The pathways Cortex CORTEX Lateral view of brain Development of cerebral cortex Lateral view cortex Cortical Divisions • Anatomically the cortex is divided into lobes: frontal, parietal, temporal, occipital. • • The insular cortex is also often considered as a separate lobe, the insular lobe The limbic lobe consists of regions of cortex such as the cingulate and para hippocampal gyri. • • Functionally the cortex is divided into numbered areas first proposed by Brodmann in 1909 Brodmann’s areas were described based on cytoarchitecture; later they were found to be functionally significant 6 Cortex: Coronal Section Cortical Connections Intracortical fibers – short, project to nearby cortical areas Commissural fibers – Connect homologous areas of the two hemispheres – Corpus callosum – Anterior commissure – Posterior commissure 8 Cortical Connections Association fibers – gyrus to gyrus and lobe to lobe in the same hemisphere – arcuate fibers connect adjacent gyri – long association fibers connect distant gyri Projection fibers – Projections from cortex to subcortical neurons (e.g. thalamus, corpus striatum, brainstem, and spinal cord – Internal capsule carries most of these connections. Topographic organization of functional areas Tonotopic mapping of auditory cortex Homunculus Ratunculus Species comparison The Ventricular System of the Human Brain The Major Arteries of the Brain • PN01202 .JPG Types of Cerebral Cortex • • • • Neocortex – Newest in evolution – About 90% of total – 6 layers, most complex Paleocortex – Associated with olfactory system, the parahippocampal gyrus, uncus – fewer than 6 layers Archicortex – Hippocampal formation; limbic system – 3 layers, most primitive Mesocortex – Cingulate gyrus, insular cortex – Transitional between archicortex and neocortex 16 Histology of the Cerebral Cortex • • • The neocortex has 6 layers designated I to VI Pyramidal cells are the characteristic neuron of the cortex and have axons that project to other cortical or subcortical regions. Cortical neurons such as stellate cells serve as interneurons for local processing. 17 Cortical Columns • • • • • Run vertically across all six layers Thousands of neurons in synaptic contact Main input layer is layer IV which receives thalamic input Are the basic units of the peripheral representation in the sensory cortex (e.g. retinotopy or tonotopy) Within a column neurons have similar response properties (e.g. characteristic frequency in the auditory cortex). 18 Cortical termination Corticothalamic Thalamocortical afferents terminate in: •Layer 4, spill into 3 and 5 •Layer 6 •Layer 1 spill into 2 Individual nuclei have projections to combinations of layers, partly depending on cells size. Thalamus Thalamus Divisions of thalamus For the most part when we refer to the ‘thalamus’ we really mean the dorsal thalamus. Most of the following material refers to the dorsal thalamus but you should be aware of the ventral thalamus that consists of the: •thalamic reticular nucleus •ventral lateral geniculate nuc. •zona incerta . The dorsal thalamus is divided into a number of nuclei. A basic definition of a thalamic nucleus is “a circumscribed region of cytoarchitecture receiving a particular set of afferent connections and projecting within the borders of a particular cortical field or fields.” Medial view Massa intermedia Medial wall of thalamus. Mammillary body Hypothalamus horizontal Lateral ventricle Corpus callosum thalamus Basal ganglia Internal capsule 3rd ventricle Human thalamus The human thalamus The rat thalamus Corpus callosum hippocampus Nissl stain Dorsal lateral geniculate nucleus Corpus callosum thalamus 3rd ventricle Myelin stain Thalamic nuclei can be categorized on their location within the thalamus Representative thalamic nuclei Name Afferents Cortical target Lateral geniculate (LGd) Retina Visual cortex (Brodmann area 17) Ventroposterior lateral (VPL) Medial lemniscus (Dorsal columns) Spinothalamic tract Primary somatosensory cortex. (Brodmann areas 3, 1, 2) Ventroposterior medial (VPM) Trigeminal nuclear complex Primary somatosensory cortex (Brodmann areas 3, 1, 2) Ventrolateral (VLp) Deep cerebellar nuclei Vestibular nuclei Globus pallidus Primary motor cortex (Brodmann area 4) Medial Geniculate Inferior colliculus Auditory cortex (Brodmann areas 41,42) generalized scheme Generalized scheme of thalamic circuitry including interneurons and the thalamic reticular nucleus p6 Topography Somatotopy (VPL) The thalamic glomerulus •D thalamocortical neuron dendrite •T1 principal afferent (Glu) •T2 local circuit neuron presynaptic dendrite (GABA) •G glial cell The glomerulus Tonic and burst mode Basal ganglia Basal Ganglia Relationship of Basal Ganglia to descending motor pathways Basal Ganglia definitions Caudate nucleus Striatum Putamen Basal Ganglia Globus pallidus Corpus Striatum Pallidum Substantia Nigra Subthalamic nucleus Basal ganglia Basal ganglia Basal ganglia: Coronal section Basal Ganglia horizontal Section Lateral ventricle Corpus callosum thalamus Basal ganglia Internal capsule 3rd ventricle Basal ganglia circuits The direct pathway Direct pathway GABA GABA Direct pathway GABA GABA The indirect pathway Substantia nigra Basal Ganglia circuitry Direct pathway facilitates movement Indirect pathway inhibits movement Cerebellum Cerebellum Relationship of cerebellum to descending motor pathways Cerebellum CEREBELLUM Overall Organization and Subdivisions Organization of cerebellar Outputs and Inputs input output Functional Organization of the Inputs Projections from the cerebellum Input to the cerebellum Open loop Cerebellar Closed Loop Primary Motor Cortex midline Thalamus (VA/VL) Deep cerebellar nuclei PONS (pontine nuclei) Proprioceptive (sensory) input a-motor neuron