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2016-10-17T14:10:37.000000Z
字数 11784
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作业
Sensory processing is clearly affected bymental state. Everyone has had the experienceof sitting in a lecture and spacingout—even though you are awake andsound waves have been hitting youreardrum, you have no idea of what wassaid in the past five minutes. But what ishappening to those signals in your brain?
检测器处理显然是影响智力的状态。每个人都有经验,坐在讲座和做白日梦即使你清醒和声波已经击中你的耳膜,你不知道在过去的五分钟什么说道。但所发生的这些信号的确是在影响你的大脑吗?
Numerous studies have examined thedifferences in ongoing neural activity andthe interaction with incoming sensoryinput across different behavioral statesfrom sleep versus wake to selectivespatial attention. Recently, these studieshave extended to the rodent, where motorstates such as active whisking (Poulet andPetersen, 2008) and locomotion (Niell andStryker, 2010; Schneider et al., 2014) havebeen shown to correlate with changes insensory representations, even in the primarysensory cortices. Despite the varietyof these states, they can all be seen tovary along the dimension of arousal.
无数的研究已经证明这种差异在持续的神经活动和互动与传入对于不同的行为从睡眠与觉醒的空间选择性会产生。最近,这些研究扩展到啮齿类动物,在运动状态,如主动咀嚼(Poulet andpetersen,2008)和运动(Niell andstryker,2010;Schneider等人,2014)被证明与感觉的陈述相关的变化,即使在初级感觉皮层。尽管有各种各样的这些状态,他们都可以看到沿唤醒维度tovary。
Psychologists have encapsulated theeffects of arousal on behavioral performancein what is known as the Yerkes-Dodson curve (Yerkes and Dodson,1908), observing that performance on difficulttasks generally peaks at an optimal intermediatelevel of arousal (Figure 1). If yourarousal level is too low, you are spaced outor disengaged, and if your arousal level istoo high, then you are anxious anddistracted—in both cases performance suffers. In this issue of Neuron, McGinleyet al. (2015) deliver a tour-de-force explorationof the Yerkes-Dodson relationship inthe auditory system ofmice, finding it manifesteda cross multiple levels from theresting membrane potential of individual neurons to sensory-evoked synaptic inputs to behavioral performance.
心理学家对行为表现的影响封装的觉醒是耶基斯德森曲线(耶基斯和多德森,1908),观察性能的重任一般峰在觉醒的最佳中级水平(图1)。如果你的觉醒水平太低,你的间隔或脱离,如果你唤醒水平太高,那么你的焦虑和分心,在这两种情况下的性能会受到影响。在这个问题上的神经元,mcginleyet铝。(2015)提供一个绝技对耶基斯德森的关系在听觉系统的小鼠,发现它manifesteda跨多层次个体神经元膜电位从静息到感觉诱发的突触输入的行为表现。
The authors use a deceptively simplemeasure of behavioral state: pupil diameter.In addition to varying with ambient light level, it is also well established that pupil diameter in humans correlates withvarious aspects of arousal, from task difficultyto affective state; it is even possibleto track the number of items a person isholding in memory based on their pupil diameter (Laeng et al., 2012). Two recentstudies in the mouse visual system haveused pupil diameter as a metric for arousal state (Reimer et al., 2014; Vinck et al.,2015), and McGinley et al. (2015) begintheir study by confirming that pupil diameter agrees with an internal neural measureof arousal, the rate of hippo campalripple waves. Notably, pupil diameter providesa continuous variable measuring arousal, rather than simply a discrete categorization into states. However, an important caveat is that a number of factors cancontribute to arousal as measured by pupil diameter, from mental effort to anxiety tophysical activity, so the precise nature ofthe arousal is not determined.
作者用一个看似来衡量行为状态:瞳孔直径。除了与环境光水平不同,它也建立在人类的瞳孔直径相关的觉醒的各个方面,从任务困难的情感状态;它甚至可以跟踪项目的人在记忆上基于他们的瞳孔直径的数量(朗等人,2012)。在小鼠视觉系统采用瞳孔直径两研究作为唤醒状态度量(Reimeret al.,2014;文克etal.,2015),和金尼等人。(2015)来确认,瞳孔直径与内部神经的兴奋同意研究,河马campalripple波率。值得注意的是,瞳孔直径的测量提供了连续可变的觉醒,而不是简单的分类成离散状态。然而,值得注意的是,许多因素可以促进觉醒的瞳孔直径的测量,从精神焦虑的体育活动,因此精确性的觉醒是不确定的。
Using pupil diameter as their metric, theauthors first examine neural correlates of arousal using whole-cell recordings ofspontaneous activity in auditory cortexof awake head-fixed mice on a running wheel. Remarkably, they find that arousal,as measured by pupil diameter, predicts variations in membrane potential. In fact,on the timescale of seconds there isover 50% coherence between pupil diameterand membrane potential. In otherwords, pupil diameter can predict morethan half the variance in membrane potentialof neurons in auditory cortex.
利用瞳孔直径为度量,作者首先考察唤醒使用全细胞记录的自发活动在清醒小鼠跑轮固定在听觉皮层的神经相关。值得注意的是,他们发现,觉醒,作为测量瞳孔直径,预测在膜电位的变化。事实上,在几秒钟时间有超过50%学生、膜电位之间的连贯性。换句话说,瞳孔直径可以预测比听觉皮层神经元膜电位的半方差。
The authors demonstrate that, just asthe Yerkes-Dodson curve predicts, therelationship of membrane potential with arousal is U-shaped, as is the variabilityof membrane potential. At low levels of arousal, slow-wave oscillations begin,which increase variability and raise themembrane potential. At high levels of arousal, there is a tonic depolarization ofmembrane potential and high-frequency oscillations, which is often referred to asthe desynchronized state. In between, atwhat might be expected to be the optimal point on the Yerkes-Dodson curve, themembrane potential sits relatively lowand quiet. This is also reflected in spontaneous firing rate, which is lowest at intermediate levels of arousal.
作者表明,正如耶基斯德森曲线预测,与觉醒的关系膜电位是U形的,为的是变异性膜电位。在低层次的觉醒,慢波振荡开始,增加和提高膜电位的变化。在高层次的觉醒,有滋补的去极化膜电位和高频振荡,这是通常被称为非同步状态。在什么可能是耶基斯-多德森曲线的最佳点,膜电位较低,安静的坐着。这也反映在自发的发射率,这是最低的在中间水平的觉醒。
Again, sensory-evoked responses,both at the level of synaptic potential sand multi-unit firing, show a U-shapedcurve, now inverted with maximal evoked responses at the mid-point. Thus, adecrease in background variance and increasein evoked response work togetherto maximize the signal-to-noise ratio at intermediate levels of arousal.
再次,感觉诱发电位,在突触电位砂多单元发射的水平,显示u-shapedcurve,现在倒最大诱发反应的中点。因此,在背景方差减少和增加诱发反应共同最大化在觉醒的中级水平的信噪比。
To tie this all back to the initial Yerkes-Dodson finding, the authors find that performanceof the detection task follows theU-shaped prediction. When animals areat low arousal, they often miss the stimulus,and at high arousal they often falsealarm. Furthermore, the peak of the performancecurve was at a similar level ofarousal (pupil diameter) as the peak forneural encoding of the sensory input.
把这一切回到最初的耶基斯德森发现,发现性能的检测任务:U形预测。当动物处于低唤醒,他们经常错过的刺激,并在高唤醒他们经常误报警。此外,该曲线的峰值是在同一水平ofarousal(瞳孔直径)为高峰forneural编码的感官输入。
An important aspect of their approachis that the mice, rather than beingrestricted to periods of high performance,were allowed to drift between behavioral states, which enabled the authors tomap out the continuum of arousal. This provides a clear demonstrationthat, just because an animalis awake and performing,it is not in a specificwell-defined state. In fact,just as one would measure depth of an esthesia in ananesthetized experiment, ifone wants to compare across experiments it is necessary toeither control (as best aspossible) or measure (asbest as possible) an animal’s behavioral state.
一个重要的方面,他们应用小鼠,而不是国家对高性能的时期,被允许漂移之间的行为状态,使作者绘制了觉醒的连续体。这提供了一个阐述清楚,只是因为一个动物清醒和表演,它不是一个specificwell定义的状态。事实上,正如人会衡量在ananesthetized实验麻醉深度,如果要比较实验组控制是必要的(尽可能)或测量(石棉越好)动物的行为状态。
This study also providesinsight into another recentlyused measure of behavioral state: locomotion. Like otherstudies, they find that loco motionis accompanied (andin fact preceded) by pupil dilation, suggesting it is often a consequence of arousal. Indeed, in the experiments herethe effect of locomotion was not greatlydifferent than high arousal alone, exceptfor a few measures such as false alarmrate and evoked firing in MGN. However,in this study locomotion was always associatedwith arousal. Other recent studiesin the visual system (Reimer et al., 2014;Vinck et al., 2015) have been able to segregate arousal from locomotion andfound that while arousal accounts formany effects correlated with loco motion,there are distinct contributions of loco motion alone as well.
本研究也为进入另一个好的衡量行为状态:运动。像其他的研究,他们发现,伴随局部运动(实际上之前)的瞳孔放大,这往往是一个后果的觉醒。事实上,在运动的实验效果并不比在不同高觉醒本身,除了少数的措施如虚警率和诱发放电的MGN。然而,在这项研究中,运动总是与觉醒。其他最近的研究视觉系统(Reimer et al.,2014;文克et al.,2015)已经能够将觉醒运动发现而唤醒许多影响机车运动相关账户,有明显的贡献以及局部运动。
The different coupling between loco motionand arousal in these studies illustrates a limitation in using loco motor speed as a single scalar metric ofbehavior. An animal can run for many reasons—in some cases it may represent hyper-arousal, such as startle, whereas inothers it may represent an optimal state,such as goal-directed navigation. Furthermore,locomotor speed itself is an importantvariable that the brain is likely torepresent independently from arousal state, and in fact recent studies havedemonstrated continuous encoding of loco motor speed in visual cortex (Saleemet al., 2013). Thus, locomotion likely representsboth an internal state that ispartially correlated with arousal, as wellas a physical variable that is importantfor both navigation and processing sensoryinformation relative to self-motionand self-generated noise.
这些研究中的局部运动唤起不同的耦合关系说明使用运动速度作为一个标量度量行为的限制。一种动物能跑的原因很多,在某些情况下,它可能代表的超兴奋的,如惊吓,而其他可能代表一个最佳的状态,如目标的导航。此外,运动速度是一个重要变项,大脑可能代表独立于觉醒状态,事实上,最近的研究表明连续的视觉皮层中运动速度编码(saleemet al.,2013)。因此,运动可能既象征一个内部状态,唤醒部分相关,以及一个物理变量,对导航和处理感觉信息的相对自我和自我产生的噪声。
The effects of state showa striking difference across sensory modalities. As shownhere, in auditory cortex both high arousal and locomotion are coupled with adecrease in sensory-evoked responses,whereas other studies have shown that invisual cortex these are associated with anincrease in response gain. These differences maybe due todiffering requirementsfor sensory processing. For example, invision movement through the environment predictably interacts with the sensory input, whereas in audition movement cancause self-generated noise that needs tobe canceled. On the other hand, thesemay also represent differences in wheneach sensory modality is engaged ethologically—vision may bemost important when navigating, whereas audition may be mostimportant for detecting predators.
国家昭和显著差异的跨感官的方式的影响。为赞赏,听觉皮层高觉醒和运动耦合减少诱发反应,而其他的一些研究已经表明,在视觉皮层有响应增益增加。这些差异可能是由于感觉到不同要求的处理。例如,通过视觉运动环境与感觉输入可预见的相互作用,而在试听运动能引起自我产生的噪声,需要取消。另一方面,这些也代表了当各个感官差异是最重要的行为可能从事视觉导航时,而可能是重要的检测大鳄试镜。
The findings presented by McGinleyet al. (2015), along with the diverse effects seen across sensory modalities, raise thequestion as to which neural circuits underlie the state changes. A likely candidate forglobal changes in state, such as arousal, is neuro modulation, and evidence points to both norepinephrine(NE) andacetylcholine(Ach). Pupil dilation is often thought to beassociated with noradrenergic tone (Aston-Jones and Cohen, 2005), and a recentstudy by Polack et al. (2013) showed that NE was necessary for the elevation in base line membrane potential with locomotion,similar to the correlated changes in membranepotential and pupil diameter demonstratedhere. On the other hand, studies have demonstrateda role for cholinergic inputsto cortex in regulating thestrength of response to a visualstimulus (Fu et al., 2014; Pintoet al., 2013). These findingsmay be reconciled by a modelin which NE regulates a neuron’s‘‘set point’’ in terms ofresting membrane potential,whereas Ach regulates sensory-evoked responses, whichis supported by pharmacological and lesion experiments insoma to sensory cortex (Constantinopleand Bruno, 2011).
通过mcginleyet Al的结果。(2015),随着跨感官的方式看到了不同的影响,以提高问题的神经电路的基础状态的变化。一个可能的候选人的整体状态的变化,如兴奋,是神经调制,和证据表明,去甲肾上腺素(NE)和乙酰胆碱(ACh)。瞳孔的扩张通常被认为与去甲肾上腺素的语气(Aston Jones和科恩,2005),并由波兰等人最近的研究。(2013)表明,NE在运动基线膜电位的升高是必要的,类似于相关改变膜电位和瞳孔直径demonstratedhere。另一方面,研究在调节反应强度一visualstimulus胆碱能输入皮质种角色(Fu et al.,2014;Pintoet al,2013)。这些研究结果可能是和解的模型NE调节神经元的''set点”方面ofresting膜电位,而乙酰胆碱诱发反应的调节,是通过药物和病变的实验insoma感觉皮层(constantinopleand支持布鲁诺,2011)。
This raises the possibility thatthese well-trained mice had learned toregulate their state for the task at hand,although this remains to be tested bymeasuring state fluctuations in untrainedmice. If so, investigating the circuit mechanismsunderlying the acquisition of this metacognitive control could have importantimplications for both educationaland therapeutic interventions.
这就提出了一个可能性这些训练有素的老鼠学会了调整自身状态为手头的任务,尽管这仍有待检验测定untrainedmice状态波动。如果是这样的话,对电路基础这一元认知控制的收购可以为教育和治疗干预的重要意义。
Interestingly, the initial study by YerkesandDodson(1908)was actually performedin mice, measuring the rate of learning in avisual task, and was subsequently adoptedin human psychology. The work of McGinley et al. (2015) therefore representsa full circle, from mice to humans and backto mice. With new measures of behavioral state and powerful tools for observingand manipulating neural circuits, experimentsin both mice and humans can now continue to explore how neural dynamics give rise to the ups and downs in our dailyexperience.
有趣的是,通过yerkesanddodson的初步研究(1908)实际上是对小鼠,测定在视觉任务的学习率,并随后在人类心理学。对金尼等人的工作。(2015)因此,一个完整的圆,从小鼠到人类和小鼠回。为观察和操纵的神经回路的行为状态和强大的工具的新措施,实验小鼠和人类现在可以继续探讨神经动力学的产生在我们日常所遭遇的跌宕起伏。
摘要:
This article mainly tells about a method using in detecte the potential signature of optimal states for sensoory signal with 5 ways.
The different coupling between loco motionand arousal in these studies illustrates a limitation in using loco motor speed as a single scalar metric ofbehavior. An animal can run for many reasons—in some cases it may represent hyper-arousal, such as startle, whereas inothers it may represent an optimal state,such as goal-directed navigation.
sensory processing is clearly affected bymental state