Guarding the gateway to cortex with attention in visual thalamus

The massive visual input from the eye to the brain requires selective processing of some visual information at the expense of other information, a process referred to as visual attention. Increases in the responses of visual neurons with attention have been extensively studied along the visual processing streams in monkey cerebral cortex, from primary visual areas to parietal and frontal cortex. Here we show, by recording neurons in attending macaque monkeys (Macaca mulatta), that attention modulates visual signals before they even reach cortex by increasing responses of both magnocellular and parvocellular neurons in the first relay between retina and cortex, the lateral geniculate nucleus (LGN). At the same time, attention decreases neuronal responses in the adjacent thalamic reticular nucleus (TRN). Crick argued for such modulation of the LGN by observing that it is inhibited by the TRN, and suggested that "if the thalamus is the gateway to the cortex, the reticular complex might be described as the guardian of the gateway", a reciprocal relationship we now show to be more than just hypothesis. The reciprocal modulation in LGN and TRN appears only during the initial visual response, but the modulation of LGN reappears later in the response, suggesting separate early and late sources of attentional modulation in LGN.     

Parallel colour-opponent pathways to primary visual cortex

The trichromatic primate retina parses the colour content of a visual scene into 'red/green' and 'blue/yellow' representations. Cortical circuits must combine the information encoded in these colour-opponent signals to reconstruct the full range of perceived colours. Red/green and blue/yellow inputs are relayed by the lateral geniculate nucleus (LGN) of thalamus to primary visual cortex (V1), so understanding how cortical circuits transform these signals requires understanding how LGN inputs to V1 are organized. Here we report direct recordings from LGN afferent axons in muscimol-inactivated V1. We found that blue/yellow afferents terminated exclusively in superficial cortical layers 3B and 4A, whereas red/green afferents were encountered only in deeper cortex, in lower layer 4C. We also describe a distinct cortical target for 'blue-OFF' cells, whose afferents terminated in layer 4A and seemed patchy in organization. The more common 'blue-ON' afferents were found in 4A as well as lower layer 2/3. Chromatic information is thus conveyed to V1 by parallel, anatomically segregated colour-opponent systems, to be combined at a later stage of the colour circuit.     

Mixed parvocellular and magnocellular geniculate signals in visual area V4.

Visual information from the retina is transmitted to the cerebral cortex by way of the lateral geniculate nucleus (LGN) in the thalamus. In primates, most of the retinal ganglion cells that project to the LGN belong to one of two classes, P and M, whose axons terminate in the parvocellular or magnocellular subdivisions of the LGN. These cell classes give rise to two channels that have been distinguished anatomically, physiologically and behaviourally. The visual cortex also can be subdivided into two pathways, one specialized for motion processing and the other for colour and form information. Several lines of indirect evidence have suggested a close correspondence between the subcortical and cortical pathways, such that the M channel provides input to the motion pathway and the P channel drives the colour/form pathway. This hypothesis was tested directly by selectively inactivating either the magnocellular or parvocellular subdivision of the LGN and recording the effects on visual responses in the cortex. We have previously reported that, in accordance with the hypothesis, responses in the motion pathway in the cortex depend primarily on magnocellular LGN. We now report that in the colour/form pathway, visual responses depend on both P and M input. These results argue against a simple correspondence between the subcortical and cortical pathways.     

Activation of specific interneurons improves V1 feature selectivity and visual perception

Inhibitory interneurons are essential components of the neural circuits underlying various brain functions. In the neocortex, a large diversity of GABA (γ-aminobutyric acid) interneurons has been identified on the basis of their morphology, molecular markers, biophysical properties and innervation pattern. However, how the activity of each subtype of interneurons contributes to sensory processing remains unclear. Here we show that optogenetic activation of parvalbumin-positive (PV+) interneurons in the mouse primary visual cortex (V1) sharpens neuronal feature selectivity and improves perceptual discrimination. Using multichannel recording with silicon probes and channelrhodopsin-2 (ChR2)-mediated optical activation, we found that increased spiking of PV+ interneurons markedly sharpened orientation tuning and enhanced direction selectivity of nearby neurons. These effects were caused by the activation of inhibitory neurons rather than a decreased spiking of excitatory neurons, as archaerhodopsin-3 (Arch)-mediated optical silencing of calcium/calmodulin-dependent protein kinase IIα (CAMKIIα)-positive excitatory neurons caused no significant change in V1 stimulus selectivity. Moreover, the improved selectivity specifically required PV+ neuron activation, as activating somatostatin or vasointestinal peptide interneurons had no significant effect. Notably, PV+ neuron activation in awake mice caused a significant improvement in their orientation discrimination, mirroring the sharpened V1 orientation tuning. Together, these results provide the first demonstration that visual coding and perception can be improved by increased spiking of a specific subtype of cortical inhibitory interneurons.     

Disruption of retinogeniculate afferent segregation by antagonists to NMDA receptors.

Afferent activity has an important role in the formation of connections in the developing mammalian visual system. But the extent to which the activity of target neurons shapes patterns of afferent termination and synaptic contact is not known. In the ferret's visual pathway, retinal ganglion cell axons from each eye segregate early in development into eye-specific laminae in the lateral geniculate nucleus (LGN). The dorsal laminae (termed laminae A and A1) then segregate further into inner and outer sublaminae that retain input from on-centre and off-centre retinal axons, respectively. Thus, individual retinogeniculate axons form terminal arbors within laminae A and A1 that are restricted to one inner or outer sublamina. We report here that blockade of N-methyl-D-aspartate (NMDA) receptors on LGN cells with specific antagonists during the period of sublamina formation prevents retinal afferents from segregating into 'On' and 'Off' sublaminae. Retinogeniculate axons have arbors that are not restricted appropriately, or are restricted in size but inappropriately positioned within the eye-specific laminae. NMDA receptor antagonists may specifically disrupt a mechanism by which LGN neurons detect correlated afferent and target activity, and have been shown to reduce retinogeniculate transmission more generally, causing LGN cells to have markedly reduced levels of activity. These results therefore indicate that the activity of postsynaptic cells can significantly influence the patterning of inputs and the structure of presynaptic afferents during development.     

一种12-bit 1MSPs SAR ADC的设计

设计了工作在5 V单电源电压下,典型采样速度为1 MSPs的12位低功耗逐次逼近型模数转换器。设计中D/A转换器采用了加电容分压器的电荷分布式结构,在扩展并行D/A转换器分辨率的同时大大节省了芯片面积,内置的3.3 V参考电压源采用自偏置的供电方式,提高了基准电压的精度,同时也降低了功耗。使用cadence spectre工具进行仿真,后仿结果表明,设计的D/A转换器、3.3 V基准源满足12 bit A/D转换的要求,逐次逼近A/D转换器可以正常工作。     

Motion streaks provide a spatial code for motion direction.

Although many neurons in the primary visual cortex (V1) of primates are direction selective, they provide ambiguous information about the direction of motion of a stimulus. There is evidence that one of the ways in which the visual system resolves this ambiguity is by computing, from the responses of V1 neurons, velocity components in two or more spatial orientations and then combining these velocity components. Here I consider another potential neural mechanism for determining motion direction. When a localized image feature moves fast enough, it should become smeared in space owing to temporal integration in the visual system, creating a spatial signal-a 'motion streak'-oriented in the direction of the motion. The orientation masking and adaptation experiments reported here show that these spatial signals for motion direction exist in the human visual system for feature speeds above about 1 feature width per 100 ms. Computer simulations show that this psychophysical finding is consistent with the known response properties of V1 neurons, and that these spatial signals, when appropriately processed, are sufficient to determine motion direction in natural images.     

Preferential electrical coupling regulates neocortical lineage-dependent microcircuit assembly

Radial glial cells are the primary neural progenitor cells in the developing neocortex. Consecutive asymmetric divisions of individual radial glial progenitor cells produce a number of sister excitatory neurons that migrate along the elongated radial glial fibre, resulting in the formation of ontogenetic columns. Moreover, sister excitatory neurons in ontogenetic columns preferentially develop specific chemical synapses with each other rather than with nearby non-siblings. Although these findings provide crucial insight into the emergence of functional columns in the neocortex, little is known about the basis of this lineage-dependent assembly of excitatory neuron microcircuits at single-cell resolution. Here we show that transient electrical coupling between radially aligned sister excitatory neurons regulates the subsequent formation of specific chemical synapses in the neocortex. Multiple-electrode whole-cell recordings showed that sister excitatory neurons preferentially form strong electrical coupling with each other rather than with adjacent non-sister excitatory neurons during early postnatal stages. This preferential coupling allows selective electrical communication between sister excitatory neurons, promoting their action potential generation and synchronous firing. Interestingly, although this electrical communication largely disappears before the appearance of chemical synapses, blockade of the electrical communication impairs the subsequent formation of specific chemical synapses between sister excitatory neurons in ontogenetic columns. These results suggest a strong link between lineage-dependent transient electrical coupling and the assembly of precise excitatory neuron microcircuits in the neocortex.     

NgR在中菊头蝠(Rhinolophus affinis)脑中分布

Nogo和其受体相互作用可能在抑制神经再生中发挥着重要作用.运用免疫细胞化学方法,本研究观察了Nogo受体(NgR)在中菊头蝠脑中的分布.结果显示NgR在中菊头蝠脑皮层各层均有表达.在海马,NgR主要分布在CA1、CA3和DG区的神经细胞胞膜、胞质或/和突起上.杏仁核、丘脑、室旁核、视上核、视交叉上核等也有NgR阳性神经细胞着色.在脑的白质,轴突着色明显.小脑的分子层、Purkinje细胞层和颗粒细胞层均有NgR免疫阳性反应,其中阳性反应的颗粒细胞最多.这些提示NgR可能介导其配基对中菊头蝠脑多个区域的神经细胞起作用.     

Influence of the thalamus on spatial visual processing in frontal cortex

Each of our movements activates our own sensory receptors, and therefore keeping track of self-movement is a necessary part of analysing sensory input. One way in which the brain keeps track of self-movement is by monitoring an internal copy, or corollary discharge, of motor commands. This concept could explain why we perceive a stable visual world despite our frequent quick, or saccadic, eye movements: corollary discharge about each saccade would permit the visual system to ignore saccade-induced visual changes. The critical missing link has been the connection between corollary discharge and visual processing. Here we show that such a link is formed by a corollary discharge from the thalamus that targets the frontal cortex. In the thalamus, neurons in the mediodorsal nucleus relay a corollary discharge of saccades from the midbrain superior colliculus to the cortical frontal eye field. In the frontal eye field, neurons use corollary discharge to shift their visual receptive fields spatially before saccades. We tested the hypothesis that these two components-a pathway for corollary discharge and neurons with shifting receptive fields-form a circuit in which the corollary discharge drives the shift. First we showed that the known spatial and temporal properties of the corollary discharge predict the dynamic changes in spatial visual processing of cortical neurons when saccades are made. Then we moved from this correlation to causation by isolating single cortical neurons and showing that their spatial visual processing is impaired when corollary discharge from the thalamus is interrupted. Thus the visual processing of frontal neurons is spatiotemporally matched with, and functionally dependent on, corollary discharge input from the thalamus. These experiments establish the first link between corollary discharge and visual processing, delineate a brain circuit that is well suited for mediating visual stability, and provide a framework for studying corollary discharge in other sensory systems.     

Changes in geniculate cell size following brief monocular blockade of retinal activity in kittens

When a kitten is subjected to monocular lid suture early in life, cells in laminae of the lateral geniculate nucleus (LGN) connected to the sutured eye grow less than normal and cells in those laminae connected to the non-sutured eye grow more than normal. These changes are seen primarily in the binocular segment of the LGN, which corresponds to the central visual field, and are due to competition either between intracortical afferents originating from the different LGN laminae, or directly among cells within the LGN. The afferent deprivation induced by lid suture, however, is not complete, as retinal ganglion cells fire tonically both in darkness and in light. It is generally thought that this tonic retinal activity is necessary to maintain neuronal excitability at normal threshold in the central visual pathway. In the visual cortex of developing kittens, we previously showed a long-lasting change in ocular dominance of binocular cells by a brief blockade of retinal activity in one optic nerve. We report here that a complete blockade of retinal activity in one eye causes major changes in LGN cell size within 1 week. These changes occur throughout the LGN, including the monocular segment where binocular competition does not occur. The results indicate that tonic retinal activity may have an important role in the control of geniculate cell size.     

Clonally related visual cortical neurons show similar stimulus feature selectivity

A fundamental feature of the mammalian neocortex is its columnar organization. In the visual cortex, functional columns consisting of neurons with similar orientation preferences have been characterized extensively, but how these columns are constructed during development remains unclear. The radial unit hypothesis posits that the ontogenetic columns formed by clonally related neurons migrating along the same radial glial fibre during corticogenesis provide the basis for functional columns in adult neocortex. However, a direct correspondence between the ontogenetic and functional columns has not been demonstrated. Here we show that, despite the lack of a discernible orientation map in mouse visual cortex, sister neurons in the same radial clone exhibit similar orientation preferences. Using a retroviral vector encoding green fluorescent protein to label radial clones of excitatory neurons, and in vivo two-photon calcium imaging to measure neuronal response properties, we found that sister neurons preferred similar orientations whereas nearby non-sister neurons showed no such relationship. Interestingly, disruption of gap junction coupling by viral expression of a dominant-negative mutant of Cx26 (also known as Gjb2) or by daily administration of a gap junction blocker, carbenoxolone, during the first postnatal week greatly diminished the functional similarity between sister neurons, suggesting that the maturation of ontogenetic into functional columns requires intercellular communication through gap junctions. Together with the recent finding of preferential excitatory connections among sister neurons, our results support the radial unit hypothesis and unify the ontogenetic and functional columns in the visual cortex.     

非完美CSI下大规模MIMO DF中继多用户对全双工通信系统性能研究

通过解码转发（DF）协议中继,研究了在非完美信道状态信息（CSI）下,多用户对双向全双工通信系统的性能.通过配备大量天线,可以大大增加通信系统的频谱效率（SE）.对到达中继端的信号采用迫零（ZF）线性接收处理并预编码,推导了SE的近似表达式,提出了与天线数量相关的功率缩放定律,讨论了在导频功率固定和变化条件下缩放因子的取值.所提的功率缩放定律可以在保持稳定的频谱效率的同时,将发射功率降低.研究了在非完美信道状态信息下,经过解码转发协议中继的多用户对双向全双工通信系统的性能.     

An RNA gene expressed during cortical development evolved rapidly in humans

The developmental and evolutionary mechanisms behind the emergence of human-specific brain features remain largely unknown. However, the recent ability to compare our genome to that of our closest relative, the chimpanzee, provides new avenues to link genetic and phenotypic changes in the evolution of the human brain. We devised a ranking of regions in the human genome that show significant evolutionary acceleration. Here we report that the most dramatic of these 'human accelerated regions', HAR1, is part of a novel RNA gene (HAR1F) that is expressed specifically in Cajal-Retzius neurons in the developing human neocortex from 7 to 19 gestational weeks, a crucial period for cortical neuron specification and migration. HAR1F is co-expressed with reelin, a product of Cajal-Retzius neurons that is of fundamental importance in specifying the six-layer structure of the human cortex. HAR1 and the other human accelerated regions provide new candidates in the search for uniquely human biology.     

How did brains evolve?

Three reports on mammalian brain evolution analyse the same comparative data on brain component volumes but come to partially conflicting conclusions. Clark et al. conclude from their analysis of volumetric brain proportions ("cerebro-types") that cerebellum size is invariant across mammalian taxonomic groups, the neocortex and cerebellum do not co-vary in size (in contradiction to ref. 1), and cerebrotype-based measures identify directional changes in brain architecture. Here I provide evidence that calls each of these conclusions into question. The failure of the cerebrotype measure to identify species differences in brain architecture that are independent of gross brain size undermines the proposal by Clark et al. that it could be useful for detecting evolutionary patterns and phylogenetic relationships.     

Requirement for subplate neurons in the formation of thalamocortical connections

The neurons of layer 4 in the adult cerebral cortex receive their major ascending inputs from the thalamus. In development, however, thalamic axons arrive at the appropriate cortical area long before their target layer 4 neurons have migrated into the cortical plate. The axons accumulate and wait in the zone below the cortical plate, the subplate, for several weeks before invading the cortical plate. The subplate is a transient zone that contains the first postmitotic neurons of the telencephalon. These neurons mature well before other cortical neurons, and disappear by cell death after the thalamic axons have grown into the overlying cortical plate. The close proximity of growing thalamocortical axons and subplate neurons suggests that they might be involved in interactions important for normal thalamocortical development. Here we show that early in development the deletion of subplate neurons located beneath visual cortex prevents axons from the lateral geniculate nucleus of the thalamus from recognizing and innervating visual cortex, their normal target. In the absence of subplate neurons, lateral geniculate nucleus axons continue to grow in the white matter past visual cortex despite the presence of their target layer 4 neurons. Thus the transient subplate neurons are necessary for appropriate cortical target selection by thalamocortical axons.     

硅酸镓镧、钽酸镓镧和铌酸镓镧的声表面波传播特性

计算了硅酸镓镧、钽酸镓镧和铌酸镓镧的不同切向和传播方向的声表面波(SAW)传播特性，并对其和石英晶体上相应切向和传播方向的SAW传播特性做了对比分析．计算结果表明，这3种材料的SAW传播速度一般比石英低1km／s左右；和石英一样，具有零温度切向和纯模方向；机电耦合系数k^2却远高于石英，其中钽酸镓镧的k^2最大．对不同材料常数进行了对比计算，结果表明，不同材料常数的SAW传播特性有明显的差异，其中以频率温度系数差异最大．     

Eye-specific effects of binocular rivalry in the human lateral geniculate nucleus

When dissimilar images are presented to the two eyes, they compete for perceptual dominance so that each image is visible in turn for a few seconds while the other is suppressed. Such binocular rivalry is associated with relative suppression of local, eye-based representations that can also be modulated by high-level influences such as perceptual grouping. However, it is currently unclear how early in visual processing the suppression of eye-based signals can occur. Here we use high-resolution functional magnetic resonance imaging (fMRI) in conjunction with a new binocular rivalry stimulus to show that signals recorded from the human lateral geniculate nucleus (LGN) exhibit eye-specific suppression during rivalry. Regions of the LGN that show strong eye-preference independently show strongly reduced activity during binocular rivalry when the stimulus presented in their preferred eye is perceptually suppressed. The human LGN is thus the earliest stage of visual processing that reflects eye-specific dominance and suppression.     

An unexpected specialization for horizontal disparity in primate primary visual cortex

The horizontal separation of the eyes means that objects nearer or farther than the fixation point project to different locations on the two retinae, differing principally in their horizontal coordinates (horizontal binocular disparity). Disparity-selective neurons have generally been studied with disparities applied in only one direction (often horizontal), which cannot determine whether the encoding is specialized for processing disparities along the horizontal axis. It is therefore unclear if disparity selectivity represents a specialization for naturally occurring disparities. I used random dot stereograms to study disparity-selective neurons from the primary visual cortex (V1) of awake fixating monkeys. Many combinations of vertical and horizontal disparity were used, characterizing the surface of responses as a function of two-dimensional disparity. Here I report that the response surface usually showed elongation along the horizontal disparity axis, despite the isotropic stimulus. Thus these neurons modulated their firing rate over a wider range of horizontal disparity than vertical disparity. This demonstrates that disparity-selective cells are specialized for processing horizontal disparity, and that existing models of disparity selectivity require substantial revision.     

Grouping of image fragments in primary visual cortex.

In the visual world, objects are partially occluded by nearer objects, separating them into image fragments. However, the image fragments of the object can easily be grouped and organized together by the visual system. Psychophysical data and theoretical analysis indicate that such perceptual grouping might be mediated in the early stages of visual processing. Here I show that some orientation-selective cells in the primary visual cortex (V1) have response properties that can mediate the grouping of image fragments. These cells stopped responding to a stimulus bar when it was partly occluded by a small patch. The cells also did not respond when the patch had uncrossed disparity so that it appeared to be behind the bar. However, the cells began responding again when the patch had crossed disparity so that it appeared to be in front of the bar. These results indicate that cells as early as V1 have the computational power to make inferences about the nature of partially invisible forms seen behind occluding structures.