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Modern Brain Mapping – What Do We Map Nowadays?

机译:现代人脑图谱-我们现在要映射什么?

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Introduction The problem of function localization in the brain is one of the most fundamental in neuroscience. There are two opposite paradigms relating to the problem: “modularism,” also known as “localism,” versus “holism,” which have been discussed for a long time ( 1 , 2 ). The debate in favor of one or another view can still be traced at all methodological levels – from the cell to the system. In this opinion paper we want to raise a question – what is meant nowadays by brain mapping? In addition, we want to highlight the necessity of being aware of occasionally occurring discontinuity in the research at different methodological scales. This problem is evident for experts in the field, but not always sufficiently so for early career researches. We will try to describe the difficulties of modern brain mapping primarily by looking at one of the currently best-studied functions – motor function. History of Opposition of Modular Versus Holistic Conceptions of Brain Organization A somewhat artificial opposition of “modular” and “holistic” organization of the brain has been evident in neuroscience from 18th century, and started mostly as a disagreement between physiologists working on animals and clinicians studying brain lesions in humans ( 1 – 3 ). A first revision of the term “function” by a clinician and a step away from hardwired localism was performed by the neurologists J. H. Jackson at the end of 19th century who wrote that “localization of a symptom is not localization of a function” ( 4 ). In the beginning of the 20th century, a paradigm shift occurred toward gestalt psychology, which changed the trend of research at the macro-scale level towards a more holistic view ( 2 ). A prime example of a confrontation at the micro-scale at the same time was the debate of Golgi and Cajal regarding the essence of a neuron ( 5 ). In 1937, a neurosurgeon W. Penfield performed the first cortical cartography in humans and published an iconic description of sensory and motor homunculi ( 6 ). In the second quarter of the 20th century, the concept of a function as a goal-dependent entity appeared in the form of theory of movements ( 7 ) and theory of functional systems ( 8 ), both viewing a function as a non-rigid goal-dependent entity. To date, it is usually postulated that localism and holism have been replaced by “connectionism,” with many studies nowadays trying to find interactions between brain regions and not the function of these regions by themselves ( 9 , 10 ). However, it seems there is still a tendency to favor localism, especially in the cognitive sciences ( 11 , 12 ). Perhaps, this is due to the fact that modern non-invasive methods, such as PET, fMRI or TMS, are mostly associated with functional mapping of the brain based on M. Minsky’s philosophy that “minds are what brains do.” Discontinuity of the Motor Research of Different Methodological Scales The question remains open of how cortical “activation” at the macro-level, viewed for example with fMRI or EEG, is linked to micro-scale phenomena such as single neuron activity in the spinal cord in awake animals (especially in humans) ( 13 , 14 ). There is a large community of researches studying ways of activation of a particular alpha-motor neurons in the spinal cord ( 15 – 17 ); scientists working on the level of a single neuron usually associate it to a specific task ( 16 , 18 ). Thereby, those who work with slices of the spinal cord are well aware of how to activate a certain motoneuron ( 19 ), but it is still difficult to bridge these phenomena with activation of the cortex ( 15 ). Modern macro-scale approaches connecting peripheral and central recording, such as TMS-EEG and corticomuscular coherence, including biofeedback, are trying to overcome this gap ( 20 ). A good example of a discontinuity in motor research at different methodological scales is the phenomena of convergence and divergence of motor cortex organization. They are well known in micro- and meso-scale studies. For example, in the invasive brain–computer interface (BCI) research, principles like neural degeneracy and neuronal multitasking were formulated ( 21 ). However, these phenomena are still widely overlooked in the research at the macro-scale level ( 22 ). For instance, a commonly used term in macro-scale research is an “area of a muscle cortical representation” ( 23 , 24 ), which is suitable for practical use like presurgical motor mapping ( 25 ), but is physiologically dubious considering the proven fact that some pyramidal cells may broadly innervate corresponding alpha motoneurons relating to activation of different muscles, even of different limb segments ( 26 – 28 ). Goal-Driven Concept of a Function – Problem of Awake Versus Anesthetized Animals Clearly demonstrated in many studies, context and goal-dependency of a motor function ( 29 , 30 ) brings us back to the necessity to revise the concept of a function as an environment and goal-dependent entity. Since a large amo
机译:简介大脑中的功能定位问题是神经科学中最基本的问题之一。关于这个问题有两种相反的范例:“模块化”,也称为“局部主义”与“整体主义”,已经讨论了很长时间(1、2)。仍然可以在所有方法论层面(从单元到系统)追溯支持一种或另一种观点的辩论。在这篇意见书中,我们想提出一个问题–如今的脑图绘制意味着什么?此外,我们想强调必须注意在不同方法学规模的研究中偶尔发生的不连续性。这个问题对于本领域的专家是显而易见的,但对于早期职业研究而言并不总是如此。我们将主要通过研究目前研究最深入的功能之一-运动功能来描述现代大脑映射的困难。反对模块化与整体大脑组织构想的历史从18世纪开始,神经科学就已经明显地有人为地反对大脑的“模块化”和“整体”组织,而这主要是因为从事动物研究的生理学家与临床医生之间的分歧人类的脑部病变(1-3)。 19世纪末,神经学家JH Jackson对临床医生对“功能”一词进行了首次修订,并摆脱了局限性的局限性。他写道:“症状的局部化不是功能的局部化”(4) 。在20世纪初,范式发生了向格式塔心理学的转变,这将宏观研究的趋势转向了更全面的观点(2)。在微观尺度上对抗的一个主要例子是高尔基和卡哈尔关于神经元本质的争论(5)。 1937年,神经外科医生W. Penfield对人类进行了第一次皮层成像,并发表了对感觉和运动性单向性的标志性描述(6)。在20世纪第二个季度,以运动理论(7)和功能系统理论(8)的形式出现了作为目标相关实体的功能概念,都将功能视为非刚性目标依赖实体。迄今为止,通常假定局部主义和整体主义已被“联系主义”所取代,如今许多研究试图寻找大脑区域之间的相互作用,而不是自己寻找这些区域的功能(9、10)。然而,似乎仍然有一种偏爱局部主义的趋势,尤其是在认知科学领域(11、12)。也许是由于这样的事实,即现代无创方法(例如PET,fMRI或TMS)大多与基于M. Minsky的思想“大脑是做什么的”的大脑功能映射有关。不同方法学规模的运动研究的不连续性还有一个问题仍然存在,那就是在宏观水平上皮层的“激活”如何与例如fMRI或EEG相联系,如何与微观现象如脊髓中单个神经元活动联系在一起。唤醒动物(尤其是人类)(13,14)。有大量的研究团体研究脊髓中特定的α运动神经元的激活方式(15 – 17)。研究单个神经元水平的科学家通常将其与特定任务相关联(16,18)。因此,那些研究脊髓切片的人都清楚如何激活特定的运动神经元(19),但是仍然很难通过激活皮质(15)来弥合这些现象。连接外围和中央记录的现代宏观方法,例如TMS-EEG和皮质生物相干性,包括生物反馈,正在试图克服这一差距(20)。运动学研究在不同方法学规模上的不连续性的一个很好的例子是运动皮层组织的收敛和发散现象。它们在微观和中观研究中是众所周知的。例如,在侵入性脑机接口(BCI)研究中,制定了诸如神经变性和神经元多任务处理等原理(21)。然而,这些现象在宏观层面的研究中仍然被广泛忽略(22)。例如,在宏观研究中通常使用的术语是“肌肉皮层代表区域”(23、24),它适合于诸如手术前运动作图(25)之类的实际使用,但考虑到已证实的事实,在生理上是可疑的一些锥体细胞可能广泛地支配与不同肌肉,甚至不同肢体节段的激活有关的相应的α运动神经元(26 – 28)。目标驱动的功能概念–清醒与麻醉动物的问题在许多研究中清楚地表明,运动功能的背景和目标依赖性(29,30)使我们回到修改功能作为环境的概念的必要性和目标相关实体。由于大弹药

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