Search

Rhythmic brain activity has been implicated in many brain functions and it is sensible to neuromodulation, but so far very few studies have investigated this activity on the cellular levelin vitroin human tissue samples. In this study we revealed and characterized a novel rhythmic network activity in human neocortex. Intracellular patch-clamp recordings showed that giant depolarizing potentials (GDPs) were frequently found in human cortical neurons. GDPs appeared in a low frequency band (∼ 0.3 Hz) similar to that described for slow oscillationsin vivoand displayed large amplitudes and long decay times. Under the same experimental conditions, no rhythmic activity was found in L2/3 of the rat neocortex. GDPs were predominantly observed in a subset of L2/3 interneurons considered to be large basket cells based on previously described morphological features. In addition, GDPs are highly sensitive to norepinephrine (NE) and acetylcholine (ACh), two neuromodulators known to modulate low frequency oscillations. NE increased the frequency of the GDPs by enhancing β-adrenergic receptor activity while ACh decreased GDP frequency through M4muscarinic receptor-activation. Multi-electrode array (MEA) recordings demonstrated that NE promoted synchronous oscillatory network activity while the application of ACh led to a desynchronization of neuronal activity. Our data indicate that the human neocortex is more prone to generate slow wave activity, which was reflected by more pronounced GDPs in L2/3 large basket cells. The distinct modulation of GDPs and slow wave activity by NE and ACh exerts a specific modulatory control over the human neocortex.

Cerebral cortex 32(10), 2095-2111 (2022). doi:10.1093/cercor/bhab340, Published by Oxford Univ. Press, Oxford

The neuromodulator acetylcholine (ACh) plays an important role in arousal, attention, vigilance, learning and memory. ACh is released during different behavioural states and affects the brain microcircuit by regulating neuronal and synaptic properties. Here, we investigated how a low concentration of ACh (30 μM) affects the intrinsic properties of electrophysiologically and morphologically identified excitatory and inhibitory neurons in layer 4 (L4) of rat barrel cortex. ACh altered the membrane potential of L4 neurons in a heterogeneous manner. Nearly all L4 regular spiking (RS) excitatory neurons responded to bath-application of ACh with a M4 muscarinic ACh receptor-mediated hyperpolarisation. In contrast, in the majority of L4 fast spiking (FS) and non-fast spiking (nFS) interneurons 30 μM ACh induced a depolarisation while the remainder showed a hyperpolarisation or no response. The ACh-induced depolarisation of L4 FS interneurons was much weaker than that in L4 nFS interneurons. There was no clear difference in the response to ACh for three morphological subtypes of L4 FS interneurons. However, in four morpho-electrophysiological subtypes of L4 nFS interneurons, VIP+-like interneurons showed the strongest ACh-induced depolarisation; occasionally, even action potential firing was elicited. The ACh-induced depolarisation in L4 FS interneurons was exclusively mediated by M1 muscarinic ACh receptors; in L4 nFS interneurons it was mainly mediated by M1 and/or M3/5 muscarinic ACh receptors. In a subset of L4 nFS interneurons, a co-operative activation of muscarinic and nicotinic ACh receptors was also observed. The present study demonstrates that low-concentrations of ACh affect different L4 neuron types in a cell-type specific way. These effects result from a specific expression of different muscarinic and/or nicotinic ACh receptors on the somatodendritic compartments of L4 neurons. This suggests that even at low concentrations ACh may tune the excitability of L4 excitatory and inhibitory neurons and their synaptic microcircuits differentially depending on the behavioural state during which ACh is released.

Neocortical layer 6 plays a crucial role in sensorimotor coordination and integration through functionally segregated circuits linking intracortical and subcortical areas. However, because of the high neuronal heterogeneity and sparse intralaminar connectivity data on the cell-type specific synaptic microcircuits in layer 6 remain few and far between. To address this issue, whole-cell recordings combined with morphological reconstructions have been used to identify morphoelectric types of layer 6A pyramidal cells (PCs) in rat barrel cortex. Cortico-thalamic (CT), corticocortical (CC) and cortico-claustral (CCla) pyramidal cells have been distinguished based on to their distinct dendritic and axonal morphologies as well as their different electrophysiological properties. Here we demonstrate that these three types of layer 6A pyramidal cells innervate neighboring excitatory neurons with distinct synaptic properties: CT PCs establish weak facilitating synapses to other L6A PCs; CC PCs form synapses of moderate efficacy; while synapses made by putative CCla PCs display the highest release probability and a marked short-term depression. Furthermore, for excitatory-inhibitory synaptic connections in layer 6 we were able to show that both the presynaptic PC type and the postsynaptic interneuron type govern the dynamic properties of the of the respective synaptic connections. We have identified a functional division of local layer 6A excitatory microcircuits which may be responsible of the differential temporal engagement of layer 6 feed-forward and feedback networks. Our results provides a basis for further investigations on the long-range cortico-cortical, cortico-thalamic and cortico-claustral pathways.

Neuroscience : an international journal under the editorial direction of IBRO 456, 114-130 (2021). doi:10.1016/j.neuroscience.2020.06.006 special issue: "Metabotropic Regulation of Synaptic Plasticity / Edited by David Fernández de Sevilla, Ángel Nuñez, Alfonso Araque, Washington Buño", Published by Elsevier Science, Amsterdam [u.a.]

Acetylcholine (ACh) is known to regulate cortical activity during different behavioral states, e.g. wakefulness and attention. Here we show a differential expression of muscarinic ACh receptors (mAChRs) and nicotinic AChRs (nAChRs) in different layer 6A (L6A) pyramidal cell (PC) types of somatosensory cortex. At low concentrations, ACh induced a persistent hyperpolarization in corticocortical (CC) but a depolarization in corticothalamic (CT) L6A PCs via M4 and M1 mAChRs, respectively. At ∼1 mM ACh depolarized exclusively CT PCs via α4β2 subunit-containing nAChRs without affecting CC PCs. Miniature EPSC frequency in CC PCs was decreased by ACh but increased in CT PCs. In synaptic connections with a presynaptic CC PC, glutamate release was suppressed via M4 mAChR activation but enhanced by nAChRs via α4β2 nAChRs when the presynaptic neuron was a CT PC. Thus, in layer 6A the interaction of mAChRs and nAChRs results in an altered excitability and synaptic release, effectively strengthening corticothalamic output while weakening corticocortical synaptic signaling.

Adenosine is considered to be a key regulator of sleep homeostasis by promoting slow-wave sleep through inhibition of the brain's arousal centers. However, little is known about the effect of adenosine on neuronal network activity at the cellular level in the neocortex. Here, we show that adenosine differentially modulates synaptic transmission between different types of neurons in cortical layer 4 (L4) through activation of pre- and/or postsynaptically located adenosine A1 receptors. In recurrent excitatory connections between L4 spiny neurons, adenosine suppresses synaptic transmission through activation of both pre- and postsynaptic A1 receptors. In reciprocal excitatory and inhibitory connections between L4 spiny neurons and interneurons, adenosine strongly suppresses excitatory transmission via activating presynaptic A1 receptors but only slightly suppresses inhibitory transmission via activating postsynaptic A1 receptors. Adenosine has no effect on inhibitory transmission between L4 interneurons. The effect of adenosine is concentration dependent and first visible at a concentration of 1 μM. The effect of adenosine is blocked by the specific A1 receptor antagonist, 8-cyclopentyltheophylline or the nonspecific adenosine receptor antagonist, caffeine. By differentially affecting excitatory and inhibitory synaptic transmission, adenosine changes the excitation-inhibition balance and causes an overall shift to lower excitability in L4 primary somatosensory (barrel) cortical microcircuits.

Recent years have seen substantial progress in studying the structural and functional properties of GABAergic interneurons and their roles in the neuronal networks of barrel cortex. Although GABAergic interneurons represent only about 12% of the total number of neocortical neurons, they are extremely diverse with respect to their structural and functional properties. It has become clear that barrel cortex interneurons not only serve the maintenance of an appropriate excitation/inhibition balance but also are directly involved in sensory processing. In this review we present different interneuron types and their axonal projection pattern framework in the context of the laminar and columnar organization of the barrel cortex. The main focus is here on the most prominent interneuron types, i.e. basket cells, chandelier cells, Martinotti cells, bipolar/bitufted cells and neurogliaform cells, but interneurons with more unusual axonal domains will also be mentioned. We describe their developmental origin, their classification with respect to molecular, morphological and intrinsic membrane and synaptic properties. Most importantly, we will highlight the most prominent circuit motifs these interneurons are involved in and in which way they serve feed-forward inhibition, feedback inhibition and disinhibition. Finally, this will be put into context to their functional roles in sensory signal perception and processing in the whisker system and beyond.

Excitatory layer 4 (L4) neurons in the ‘barrel field’ of the rat somatosensory cortex represent an important component in thalamocortical information processing. However, no detailed information exists concerning the quantitative geometry of synaptic boutons terminating on these neurons. Thus, L4 synaptic boutons were investigated using serial ultrathin sections and subsequent quantitative 3D reconstructions. In particular, parameters representing structural correlates of synaptic transmission and plasticity such as the number, size and distribution of pre- and postsynaptic densities forming the active zone (AZ) and of the three functionally defined pools of synaptic vesicles were analyzed. L4 synaptic boutons varied substantially in shape and size; the majority had a single, but large AZ with opposing pre- and postsynaptic densities that matched perfectly in size and position. More than a third of the examined boutons showed perforations of the postsynaptic density. Synaptic boutons contained on average a total pool of 561 ± 108 vesicles, with ~5 % constituting the putative readily releasable, ~23 % the recycling, and the remainder the reserve pool. These pools are comparably larger than other characterized central synapses. Synaptic complexes were surrounded by a dense network of fine astrocytic processes that reached as far as the synaptic cleft, thus regulating the temporal and spatial glutamate concentration, and thereby shaping the unitary EPSP amplitude. In summary, the geometry and size of AZs, the comparably large readily releasable and recycling pools, together with the tight astrocytic ensheathment, may explain and contribute to the high release probability, efficacy and modulation of synaptic transmission at excitatory L4 synaptic boutons. Moreover, the structural variability as indicated by the geometry of L4 synaptic boutons, the presence of mitochondria and the size and shape of the AZs strongly suggest that synaptic reliability, strength and plasticity is governed and modulated individually at excitatory L4 synaptic boutons.

Cerebral cortex 25(3), 772-787 (2015). doi:10.1093/cercor/bht274, Published by Oxford Univ. Press, Oxford

We dissect Rossler equation and give a linear core, which can capture the essence of Rossler oscillator. The maximal transversal Lyapunov exponents calculated from the core are in good agreement with those obtained from Rossler equation. The idea is also tested for coupled Chua's circuit systems. Numerical experiments are given.

Synchronization stability in complex networks is a topic of theoretical interest and practical importance. Increasing effort has been devoted to the enhancement of synchronizability of networks, or more specifically, the design of synchronizable networks. However, most previous attempts turn the coupling weight/gradient or change the topological interactions, which sometimes is not manageable. In this paper, by adopting a simple kind of discontinuous coupling strategy: the uniform on-off coupling scheme, with on-off period being comparable to the timescale of node dynamics, the problem is solved within the framework of the master stability function. The results show that, this strategy can greatly increase the stable region of synchronization, which means the size of synchronizable networks can be much larger than the traditional case, without any changes of their connections. Furthermore, the synchronization speed can be accelerated considerably, which is even higher than the previous optimal case. The mechanism of the facilitation is revealed and shows that the continuous coupling in fact is one of the worst choices for synchronization in the view of discontinuous coupling strategy. The coupling cost required for synchronization is also examined, which is approximately the same as the continuous coupling.

The fate of the subplate (SP) is still a matter of debate. The SP and layer 6 (which is ontogenetically the oldest and innermost neocortical lamina) develop coincidentally. Yet, the function of sublamina 6B is largely unknown. It has been suggested that it consists partly of neurons from the transient SP, however, experimental evidence for this hypothesis is still missing. To obtain first insights into the neuronal complement of layer 6B in the somatosensory rat barrel cortex, we used biocytin stainings of SP neurons (aged 0-4 postnatal days, PND) and layer 6B neurons (PND 11-35) obtained during in vitro whole-cell patch-clamp recordings. Neurons were reconstructed for a quantitative characterization of their axonal and dendritic morphology. An unsupervised cluster analysis revealed that the SP and layer 6B consist of heterogeneous but comparable neuronal cell populations. Both contain 5 distinct spine-bearing cell types whose relative fractions change with increasing age. Pyramidal cells were more prominent in layer 6B, whereas non-pyramidal neurons were less frequent. Because of the high morphological similarity of SP and layer 6B neurons, we suggest that layer 6B consists of persistent non-pyramidal neurons from the SP and cortical L6B pyramidal neurons.

The synchronous dynamics of complex networks of pulse-coupled bursting neurons with heterogeneous coupling is studied. The results reveal that the generation and the stability of the completely synchronous state in such networks only depends on the input weight of each neuron, which is independent of the network structure. If the input weight is heterogenous in the networks, then a subgroup of bursting neurons, each having the same input weight, can be in almost completely synchronous states, and different subgroups with different input weights can be in burst synchronous states. The stability of this input weight determined synchronization is numerically studied by Lyapunov exponents, and the results show that the complete or burst synchronization in any complex neuronal networks is fully determined by the input weight of each neuron.

We study the dynamical localization in the system of a two-level Rydberg atom interacting exactly with a pulsed standing microwave. This system approaches an atom optics realization of usual delta-kicked rotor under the rotating-wave approximation (RWA). We find that the non-energy-conserving processes neglected in the RWA have a strong effect on the classical and quantum diffusion, which, compared with the results of the RWA, can enhance, reduce and even completely suppress the momentum diffusion under certain kicked conditions.

We study the dynamics of unidirectionally coupled linear arrays of chaotic oscillators in star configuration, and find that if the central driving oscillator is different from the oscillators come off from it in the linear arrays, then the oscillators in different linear arrays with the same position with respect to the central one are synchronized but no chaotic synchronization occurs in any one linear array. Such a synchronization, called layered synchronization, severely depends on the dynamical coupling symmetry. Numerical calculation shows that, for the Lorenz oscillator, the strange attractor spiralling around two focus points will become a strange attractor spiralling around one focus point down the linear arrays. The stabilities of the synchronized states are enhanced by chaos, and degraded when the oscillators are far from the central driving oscillator.

Excitatory connections between neocortical layer 4 (L4) and L6 are part of the corticothalamic feedback microcircuitry. Here we studied the intracortical element of this feedback loop, the L4 spiny neuron-to-L6 pyramidal cell connection. We found that the distribution of synapses onto both putative corticothalamic (CT) and corticocortical (CC) L6 pyramidal cells (PCs) depends on the presynaptic L4 neuron type but is independent of the postsynaptic L6 PC type. L4 spiny stellate cells establish synapses on distal apical tuft dendrites of L6 PCs and elicit slow unitary excitatory postsynaptic potentials (uEPSPs) in L6 somata. In contrast, the majority of L4 star pyramidal neurons target basal and proximal apical oblique dendrites of L6 PCs and show fast uEPSPs. Compartmental modeling suggests that the slow uEPSP time course is primarily the result of dendritic filtering. This suggests that the dendritic target specificity of the 2 L4 spiny neuron types is due to their different axonal projection patterns across cortical layers. The preferential dendritic targeting by different L4 neuron types may facilitate the generation of dendritic Ca(2+) or Na(+) action potentials in L6 PCs; this could play a role in synaptic gain modulation in the corticothalamic pathway.

The combination of patch clamp recordings from two (or more) synaptically coupled neurons (paired recordings) in acute brain slice preparations with simultaneous intracellular biocytin filling allows a correlated analysis of their structural and functional properties. With this method it is possible to identify and characterize both pre- and postsynaptic neurons by their morphology and electrophysiological response pattern. Paired recordings allow studying the connectivity patterns between these neurons as well as the properties of both chemical and electrical synaptic transmission. Here, we give a step-by-step description of the procedures required to obtain reliable paired recordings together with an optimal recovery of the neuron morphology. We will describe how pairs of neurons connected via chemical synapses or gap junctions are identified in brain slice preparations. We will outline how neurons are reconstructed to obtain their 3D morphology of the dendritic and axonal domain and how synaptic contacts are identified and localized. We will also discuss the caveats and limitations of the paired recording technique, in particular those associated with dendritic and axonal truncations during the preparation of brain slices because these strongly affect connectivity estimates. However, because of the versatility of the paired recording approach it will remain a valuable tool in characterizing different aspects of synaptic transmission at identified neuronal microcircuits in the brain.

An elementary feature of sensory cortices is thought to be their organisation into functional signal-processing units called ‘cortical columns’. These elementary units process sensory information arriving from peripheral receptors; they are vertically oriented throughout all cortical layers and contain several thousands of excitatory and inhibitory synaptic connections. To understand how sensory signals are transformed into electrical activity in the neocortex it is necessary to elucidate the spatial-temporal dynamics of cortical signal processing and the underlying neurons and synaptic ‘microcircuits’.

We investigate the coevolution dynamics of node activities and coupling strengths in coupled chaotic oscillators via a simple threshold adaptive scheme. The coupling strength is synchronous activity regulated, which in turn is able to boost the synchronization remarkably. In the case of weak coupling, the globally coupled oscillators present a highly clustered functional connectivity with a power-law distribution in the tail with $\ensuremath{\gamma}\ensuremath{\simeq}3.1$, while for strong coupling, they self-organize into a network with a heterogeneously rich connectivity at the onset of synchronization but exhibit rather sparse structure to maintain the synchronization in noisy environment. The relevance of the results is briefly discussed.

An analytical expression for the synchronization time in coupled-map networks is given. By means of the expression, the synchronization time for any given network can be predicted accurately. Furthermore, for networks in which the distributions of nontrivial eigenvalues of coupling matrices have some unique characteristics, analytical results for the minimal synchronization time are given.

We study the dynamic stabilities of unidirectionally coupled linear arrays of chaotic oscillators with time-delay feedbacks in star configuration, and find that if all oscillators in the network are identical, then the oscillators in the linear arrays can anticipate the driving oscillators, and simultaneously the oscillators in the linear arrays with the same position with respect to the central one are in synchronous chaotic state. Compared with the anticipated synchronization, the layered synchronization is first generated and last destroyed as the coupling constant is increased. This coexistence of anticipated and layered chaotic synchronization is destroyed by long time feedback. If the driving and driven oscillators are different, then only layered chaotic synchronization is possible.

Stable propagation of noise-induced synchronous spiking in uncoupled linear neuron arrays is studied numerically. The chaotic neurons in the unidirectionally coupled linear array are modeled by Hindmarsh-Rose neurons. Stability analysis shows that the synchronous chaotic spiking can be successfully transmitted to cortical areas through layered synchronization in the neural network under certain conditions of the network structure.

We study the time-delay and unidirectionally coupled ring and linear arrays of chaotic systems, and find that under certain conditions, the linear array can spatial periodically ``copy'' the chaotic dynamics of the ring with very long anticipation times. Numerical calculations of the Lyapunov exponents show that the delay times can result in unsynchronized chaotic waves, periodic waves, and stable states in the ring that are replicated in the linear array, but have no effect on the absolute stability of the anticipatory synchronization. Our results show that such configurations can both enhance the absolute stability of the synchronization manifolds and minimize the effects of convective instabilities.

The dynamics of brain functional motifs are studied. It is shown that different rhythms can occur in the motifs when time delay is taken into account. These rhythms include synchronization, winnerless competition (WLC) and two plus one (TPO). The main discovery is that the transition from WLC to synchronization can be induced simply by time delay. It is also concluded that some medium time delay is needed to achieve WLC in the realistic case. The motifs composed of heterogeneous neurons are also considered.

We study the fast synchronization in complex networks of coupled Hindmarsh-Rose neurons. The relation between the maximal Lyapunov exponent corresponding to the least stable transverse mode and the speed of synchronization is given, based on which we can obtain an optimal value of global coupling strength, with which the network synchronizes with the minimal synchronization time. The speed limits of several kinds of complex networks (small-world, scale-free, and modular) with different eigenratios are studied. Finally, we extend to the modified Hodgkin-Huxley neuron case.