors modify the neuronal functional properties within the nuclei of the avian song system remains an open question. In adult songbirds, neurons of the song premotor nucleus HVC (used as a proper name) exhibit selective responses for the bird's own song (BOS). Here we examine whether, outside the breeding season, when songs are less stereotyped, HVC neurons of male canaries still respond selectively to the BOS produced during this period. In an initial experiment, single-unit recordings (n = 114) revealed that the neuronal selectivity for the current BOS was attenuated in males exposed to a short-day photoperiod (typical of the nonbreeding season) compared with that found in males exposed to a long-day photoperiod. In long-day conditions, 35% of the cells responded to the BOS, whereas only 12% did in short-day conditions; there were four times more selective cells (d' > 1) in long-day than in short-day conditions. To determine whether these effects were the consequence of differences in acoustic features between breeding and nonbreeding songs, neurons (n = 72) recorded in short-day conditions were tested with both a short-day BOS and a long-day BOS. A low percentage of neurons exhibited responses to short-day or to long-day BOS (11% for each song). Responses of putative interneurons (spike duration < 0.4 ms) and of putative relay cells were similarly attenuated by the short-day conditions. These results strongly suggest that, in canary, rather than being a fixed property, the selectivity for the BOS moves along a continuum and peaks when the day length mimics the breeding conditions.

Acoustic Stimulation↗

Noradrenergic induction of selective plasticity in the frequency tuning of auditory cortex neurons.

Neuromodulators have long been viewed as permissive factors in experience-induced cortical plasticity, both during development and in adulthood. Experiments performed over the last two decades have reported the potency of acetylcholine to promote changes in functional properties of cortical cells in the auditory, visual, and somatosensory modality. In contrast, very few attempts were made with the monoaminergic systems. The present study evaluates how repeated presentation of brief pulses of noradrenaline (NA) concomitant with presentation of a particular tone frequency changes the frequency tuning curves of auditory cortex neurons determined at 20 dB above threshold. After 100 trials of NA-tone pairing, 28% of the cells (19/67) exhibited selective tuning modifications for the frequency paired with NA. All the selective effects were obtained when the paired frequency was within 1/4 of an octave from the initial best frequency. For these cells, selective decreases were prominent (15/19 cases), and these effects lasted > or =15 min after pairing. No selective effects were observed under various control conditions: tone alone (n = 10 cells), NA alone (n = 11 cells), pairing with ascorbic acid (n = 6 cells), or with GABA (n = 20 cells). Selective effects were observed when the NA-tone pairing was performed in the presence of propranolol (4/10 cells) but not when it was performed in the presence phentolamine (0/13 cells), suggesting that the effects were mediated by alpha receptors. These results indicate that brief increases in noradrenaline concentration can trigger selective modifications in the tuning curves of cortical neurons that, in most of the cases, go in opposite direction compared with those usually reported with acetylcholine.

Acoustic Stimulation↗

Evoked oscillations in unit recordings from the thalamo-cortical auditory system: an aspect of temporal processing or the reflection of hyperpolarized brain states?

Since the beginning of the 90's several lines of research have brought new insights concerning the temporal aspects of sensory processing. Initial observations showing that sensory stimuli can trigger evoked, or induced, oscillations has generated a large number of studies where, explicitly or implicitly, the large-scale rhythmic activity of thalamo-cortical neurons was viewed as a key factor to synchronize neurons responding to different dimensions of a given stimulus, and therefore to solve the so-called "binding problem" (for reviews see Eckhorn 2000, Singer 1990, 1999). This line of research contrasts with the rate coding concept which has been used over the last 40 years to describe the functional properties of neurons in sensory systems and the plasticity of sensory systems both during development and in adulthood. We review here results obtained in the thalamo-cortical auditory system. After a characterization of the oscillatory events, we determined their origins by inactivating each component of the thalamo-cortical loop. Then we compared, within the same animals, the oscillations obtained in anesthetized and unanesthetized conditions. Our data strongly suggest that to clarify the functional roles of evoked oscillations, and more generally of any aspects of temporal coding, we need to study them on awake animals.

Animals↗

The thalamo-cortical auditory receptive fields: regulation by the states of vigilance, learning and the neuromodulatory systems.

The goal of this review is twofold. First, it aims to describe the dynamic regulation that constantly shapes the receptive fields (RFs) and maps in the thalamo-cortical sensory systems of undrugged animals. Second, it aims to discuss several important issues that remain unresolved at the intersection between behavioral neurosciences and sensory physiology. A first section presents the RF modulations observed when an undrugged animal spontaneously shifts from waking to slow-wave sleep or to paradoxical sleep (also called REM sleep). A second section shows that, in contrast with the general changes described in the first section, behavioral training can induce selective effects which favor the stimulus that has acquired significance during learning. A third section reviews the effects triggered by two major neuromodulators of the thalamo-cortical system--acetylcholine and noradrenaline--which are traditionally involved both in the switch of vigilance states and in learning experiences. The conclusion argues that because the receptive fields and maps of an awake animal are continuously modulated from minute to minute, learning-induced sensory plasticity can be viewed as a "crystallization" of the receptive fields and maps in one of the multiple possible states. Studying the interplays between neuromodulators can help understanding the neurobiological foundations of this dynamic regulation.

Animals↗

Bursts in the medial geniculate body: a comparison between anesthetized and unanesthetized states in guinea pig.

Thalamic high frequency bursts have long been described under anesthesia and during slow-wave sleep (SWS). More recently, studies in the lateral geniculate nucleus have pointed out that they are also present during waking (W). Here, we compared the bursts recorded in the medial geniculate body of guinea pigs under anesthesia or during periods of W and SWS. The tuning of single units was tested between threshold and 80 dB SPL in two conditions: (1) in restrained, undrugged, non-sleep-deprived guinea pigs (n=101 cells) and (2) under pentobarbital anesthesia (n=53 cells). Off-line analyses allowed us to distinguish single action potentials (APs) from bursts. A burst was defined as a group of APs with an interspike interval < or =4 ms, preceded by a silent period > or =100 ms. We found that auditory thalamus bursts occur in synchronized electroencephalogram states (SWS and anesthesia), but also during W. Although the burst characteristics did not differ among the three states, group data showed that the proportion of bursts within spike trains was the greatest under anesthesia. This observation resulted from two types of effects: (1) the percentage of non-bursting cells was lowest under anesthesia and (2) some cells under anesthesia exhibited up to 90% of bursts, whereas during W or SWS the highest proportion of bursts did not exceed 40%. The presence of these bursts is discussed with regards to the known fluctuations of membrane potential which occur in these various states.

Acoustic Stimulation↗

Does the radial arm maze necessarily test spatial memory?

Since its design 25 years ago (Olton & Samuelson, 1976), the eight-arm radial maze has become very popular and is now widely used to assess spatial memory in rodents. Two versions of the full-baited maze protocol are present in the literature: with or without confinement between the visit of each arm. The confinement was introduced by Olton himself as early as 1977 (Olton, Collison, & Werz, 1977) to eliminate stereotypic behaviors that he had previously observed. It is widely regarded that the confinement prevents rodents from developing these response patterns, and as such it is considered an improved procedure to test spatial memory. Surprisingly, to the best of our knowledge, no study has been especially designed to demonstrate the efficacy of the confinement in blocking the stereotypic behaviors of the animals. The present study compares the strategies of rats trained with or without a confinement procedure. The results show that, after nine days of training, rats submitted to a 5- or a 10-s confinement reach the same level of performance as rats without confinement. The confinement totally prevents stereotypic behaviors like clockwise serial searching strategies which are often observed without confinement. Even a 0-s confinement is sufficient to prevent clockwise strategies, but rats seem to develop other stratagems which do not imply spatial memory. Furthermore, rats previously trained without confinement are unable to perform the task when confinement is introduced on a test day. In contrast, rats previously trained with confinement perform the task correctly when the confinement is no longer present. Thus, without confinement, good levels of performance can be achieved without precise spatial representations.

Animals↗

Evoked oscillations in the thalamo-cortical auditory system are present in anesthetized but not in unanesthetized rats.

Over the last decade, a large number of studies have characterized stimulus-evoked oscillations in the visual cortex of anesthetized and unanesthetized animals. Comparatively, only a few studies have been performed in auditory cortex. This study compared the tone-evoked oscillations detected from the same recording sites in the thalamo-cortical auditory system of unanesthetized and anesthetized rats. Simultaneous multiunit recordings were collected in auditory cortex, auditory thalamus, and the auditory sector of the reticular nucleus of restrained rats, which spontaneously shifted from waking (W) to slow-wave sleep (SWS) and paradoxical sleep (PS). Subsequently, the same recording sites were tested under pentobarbital anesthesia, then under high doses of diazepam, and finally under urethan anesthesia. Under these drugs, oscillations were detected in 54% of the recordings: one-half of them were stimulus-locked oscillations and were directly observed on peri-stimulus time histograms (PSTHs); one-half of them were non-stimulus-locked oscillations and were detected on autocorrelograms. Spontaneous oscillations were present for 17% of the recordings. During SWS, only non-stimulus-locked oscillations were observed for a small percentage of recordings (12%). This percentage did not differ significantly from the one of spontaneous oscillations obtained during SWS (8%). No oscillations were found in W and PS. Both under anesthesia and in SWS, the frequency range of the oscillations was 5-15 Hz, and there was no frequency difference between evoked and spontaneous oscillations. Although surprising, the absence of oscillations in awake animals may allow each neuron to process acoustic information independently of its neighbors and may in fact benefit auditory perception.

Acoustic Stimulation↗

Sex and season influence the proportion of thin spike cells in the canary HVc.

In songbirds, anatomical attributes of song nuclei exhibit sexual and seasonal differences. To extend these data to physiological correlates, neurons ( n= 374) were recorded in the HVc of male and female canaries during and outside the breeding period. Surprisingly, a particular type of action potential waveforms was observed more frequently in breeding than in non-breeding birds and in males than in females. These neurons showed both a shorter action potential duration (< 0.4 ms) and a higher firing rate (2.5 1.4 spikes/s) than the other neurons. Such characteristics are usually associated with interneurons in the songbird HVc as well as in the mammalian neocortex. Thus, these results provide the first electrophysiological evidence for an alteration of the neuronal network of HVc across sexes and seasons.

Action Potentials↗

Does head-only exposure to GSM-900 electromagnetic fields affect the performance of rats in spatial learning tasks?

The rapid expansion of mobile communication has generated intense interest, but has also fuelled ongoing concerns. In both humans and animals, radiofrequency radiations are suspected to affect cognitive functions. More specifically, several studies performed in rodents have suggested that spatial learning can be impaired by electromagnetic field exposure. However, none of these previous studies have simulated the common conditions of GSM mobile phones use. This study is the first using a head-only exposure system emitting a 900-MHz GSM electromagnetic field (pulsed at 217 Hz). The two behavioural tasks that were evaluated here have been used previously to demonstrate performance deficits in spatial learning after electromagnetic field exposure: a classical radial maze elimination task and a spatial navigation task in an open-field arena (dry-land version of the Morris water maze). The performances of rats exposed for 45 min to a 900-MHz electromagnetic field (1 and 3.5 W/kg) were compared to those of sham-exposed and cage-control rats. There were no differences among exposed, sham, and cage-control rats in the two spatial learning tasks. The discussion focuses on the potential reasons that led previous studies to conclude that learning deficits do occur after electromagnetic field exposure.

Animals↗

Muscimol diffusion after intracerebral microinjections: a reevaluation based on electrophysiological and autoradiographic quantifications.

Intracerebral muscimol injection is widely used to inactivate discrete brain structures during behavioral tasks. However, little effort has been made to quantify the extent of muscimol diffusion. The authors report here electrophysiological and autoradiographic results obtained after muscimol injection (1 microg/microl) either into the nucleus basalis magnocellularis (0.1-0.4 microl) or into the thalamic reticular nucleus (RE, 0.05-0.1 microl). In 52 rats, multiunit recordings were collected either in the RE or in the auditory thalamus during the 2 h following muscimol injection. Decreases in neuronal activity were observed up to 3 mm from the injection site; their time of occurrence was a function of the distance between the injection and recording sites. Because these decreases cannot be explained by physiological effects, they likely reflected muscimol diffusion up to the recording sites. Autoradiographic studies involved 25 rats and different experimental conditions. Optical density (OD) measures indicated that after a survival time of 15 min, a 0.05 microl injection produced a labeled area of 5.25 mm(2) at the injection site and a rostrocaudal labeling of 1.7 mm. Increasing the survival time to 60 min, or increasing the injected volume to 0.1 microl, systematically led to a larger labeled area at the injection site (8-12 mm(2)) and to a larger rostrocaudal diffusion (2.0-2.5 mm). Direct quantifications of radioactivity by a high-resolution radioimager validated the OD measures and even indicated a larger muscimol diffusion (up to 3.25 mm). Thus, these data point out that muscimol diffusion after intracerebral microinjection is larger than usually supposed. The relationships between these results and those obtained in behavioral studies are discussed.

Animals↗