ECoG activity distribution patterns detects global cortical responses following weak tactile inputs.

Many studies have suggested that the neocortex operates as a global network of functionally interconnected neurons, indicating that any sensory input could shift activity distributions across the whole brain. A tool assessing the activity distribution across cortical regions with high temporal resolution could then potentially detect subtle changes that may pass unnoticed in regionalized analyses. We used eight-channel, distributed electrocorticogram (ECoG) recordings to analyze changes in global activity distribution caused by single pulse electrical stimulations of the paw. We analyzed the temporally evolving patterns of the activity distributions using principal component analysis (PCA). We found that the localized tactile stimulation caused clearly measurable changes in global ECoG activity distribution. These changes in signal activity distribution patterns were detectable across a small number of ECoG channels, even when excluding the somatosensory cortex, suggesting that the method has high sensitivity, potentially making it applicable to human electroencephalography (EEG) for detection of pathological changes.

Diversified physiological sensory input connectivity questions the existence of distinct classes of spinal interneurons.

The spinal cord is engaged in all forms of motor performance but its functions are far from understood. Because network connectivity defines function, we explored the connectivity of muscular, tendon, and tactile sensory inputs among a wide population of spinal interneurons in the lower cervical segments. Using low noise intracellular whole cell recordings in the decerebrated, non-anesthetized cat in vivo, we could define mono-, di-, and trisynaptic inputs as well as the weights of each input. Whereas each neuron had a highly specific input, and each indirect input could moreover be explained by inputs in other recorded neurons, we unexpectedly also found the input connectivity of the spinal interneuron population to form a continuum. Our data hence contrasts with the currently widespread notion of distinct classes of interneurons. We argue that this suggested diversified physiological connectivity, which likely requires a major component of circuitry learning, implies a more flexible functionality.

Metrics for environmental compensation: A comparative analysis of Swedish municipalities.

Environmental compensation (EC) aims at addressing environmental losses due to development projects and involves a need to compare development losses with compensation gains using relevant metrics. A conceptual procedure for computing no net loss is formulated and used as a point of departure for a comparative analysis of metrics used by five Swedish municipalities as a part of their EC implementation in the spatial planning context of detailed development plans. While Swedish law does not require EC in this context, these municipalities have still decided to introduce EC requirements for development projects that occur on municipality-owned land and to promote voluntary EC among private actors in development projects on private land. There is substantial variation across the municipalities studied with respect to both metrics and attributes subject to measurement, but there are also similarities: The attributes considered when assessing the need for EC in conjunction with development are not only about nature per se, but also about recreational opportunities and other types ecosystem services; semi-quantitative metrics such as scores are common while quantitative or monetary metrics are rare; and metrics are rarely applied to assess compensatory gains, focusing instead on losses from development. Streamlining across municipalities might be warranted for increasing predictability and transparency for developers and citizens, but it also introduces considerable challenges such as a need for developing consistent guidelines for semi-quantitative metrics, and to handle substitutability issues if metrics are not only applied on individual attributes but also on groups of attributes. The broad scope of attributes used by the municipalities is in line with an international tendency to broaden EC to include not only biodiversity aspects but also ecosystem services. Moreover, the EC systems applied by the municipalities are of particular importance for highlighting the crucial role of environmental management for maintaining and enhancing biodiversity and ecosystem services not only in areas having formal protection status but also in the everyday landscape. The municipalities' experience and strengths and weaknesses associated with their EC systems are therefore relevant also in an international perspective.

Focal neocortical lesions impair distant neuronal information processing.

KEY POINTS: Parts of the fields of neuroscience and neurology consider the neocortex to be a functionally parcelled structure. Viewed through such a conceptual filter, there are multiple clinical observations after localized stroke lesions that seem paradoxical. We tested the effect that localized stroke-like lesions have on neuronal information processing in a part of the neocortex that is distant to the lesion using animal experiments. We find that the distant lesion degrades the quality of neuronal information processing of tactile input patterns in primary somatosensory cortex. The findings suggest that even the processing of primary sensory information depends on an intact neocortical network, with the implication that all neocortical processing may rely on widespread interactions across large parts of the cortex. ABSTRACT: Recent clinical studies report a surprisingly weak relationship between the location of cortical brain lesions and the resulting functional deficits. From a neuroscience point of view, such findings raise questions as to what extent functional localization applies in the neocortex and to what extent the functions of different regions depend on the integrity of others. Here we provide an in-depth analysis of the changes in the function of the neocortical neuronal networks after distant focal stroke-like lesions in the anaesthetized rat. Using a recently introduced high resolution analysis of neuronal information processing, consisting of pre-set spatiotemporal patterns of tactile afferent activation against which the neuronal decoding performance can be quantified, we found that stroke-like lesions in distant parts of the cortex significantly degraded the decoding performance of individual neocortical neurons in the primary somatosensory cortex (decoding performance decreased from 30.9% to 24.2% for n = 22 neurons, Wilcoxon signed rank test, P = 0.028). This degrading effect was not due to changes in the firing frequency of the neuron (Wilcoxon signed rank test, P = 0.499) and was stronger the higher the decoding performance of the neuron, indicating a specific impact on the information processing capacity in the cortex. These findings suggest that even primary sensory processing depends on widely distributed cortical networks and could explain observations of focal stroke lesions affecting a large range of functions.

Ubiquitous Neocortical Decoding of Tactile Input Patterns.

Whereas functional localization historically has been a key concept in neuroscience, direct neuronal recordings show that input of a particular modality can be recorded well outside its primary receiving areas in the neocortex. Here, we wanted to explore if such spatially unbounded inputs potentially contain any information about the quality of the input received. We utilized a recently introduced approach to study the neuronal decoding capacity at a high resolution by delivering a set of electrical, highly reproducible spatiotemporal tactile afferent activation patterns to the skin of the contralateral second digit of the forepaw of the anesthetized rat. Surprisingly, we found that neurons in all areas recorded from, across all cortical depths tested, could decode the tactile input patterns, including neurons of the primary visual cortex. Within both somatosensory and visual cortical areas, the combined decoding accuracy of a population of neurons was higher than for the best performing single neuron within the respective area. Such cooperative decoding indicates that not only did individual neurons decode the input, they also did so by generating responses with different temporal profiles compared to other neurons, which suggests that each neuron could have unique contributions to the tactile information processing. These findings suggest that tactile processing in principle could be globally distributed in the neocortex, possibly for comparison with internal expectations and disambiguation processes relying on other modalities.

Intracellular Dynamics in Cuneate Nucleus Neurons Support Self-Stabilizing Learning of Generalizable Tactile Representations.

How the brain represents the external world is an unresolved issue for neuroscience, which could provide fundamental insights into brain circuitry operation and solutions for artificial intelligence and robotics. The neurons of the cuneate nucleus form the first interface for the sense of touch in the brain. They were previously shown to have a highly skewed synaptic weight distribution for tactile primary afferent inputs, suggesting that their connectivity is strongly shaped by learning. Here we first characterized the intracellular dynamics and inhibitory synaptic inputs of cuneate neurons in vivo and modeled their integration of tactile sensory inputs. We then replaced the tactile inputs with input from a sensorized bionic fingertip and modeled the learning-induced representations that emerged from varied sensory experiences. The model reproduced both the intrinsic membrane dynamics and the synaptic weight distributions observed in cuneate neurons in vivo. In terms of higher level model properties, individual cuneate neurons learnt to identify specific sets of correlated sensors, which at the population level resulted in a decomposition of the sensor space into its recurring high-dimensional components. Such vector components could be applied to identify both past and novel sensory experiences and likely correspond to the fundamental haptic input features these neurons encode in vivo. In addition, we show that the cuneate learning architecture is robust to a wide range of intrinsic parameter settings due to the neuronal intrinsic dynamics. Therefore, the architecture is a potentially generic solution for forming versatile representations of the external world in different sensor systems.

Correction to: Cerebellar Modules and Their Role as Operational Cerebellar Processing Units: A Consensus paper.

In the original version of this paper, the Title should have been written with "A Consensus paper" to read "Cerebellar Modules and Their Role as Operational Cerebellar Processing Units: A Consensus paper".

Cerebellar Modules and Their Role as Operational Cerebellar Processing Units: A Consensus paper [corrected].

The compartmentalization of the cerebellum into modules is often used to discuss its function. What, exactly, can be considered a module, how do they operate, can they be subdivided and do they act individually or in concert are only some of the key questions discussed in this consensus paper. Experts studying cerebellar compartmentalization give their insights on the structure and function of cerebellar modules, with the aim of providing an up-to-date review of the extensive literature on this subject. Starting with an historical perspective indicating that the basis of the modular organization is formed by matching olivocorticonuclear connectivity, this is followed by consideration of anatomical and chemical modular boundaries, revealing a relation between anatomical, chemical, and physiological borders. In addition, the question is asked what the smallest operational unit of the cerebellum might be. Furthermore, it has become clear that chemical diversity of Purkinje cells also results in diversity of information processing between cerebellar modules. An additional important consideration is the relation between modular compartmentalization and the organization of the mossy fiber system, resulting in the concept of modular plasticity. Finally, examination of cerebellar output patterns suggesting cooperation between modules and recent work on modular aspects of emotional behavior are discussed. Despite the general consensus that the cerebellum has a modular organization, many questions remain. The authors hope that this joint review will inspire future cerebellar research so that we are better able to understand how this brain structure makes its vital contribution to behavior in its most general form.

Artificial spatiotemporal touch inputs reveal complementary decoding in neocortical neurons.

Investigations of the mechanisms of touch perception and decoding has been hampered by difficulties in achieving invariant patterns of skin sensor activation. To obtain reproducible spatiotemporal patterns of activation of sensory afferents, we used an artificial fingertip equipped with an array of neuromorphic sensors. The artificial fingertip was used to transduce real-world haptic stimuli into spatiotemporal patterns of spikes. These spike patterns were delivered to the skin afferents of the second digit of rats via an array of stimulation electrodes. Combined with low-noise intra- and extracellular recordings from neocortical neurons in vivo, this approach provided a previously inaccessible high resolution analysis of the representation of tactile information in the neocortical neuronal circuitry. The results indicate high information content in individual neurons and reveal multiple novel neuronal tactile coding features such as heterogeneous and complementary spatiotemporal input selectivity also between neighboring neurons. Such neuronal heterogeneity and complementariness can potentially support a very high decoding capacity in a limited population of neurons. Our results also indicate a potential neuroprosthetic approach to communicate with the brain at a very high resolution and provide a potential novel solution for evaluating the degree or state of neurological disease in animal models.

The Roles of the Olivocerebellar Pathway in Motor Learning and Motor Control. A Consensus Paper.

For many decades, the predominant view in the cerebellar field has been that the olivocerebellar system's primary function is to induce plasticity in the cerebellar cortex, specifically, at the parallel fiber-Purkinje cell synapse. However, it has also long been proposed that the olivocerebellar system participates directly in motor control by helping to shape ongoing motor commands being issued by the cerebellum. Evidence consistent with both hypotheses exists; however, they are often investigated as mutually exclusive alternatives. In contrast, here, we take the perspective that the olivocerebellar system can contribute to both the motor learning and motor control functions of the cerebellum and might also play a role in development. We then consider the potential problems and benefits of it having multiple functions. Moreover, we discuss how its distinctive characteristics (e.g., low firing rates, synchronization, and variable complex spike waveforms) make it more or less suitable for one or the other of these functions, and why having multiple functions makes sense from an evolutionary perspective. We did not attempt to reach a consensus on the specific role(s) the olivocerebellar system plays in different types of movements, as that will ultimately be determined experimentally; however, collectively, the various contributions highlight the flexibility of the olivocerebellar system, and thereby suggest that it has the potential to act in both the motor learning and motor control functions of the cerebellum.

No Medium-Term Spinocerebellar Input Plasticity in Deep Cerebellar Nuclear Neurons In Vivo?

The existence of input plasticity in the deep cerebellar nuclear (DCN) cells of the adult cerebellum could have profound implications for our understanding of cerebellar function. Whereas the existence of plastic changes in mossy fiber (mf) synaptic responses in DCN neurons has been demonstrated in juvenile slices, there has so far been no direct demonstration of this form of plasticity in the adult cerebellum in vivo. In the present paper, we recorded from neurons in the anterior interposed nucleus (AIN) and stimulated the spinocerebellar tracts (SCT) directly or via the skin to obtain mf activation and the inferior olive to activate climbing fibers (cfs) in the nonanesthetized, adult, decerebrated cat. We used three different types of protocols that theoretically could be expected to induce plasticity, each of which involved episodically intense afferent activation lasting for 10 min. These were conjunctive mf-cf activation, which effectively induces plasticity in cortical neurons; mf and cf activation in a pattern resembling the protocol for inducing classical conditioning; and conjunctive activation of two excitatory mf inputs. None of these protocols had any statistically significant effect on the evoked responses in the AIN neurons. We conclude that the input plasticity for excitatory mfs in the AIN cells of the adult cerebellum in vivo is likely to be less effective than that of parallel fiber synaptic inputs in cerebellar cortical cells, at least in the timespan of 1 h.

Climbing Fiber Regulation of Spontaneous Purkinje Cell Activity and Cerebellum-Dependent Blink Responses(1,2,3).

It has been known for a long time that GABAergic Purkinje cells in the cerebellar cortex, as well as their target neurons in the cerebellar nuclei, are spontaneously active. The cerebellar output will, therefore, depend on how input is integrated into this spontaneous activity. It has been shown that input from climbing fibers originating in the inferior olive controls the spontaneous activity in Purkinje cells. While blocking climbing fiber input to the Purkinje cells causes a dramatic increase in the firing rate, increased climbing fiber activity results in reduced Purkinje cell activity. However, the exact calibration of this regulation has not been examined systematically. Here we examine the relation between climbing fiber stimulation frequency and Purkinje cell activity in unanesthetized decerebrated ferrets. The results revealed a gradual suppression of Purkinje cell activity, starting at climbing fiber stimulation frequencies as low as 0.5 Hz. At 4 Hz, Purkinje cells were completely silenced. This effect lasted an average of 2 min after the stimulation rate was reduced to a lower level. We also examined the effect of sustained climbing fiber stimulation on overt behavior. Specifically, we analyzed conditioned blink responses, which are known to be dependent on the cerebellum, while stimulating the climbing fibers at different frequencies. In accordance with the neurophysiological data, the conditioned blink responses were suppressed at stimulation frequencies of ≥4 Hz.

Climbing fiber receptive fields-organizational and functional aspects and relationship to limb coordination.

Climbing fiber receptive fields are a physiological marker that have proven useful to delineate the details of the olivocerebellar circuitry. They have also proven useful as a point of reference to delineate the organization of other parts of the cerebellar circuitry. But what does the location of the climbing fiber receptive field imply and what is its relation to the presumed role of the cerebellum in coordination? Can we expect that all climbing fibers have a peripheral receptive field on the skin? In this short review, we aim to cover these issues.

Properties of bilateral spinocerebellar activation of cerebellar cortical neurons.

We aimed to explore the cerebellar cortical inputs from two spinocerebellar pathways, the spinal border cell-component of the ventral spinocerebellar tract (SBC-VSCT) and the dorsal spinocerebellar tract (DSCT), respectively, in the sublobule C1 of the cerebellar posterior lobe. The two pathways were activated by electrical stimulation of the contralateral lateral funiculus (coLF) and the ipsilateral LF (iLF) at lower thoracic levels. Most granule cells in sublobule C1 did not respond at all but part of the granule cell population displayed high-intensity responses to either coLF or iLF stimulation. As a rule, Golgi cells and Purkinje cell simple spikes responded to input from both LFs, although Golgi cells could be more selective. In addition, a small population of granule cells responded to input from both the coLF and the iLF. However, in these cases, similarities in the temporal topography and magnitude of the responses suggested that the same axons were stimulated from the two LFs, i.e., that the axons of individual spinocerebellar neurons could be present in both funiculi. This was also confirmed for a population of spinal neurons located within known locations of SBC-VSCT neurons and dorsal horn (dh) DSCT neurons. We conclude that bilateral spinocerebellar responses can occur in cerebellar granule cells, but the VSCT and DSCT systems that provide the input can also be organized bilaterally. The implications for the traditional functional separation of VSCT and DSCT systems and the issue whether granule cells primarily integrate functionally similar information or not are discussed.

Simulating spinal border cells and cerebellar granule cells under locomotion--a case study of spinocerebellar information processing.

The spinocerebellar systems are essential for the brain in the performance of coordinated movements, but our knowledge about the spinocerebellar interactions is very limited. Recently, several crucial pieces of information have been acquired for the spinal border cell (SBC) component of the ventral spinocerebellar tract (VSCT), as well as the effects of SBC mossy fiber activation in granule cells of the cerebellar cortex. SBCs receive monosynaptic input from the reticulospinal tract (RST), which is an important driving system under locomotion, and disynaptic inhibition from Ib muscle afferents. The patterns of activity of RST neurons and Ib afferents under locomotion are known. The activity of VSCT neurons under fictive locomotion, i.e. without sensory feedback, is also known, but there is little information on how these neurons behave under actual locomotion and for cerebellar granule cells receiving SBC input this is completely unknown. But the available information makes it possible to simulate the interactions between the spinal and cerebellar neuronal circuitries with a relatively large set of biological constraints. Using a model of the various neuronal elements and the network they compose, we simulated the modulation of the SBCs and their target granule cells under locomotion and hence generated testable predictions of their general pattern of modulation under this condition. This particular system offers a unique opportunity to simulate these interactions with a limited number of assumptions, which helps making the model biologically plausible. Similar principles of information processing may be expected to apply to all spinocerebellar systems.

Segregation of tactile input features in neurons of the cuneate nucleus.

Our tactile perception of external objects depends on skin-object interactions. The mechanics of contact dictates the existence of fundamental spatiotemporal input features-contact initiation and cessation, slip, and rolling contact-that originate from the fact that solid objects do not interpenetrate. However, it is unknown whether these features are represented within the brain. We used a novel haptic interface to deliver such inputs to the glabrous skin of finger/digit pads and recorded from neurons of the cuneate nucleus (the brain's first level of tactile processing) in the cat. Surprisingly, despite having similar receptive fields and response properties, each cuneate neuron responded to a unique combination of these inputs. Hence, distinct haptic input features are encoded already at subcortical processing stages. This organization maps skin-object interactions into rich representations provided to higher cortical levels and may call for a re-evaluation of our current understanding of the brain's somatosensory systems.

Spike generation estimated from stationary spike trains in a variety of neurons in vivo.

To any model of brain function, the variability of neuronal spike firing is a problem that needs to be taken into account. Whereas the synaptic integration can be described in terms of the original Hodgkin-Huxley (H-H) formulations of conductance-based electrical signaling, the transformation of the resulting membrane potential into patterns of spike output is subjected to stochasticity that may not be captured with standard single neuron H-H models. The dynamics of the spike output is dependent on the normal background synaptic noise present in vivo, but the neuronal spike firing variability in vivo is not well studied. In the present study, we made long-term whole cell patch clamp recordings of stationary spike firing states across a range of membrane potentials from a variety of subcortical neurons in the non-anesthetized, decerebrated state in vivo. Based on the data, we formulated a simple, phenomenological model of the properties of the spike generation in each neuron that accurately captured the stationary spike firing statistics across all membrane potentials. The model consists of a parametric relationship between the mean and standard deviation of the inter-spike intervals, where the parameter is linearly related to the injected current over the membrane. This enabled it to generate accurate approximations of spike firing also under inhomogeneous conditions with input that varies over time. The parameters describing the spike firing statistics for different neuron types overlapped extensively, suggesting that the spike generation had similar properties across neurons.

Changing behavior towards sustainable practices using Information Technology.

This article addresses the question of how to change individuals' behavior towards more sustainable practices using Information Technology (IT). By following a multidisciplinary and socio-technical perspective, this inquiry is answered by applying a new framework-The Commonality Framework for IT-enabled Change-on a case study of sustainable behavioral change. The framework is grounded in practice theory and is used to analyze the implementation of an IT-system aimed at changing citizens' behavior towards more sustainable transport logistics and procurement in Uppsala, Sweden. The article applies case study research design and the empirical data consists of surveys, in-depth and semi-structured interviews, observations and archival documents. The results show how the change towards sustainable practices is an entanglement of both social and technical-structural elements across time. In this process, structures such as IT are the enablers, and the actors and their social activities are the tipping-point factors that ultimately determine the success of changing individuals' behavior towards a more sustainable direction. This article provides a more balanced view of how both actor and structure related properties interact during the on-going work with change towards greater sustainability practices than earlier research has offered. More specifically, the article offers both a lower-level theory and a method from which we can analyze change processes where technology is seen in its context, and where both technology and the human actor is brought forth to center stage.

Spatio-temporal skin strain distributions evoke low variability spike responses in cuneate neurons.

A common method to explore the somatosensory function of the brain is to relate skin stimuli to neurophysiological recordings. However, interaction with the skin involves complex mechanical effects. Variability in mechanically induced spike responses is likely to be due in part to mechanical variability of the transformation of stimuli into spiking patterns in the primary sensors located in the skin. This source of variability greatly hampers detailed investigations of the response of the brain to different types of mechanical stimuli. A novel stimulation technique designed to minimize the uncertainty in the strain distributions induced in the skin was applied to evoke responses in single neurons in the cat. We show that exposure to specific spatio-temporal stimuli induced highly reproducible spike responses in the cells of the cuneate nucleus, which represents the first stage of integration of peripheral inputs to the brain. Using precisely controlled spatio-temporal stimuli, we also show that cuneate neurons, as a whole, were selectively sensitive to the spatial and to the temporal aspects of the stimuli. We conclude that the present skin stimulation technique based on localized differential tractions greatly reduces response variability that is exogenous to the information processing of the brain and hence paves the way for substantially more detailed investigations of the brain's somatosensory system.

Specific relationship between excitatory inputs and climbing fiber receptive fields in deep cerebellar nuclear neurons.

Many mossy fiber pathways to the neurons of the deep cerebellar nucleus (DCN) originate from the spinal motor circuitry. For cutaneously activated spinal neurons, the receptive field is a tag indicating the specific motor function the spinal neuron has. Similarly, the climbing fiber receptive field of the DCN neuron reflects the specific motor output function of the DCN neuron. To explore the relationship between the motor information the DCN neuron receives and the output it issues, we made patch clamp recordings of DCN cell responses to tactile skin stimulation in the forelimb region of the anterior interposed nucleus in vivo. The excitatory responses were organized according to a general principle, in which the DCN cell responses became stronger the closer the skin site was located to its climbing fiber receptive field. The findings represent a novel functional principle of cerebellar connectivity, with crucial importance for our understanding of the function of the cerebellum in movement coordination.