Brainstem (II) 李立仁 副教授 解剖學暨細胞生物學科 [email protected] Brainstem (II) Long tracts in the brainstem Corticospinal tract (voluntary movement) The posterior column‐medial lemniscus system (touch and proprioception) Anterolateral system (pain and temperature) Spinocerebellar tract (proprioception) Brainstem (II) Long tracts in the brainstem Corticospinal tract (voluntary movement) Projecting fibers carrying commands for initiation of voluntary movements originated from motor, premotor, supplementary motor and somatosensory cortices descend through the internal capsule, cerebral peduncle and basal pons in company with corticopontine and corticobulbar fibers. Brainstem (II) Long tracts in the brainstem Corticospinal tract (voluntary movement) Corticospinal axons continue into the pyramids of the medulla. At the spinomedullary junction, most corticospinal fiber decussate (pyramidal decussation) to form the lateral corticospinal tract. Those do not cross in the pyramidal decussation continue into the smaller anterior corticospinal tract that typically crosses in the spinal cord before terminating. Brainstem (II) Long tracts in the brainstem The posterior column‐medial lemniscus system (touch and proprioception) Large‐diameter afferents with soma in the DRG (1st order), conveying information about position and movement of limb and the details of tactile stimuli, enter the spinal cord and ascend through the ipsilateral posterior funiculus (column) and terminates in the ipsilateral posterior column nuclei (nuclei gracilis and cuneatus). Axons from these nuclei (2nd order neurons) cross the midline as part of the internal arcuate fibers at the level of midmedulla (sensory decussation) and ascend to the ventral posterolateral nucleus (VPL) of the thalamus. 3rd order neurons in VPL projects to primary somatosensory cortex in postcentral gyrus. Brainstem (III) Cranial nerves and nuclei Main trigeminal sensory nucleus (V) Central processes of primary afferents with cell bodies in the trigeminal ganglion (1st order) terminate in the main sensory nucleus of the trigeminal nerve. Axons of 2nd neurons cross the midline, join the medial lemniscus, and ascend to the ventral posteromedial nucleus (VPM) of the thalamus. 3rd neurons in VPM in turn project to the face area of the postcentral gyrus. Touch and proprioception of face Somatic sensory nerves Brainstem (II) Anterolateral system (pain and temperature) Small‐diameter afferents with cell body in the dorsal root ganglion (1st order), conveying pain and temperature (and tactile) information, enter the spinal cord in the lateral division of each dorsal root and terminate on the tract cells in the posterior horn. Axons of 2nd neurons cross and join the spinothalamic tract (somatotopic arranged), then ascend to the ventral posterolateral nucleus (VPL) of the thalamus. Long tracts in the brainstem Brainstem (II) Long tracts in the brainstem Spinocerebellar tract (proprioception) Lower limb and trunk: Posterior spinocerebellar tract Primary afferents (1st order) terminate on the interneurons (2nd order) in the dorsal horn (Clarke's nucleus) on the same side. Projecting axons (2nd order) ascend without crossing and form synapses in cerebella by way of inferior cerebellar peduncle. Brainstem (II) Long tracts in the brainstem Spinocerebellar tract (proprioception) Upper limb and neck: Spinocuneocerebellar tract From above T1, primary afferents enter the fasciculus cuneatus directly and terminate on neurons in the accessory cuneate nucleus (2nd order). Projecting axons (2nd order) enter into the ipsilateral cerebellum via the inferior cerebellar peduncle. Brainstem (II) Neurochemistry of the brainstem “Out of billions of neurons in the human brain, relatively few appear to contain biogenic amines – such cells number only in the thousands……many of the cells containing these transmitters are clustered together in a discrete region of the brain, the brainstem.” ‐‐ Nicholls et al., From Neuron To Brain Glutamate is the most common excitatory transmitter in neurons throughout the brain. Gamma‐aminobutyric acid (GABA) is nearly a ubiquitous inhibitory neurotransmitter. Brainstem (II) Neurochemistry of the brainstem Brainstem (II) Neurochemistry of the brainstem Norepinephrine (noradrenaline) by neurons only found in pons and medullary tegmental regions. ‐ Locus ceruleus (blue spot) near the floor of the 4th ventricle ‐‐ project mainly to cerebral cortex ‐‐ silent during sleep, active during wakefulness ‐‐ form part of the ascending reticular activating system ‐ Reticular formation in the lateral part of medulla ‐‐ send fibers to spinal cord ‐ Solitary nucleus and dorsal motor nucleus of vagus Brainstem (II) Neurochemistry of the brainstem Norepinephrine (noradrenaline) innervates the entire CNS ‐ ascending fibers to thalamus, hypothalamus, limbic forebrain and cerebral cortex (SI) ‐ descending fibers to brainstem, cerebellum and spinal cord Brainstem (II) Neurochemistry of the brainstem Dopamine used by neurons mostly located in the midbrain ‐ compact part of substantia nigra (SNc) ‐ ventral tegmental area (VTA) involved in the initiation of movement cf. Parkinson’s disease also involved in motivation and cognition cf. drug addiction and schizophrenia Brainstem (II) Neurochemistry of the brainstem Dopamine projecting fibers are grouped into three bundles ‐ nigrostriatal (mesostriatal) afferents: SNc to caudate nucleus and putamen) ‐ mesolimbic afferents: VTA to limbic structures ex. amygdala and hippocampus ‐ mesocortical afferents: VTA to cerebral cortex (frontal) Brainstem (II) Neurochemistry of the brainstem Serotonin (5‐HT) used by neurons in the raphe nuclei throughout the BS ‐ rostral raphe nuclei project to forebrain (sensory and limbic) ‐ caudal raphe nuclei project to brainstem and spinal cord and may modulate thetransmission of pain ‐ form part of the ascending reticular activating system ‐ target of antidepressants Brainstem (II) Neurochemistry of the brainstem Acetylcholine used as neurotransmitter by some neuron groups in the RF, motor and preganglionic autonomic neurons involved in sleep‐wakefulness control Brainstem (II) Reticular formation A diffusely organized area in the central portion of the brainstem) Caudally, RF is continuous with the intermediate gray matter of the spinal cord. Rostrally, RF continues into the intralaminar nuclei of the thalamus. Brainstem (II) Four major groups : Precerebellar reticular nu. coordination of muscle contraction Raphe (median) and catacholamine nu. sleep‐alertness pain modulation Brainstem (II) Central (medial) reticular nu. eye movement conscious state Lateral reticular nu. respiratory and circulatory system feeding Brainstem (II) Functions of reticular formation Control of movement Spinal motor neurons are influenced (in addition to the corticospinal pathway) by reticulospinal pathways. Reticulospinal neurons receive collaterals from spinoreticular fibers and spinothalamic fibers and inputs from red nucleus, cerebellum and somatosensory as well as motor cortices. The reticulospinal tracts carry descending motor commands generated within RF. Reticular formation Brainstem (II) Functions of reticular formation Modulates pain transmission Neurons in the periaqueductal gray (PAG) receives collaterals of spinothalamic (pain‐conducting) fibers and projects to the raphe nuclei (particularly nucleus raphe magnus) in the RF. Neurons in these areas in turn send afferents to the superficial laminae of the posterior horn, suppressing the transmission of pain information received by spinothalamic neurons. Reticular formation Brainstem (II) Reticular formation Functions of reticular formation Controls the arousal and consciousness Neurons in RF collect multiple sensory information and send ascending projections to diencephalon and telencephalon. This pathway collaborates with monoamine‐containing reticular projections form the ascending reticular activating system (ARAS) that continuously send signals to sustain cerebral activities and consciousness. Brainstem (II) Reticular formation Functions of reticular formation Controls the arousal and consciousness The ascending reticular activating system also works together with histaminergic and cholinergic projections from hypothalamus and basal forebrain regulate the sleep‐awake cycle. Brainstem (II) Blood supply of the brainstem Vertebral‐basilar system Medulla oblongata Anterior spinal a. Posterior spinal a. Posterior inferior cerebellar a. Brs. Vertebral a. Pons Brs. Basilar a. Midbrain Brs. Basilar a. Superior cerebellar aa. Posterior cerebral aa. F E D C B A Brainstem (II) General arrangement of cranial nerve nuclei (SA) (VA) Alar plate Basal plate (VE) (SE) Brainstem (II) General Arrangement Brainstem (II) Cranial nerve nuclei General Arrangement General General somatic afferents (GSA) : from skin, skeletal muscles, joints and ligaments General visceral afferents (GVA) : from visceral organ and blood vessels General somatic efferents (GSE) : to skeletal muscles derived from myotomes General visceral efferents (GVE) : to smooth and cardiac muscles and glands Special Special somatic afferents (SSA) : from visual and auditory organs Special visceral afferents (SVA) : from visceral sense (taste and smell) organs Branchial Branchial efferents (BE) : to striated muscles derived from branchial arches Brainstem (II) Cranial nerve nuclei GSE (3, 4, 6, 12) Oculomotor (III) Trochlear (IV) Abducens (VI) Hypoglossal (XII) GVE (3, 7, 9, 10) Edinger‐Westphal n. (III) Sup. Salivatory nu. (VII) Inf. Salivatory nu. (IX) Dorsal nu. Vagus (X) BE (5, 7, 9, 10, 11) Trigeminal motor nu. (V) Facial motor nu. (VII) Nu. Ambiguus (IX, X) Accessory nu. (XI) General Arrangement Brainstem (II) Cranial nerve nuclei SSA (8) Cochlear and vestibular nuclei (VIII) GSA (5, 7, 9, 10) Main trigeminal sensory nucleus (V) Mesencephalic trigeminal nucleus (V) Spinal trigeminal nucleus (V, VII, IX, X) GVA/SVA (VA) (7, 9, 10) Nu. Solitary tract (VII, IX, X) General Arrangement Brainstem (II) Cranial nerve nuclei Brainstem (III) Cranial nerves and nuclei Somatic motor nerves (3, 4, 6, 12) ‐‐‐ GSE Oculomotor nerve (III) innervates four extra‐ocular muscles Trochlear nerve (IV) innervates superior oblique Abducens nerve (IV) innervates lateral rectus Hypoglossal nerve (XII) innervates tongue muscles Brainstem (III) Cranial nerves and nuclei Oculomotor nerve (III) Somatic motor nerves Brainstem (III) Cranial nerves and nuclei Somatic motor nerves Oculomotor nerve (III) Supplies : Levator palpebrae superioris Superior rectus Medial rectus, inferior oblique, inferior rectus Brainstem (III) Cranial nerves and nuclei Oculomotor nerve (III) Somatic motor nerves Brainstem (III) Cranial nerves and nuclei Trochlear nerve (IV) Somatic motor nerves Brainstem (III) Cranial nerves and nuclei Somatic motor nerves Trochlear nerve (IV) Supplies : Superior oblique (SO4) Brainstem (III) Cranial nerves and nuclei Trochlear nerve (IV) A unique cranial nerve that : exit from the dorsal surface of the brain all the lower motor neuron fibers decussate longest intracranial course smallest number of axons Somatic motor nerves Brainstem (III) Cranial nerves and nuclei Abducens nerve (VI) Somatic motor nerves Brainstem (III) Cranial nerves and nuclei Abducens nerve (VI) Supplies : Lateral rectus Somatic motor nerves Brainstem (III) Cranial nerves and nuclei Abducens nerve (VI) Somatic motor nerves Brainstem (III) Cranial nerves and nuclei Inter‐nuclear connection ‐‐ between abducens nucleus and oculomotor nucleus Somatic motor nerves Brainstem (III) Cranial nerves and nuclei Hypoglossal nerve (XII) Somatic motor nerves Brainstem (III) Cranial nerves and nuclei Somatic motor nerves Hypoglossal nerve (XII) Hypoglossal trigone Brainstem (III) Cranial nerves and nuclei Somatic motor nerves Hypoglossal nerve (XII) Supplies : intrinsic and most extrinsic tongue muscles of same side If damaged : Tongue is deviated toward the side of lesion. Bilateral lesions of hypoglossal nerve cause difficulties in eating & speaking
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