At fourth ventricle, above spinal cord, brain regions {brainstem}| can have cervical flexure and cephalic flexure and contain midbrain and hindbrain [Parvizi and Damasio, 2001] [Zeman, 2001]. Brainstem includes cranial nerve nuclei.
hindbrain
Hindbrain contains pons near midbrain and medulla oblongata near spinal cord.
midbrain
Midbrain has noradrenergic lateral reticular system, noradrenergic locus coeruleus, serotoninergic raphe nucleus, dopaminergic basal midbrain nuclei, cholinergic sense nuclei, and histaminergic nuclei.
Histaminergic nuclei project in net over brain.
Cholinergic sense nuclei connect to cerebral cortex. Damage reduces cerebral activity and causes a dreamy state.
functions
Brainstem integrates signals for attention, sex, and consciousness. Consciousness requires higher brainstem.
Regions {area postrema} can lack blood-brain barrier and can sense large molecules.
Forward brainstem {basal forebrain}| damage affects event-time recall.
Brainstem nucleus {cochlear nucleus} receives from cochlea hair cells.
Brainstem nucleus {lateral cervical nucleus} sends to superior colliculi.
Brainstem nucleus {nucleus sagulum} sends to superior colliculi.
Brainstem nucleus {perihypoglossal nucleus} sends motor input to superior colliculi.
In vertebrates other than mammals, structure {poker chip} decides which of twenty behavior modes is most appropriate. Poker chip later evolves to become reticular formation.
Brainstem nucleus {posterior commissure nucleus} sends motor input to superior colliculi.
Brainstem nucleus {prepositus hypoglossius} receives from superior colliculi and sends to oculomotor system.
Mesencephalic reticular formation, intralaminar nuclei, and reticular nuclei {reticular formation}| {reticular activating system} {reticulum} {ascending reticular activating system} {extralemniscal system} {non-specific afferent system} {gating system} {ascending activation system} {midbrain reticular formation} {mesencephalic reticular formation} stimulate thalamus and cortex to cause waking and sleep states.
purposes
Reticular formation arouses, integrates signals, maintains consciousness, controls vital functions, modulates perception, forms and recalls memories, and coordinates motor behaviors.
purposes: consciousness
Consciousness involves thalamus reticular-activating-system ascending fibers.
damage
Damage to reticular formation causes coma, memory disorganization, sleep, reduced cortex energy, similar reactions to strong and weak stimuli, and poor behavior control.
electrical stimulation
Reticular-formation electrode stimulation can cause unpleasant feelings.
electrical stimulation: memory
Retention improves with reticular formation stimulation, which arouses brain. Stimulation does not affect retrieval. Stimulating other brain areas has no affect on retention.
biology: input
Reticular formation receives axons from sense pathways and cortex and has multisensory convergence sites. All senses activate ascending reticular formation, which mediates pain. Stimulating ascending reticular formation causes fear and avoidance behaviors.
biology: output
Interconnecting neurons with short axons run from lower brainstem to midbrain. Descending reticular formation acts on interneurons indirectly.
biology: chemicals
Serotonin affects reticular formation and attention system to synchronize cortex. Noradrenaline desynchronizes cortex.
biology: columns
Reticular formation has medial, median, and lateral columns, from anterior midbrain through pons, medulla, and spinal cord [Hobson, 1989] [Hunter and Jasper, 1949] [Magoun, 1952] [Moruzzi and Magoun, 1949] [Steriade and McCarley, 1990].
Medial column receives pyramidal tract, cerebellum, and sense axons from cortex and sends by ascending reticular activating system to intralaminar thalamic nuclei, which send to striatum and cortex, to activate cortex and control waking and sleeping.
Raphe nucleus median column, mainly dorsal raphe nucleus, sends inhibition to limbic system in median column.
Lateral reticular system for attention projects to spinal cord, hypothalamus, and brainstem lateral-column tractus-solitarius nucleus. Noradrenaline locus coeruleus lateral column sends attention information to limbic system and prefrontal lobes.
biology: evolution
Reticular formation is only in mammals but evolved from something similar in lower animals.
Brainstem nucleus {solitary tract nucleus} {tractus solitarius nucleus} receives from locus coeruleus, trigeminal nucleus, and vagus nerve and sends to parabrachial nucleus, which receives from GI tract, and ventral medial basal thalamus. Solitary tract nucleus, with taste cortex and thalamus, is for taste preferences and can detect nausea. NTS is satiation region and receives gut peptide cholecystokinin (CCK).
Brainstem nucleus {trigeminal nucleus} receives sensory C fibers and A-delta fibers caudally from spinal cord posterior horn lamina I and sends to ventral medial posterior thalamus, brainstem nucleus tractus solitarius, and brainstem parabrachial nucleus.
Brainstem nuclei {vestibular nucleus} can be for balance.
Brainstem regions {zona incerta} can send motor input to superior colliculi.
Ascending reticular-activating-system (ARAS) lowest-hindbrain component {sensory reticular formation} receives visceral, somatic, auditory, and visual axons from ascending sense axons and sends to neocortex through hypothalamus and thalamus. It has pain center and wakefulness or alertness center.
Spinal-cord brainstem bulb {medulla, brain}| {medulla oblongata} includes basal ganglia and continues major nerve tracts. It relays auditory nerve sense and motor nerves, mediating phonation and articulation. It regulates cardiac action, chewing, tasting, swallowing, coughing, sneezing, salivation, vomiting, and sucking in newborns. Respiratory center maintains respiration. Some medulla-oblongata neurons make epinephrine.
A limbic-system part {amygdala}| includes insula white matter.
location
Insula is in posterior frontal lobe and anterior temporal lobe.
input
Lateral amygdala receives sensations slowly from sensory cortex and fast from thalamus, and receives memories from medial temporal lobe. Central amygdala receives from lateral amygdala, prefrontal cortex, and basal amygdala. Basal amygdala receives from lateral amygdala, medial temporal lobe, and prefrontal cortex.
output
Amygdala dopamine neurons connect to cholinergic neurons in medial septal nucleus, nucleus accumbens, nucleus basalis magnocellularis, nucleus of diagonal band of Broca, hypothalamus regions for motivation and reward, and sense and motor cerebral cortex upper layers.
Amygdala sends to orbitofrontal prefrontal cortex, mediodorsal thalamic nucleus, and hippocampal formation.
Lateral amygdala sends to central amygdala. Central amygdala sends to lateral hypothalamus for blood pressure, paraventricular hypothalamus for hormones, motor cortex for stopping, and basal amygdala. Basal amygdala sends to central amygdala.
functions
Amygdala compares new stimulus to previous stimuli and signals differences to other brain regions. Using memory, amygdala participates in habituation and anticipation.
Amygdala {basolateral nucleus} affects aggression, dominance, submission, and territoriality behaviors. Amygdala regulates fear and emotional behavior. Amygdala regulates visceral activity. Amygdala affects vision and smell.
damage
Removal of, or injury to, amygdala does not affect memory.
drug
Cocaine affects sublenticular extended amygdala.
Medulla ganglia {basal ganglia}| include amygdala, caudate nucleus, claustrum, external-capsule fibers, globus pallidus, internal-capsule fibers, lentiform nucleus, nucleus basalis of Meynert, nucleus dorsalis, putamen, septal nuclei, substantia nigra pars reticulata, and subthalamic nucleus.
input
Basal ganglia receive from basal-midbrain-nuclei dopaminergic neurons.
output
Basal-ganglia cholinergic neurons send to motor cortex for transmission to muscles [Langston and Palfreman, 1995].
functions
Basal ganglia assemble, select, and trigger automatic movements, perceptual motor coordination, ballistic movements, and proprioceptively controlled movements, using movement plans. They track moving visual objects, control eye movements, and process visual and multisensory data. They control tremor and muscle tone. Basal ganglia coordinate with neocortex and cerebellum for posture and complex voluntary movements.
Medulla basal ganglia {caudate nucleus} can inhibit globus pallidus. Caudate nucleus receives excitatory input from cerebral cortex and inhibitory input from thalamus, substantia nigra, and raphe. Caudate nucleus is for memory and obsessive behavior.
Medulla basal ganglia {claustrum} can lie under cerebral cortex near insula and project to many cortex regions.
Putamen and globus pallidus {lenticular nuclei} look striated because they have myelinated tracts.
Brainstem regions {motor reticular formation} can facilitate spinal mediated reflexes and transmit feedback from higher centers to primary receptors.
Basal ganglia nuclei {nucleus basalis of Meynert} {nuclei of Meynert} {Meynert nuclei} {substantia innominata} can have cholinergic neurons and send to cerebral cortex.
Basal ganglia nucleus {nucleus dorsalis} receives proprioception input from spinal cord.
Brainstem regions {globus pallidus} {pallidum} can receive from red nucleus and inhibit thalamus and subthalamic nucleus. Dopamine neurons can cause rigidity if overstimulated. Choline neurons can cause hyperkinesis, chorea, and athetosis if overstimulated.
Basal ganglia connect to thalamus, then to cortex, then back to basal ganglia {polysynaptic loop}.
Medulla basal ganglia {putamen} can receive excitatory input from cerebral cortex and inhibitory input from thalamus, substantia nigra, and raphé nucleus. It inhibits globus pallidus. Putamen is for memory, motor skill, and obsessiveness. It has nearby-space maps, used in motor control.
Brainstem regions {raphé dorsalis} can send motor input to superior colliculi.
Medulla nuclei {raphé nuclei} can secrete serotonin, make peptide substance P, start light sleep, and modulate pain, using spinal-cord dorsal-horn presynaptic inhibition.
Forebrain basal ganglia {septal nuclei} {septum, medulla} can receive from hippocampus and reticular formation and send to hippocampus, hypothalamus, and midbrain. Septal nuclei have trophotropic centers. They can control aggressiveness. They organize sexual thoughts, emotions, and action. They are in or near region that causes pleasure when excited.
Putamen, globus pallidus, and caudate nucleus {corpus striatum} {striatum} are near thalamus. They look striated because they have myelinated tracts.
no layers
Corpus striatum neuron types mix but not in layers.
maps
Most maps in mammalian cortex connect to maps in corpus striatum.
functions
Corpus striatum integrates learned automatic movement sequences, such as voluntary eye movements.
input
Some striatum neurons receive from thousands of cortical neurons that send 10-Hz to 40-Hz oscillating signals {interval timer}. Stimuli synchronize oscillations. Oscillators then go on oscillating. Second stimuli make substantia nigra send dopamine to striatum. Striatum remembers signal pattern. If starting signal repeats, dopamine repeats.
output
If pattern matches, striatum signals to thalamus, which informs cortex.
Medulla regions {respiratory center} can send excitatory signals along phrenic nerve to diaphragm.
Nuclei {subthalamic nucleus} {Luys nucleus} {Luys body} {nucleus of Luys} {body of Luys} can be near hypothalamus, inhibit globus pallidus, and send to thalamus. Damage causes ballistic movement.
Fiber bridge {pons}| from hindbrain side to opposite cerebellum side holds major nerve tracts connecting cerebellum and cortex, in both directions, and connects thalamus to olive. Cerebral cortex motor regions influence pons. Pons controls heart, lungs, eye movements, muscle tone, walking, and running. It mediates protective and orientation reflexes.
A pons region {locus coeruleus}| can receive feedback from sense cortex and send to spinal cord, hypothalamus, tractus solitarius nucleus, sensory cerebral cortex, and cerebellum Purkinje cells [Foote and Morrison, 1987] [Foote et al., 1980] [Hobson, 1999]. Locus coeruleus contains few thousand neurons and is largest noradrenaline nucleus.
transmitters
Locus-coeruleus neurons contain neuropeptide Y (NPY) and galanin peptide transmitters.
functions
Locus coeruleus suppresses tonic vegetative regions. It regulates attention, pleasure, energy, motivation, and arousal. It causes deep sleep. REM sleep, cataplexy, grooming, and feeding depress it. Interruptions and multimodal somatosensory stimuli, including pain, excite it. Locus-coeruleus electrical stimulation causes fear and anxiety.
A pons region {pneumotaxic center} receives from nerves that sense alveoli stretching and inhibits breathing.
A pons regions {tegmentum} can include reticular formation and be for attention.
Midbrain ganglia {cuneate nuclei} can receive texture, form, and vibration information in medulla ipsilateral cuneate tracts and send to thalamus, cerebrum, and cerebellum. Maps are smaller than in other areas.
Midbrain nuclei {gracile nuclei} can receive texture, form, and vibration information in medulla ipsilateral gracile tracts and send to thalamus, cerebrum, and cerebellum. Maps are smaller than in other areas.
Midbrain nuclei {inferior colliculi} can send to superior colliculi. They have auditory functions and control eye movements.
Brainstem regions {inferior olive} (IO) can send to cerebellar Purkinje cells.
Midbrain striatum nucleus {nucleus accumbens} receives excitatory dopaminergic pathway from frontal lobes and ventral tegmentum and receives from basal and lateral amygdala. It sends to ventral pallidum. Nucleus accumbens mediates emotions and movements. Benzodiazepine anti-anxiety agent and antipsychotic agents block dopaminergic pathway activation. Regular drug use and other reward stimuli increase delta FosB transcription factor, which causes sensitization and degrades slowly, in nucleus accumbens.
Vertebrates other than mammals have vision cell layer {optic tectum} over large ventricle. Mammals have superior colliculi instead. In amphibia and fish, fibers from retina to tectum keep growing and changing. Mostly unimodal sense pathways go from cerebral cortex to tectum. Tectum makes eye saccades to focus attention-getting object on fovea. Divergence in individual tectum-nuclei loops explains how large recruited tectum-neuron population can form composite command observed at tectum.
Posterior upper brainstem region {parabrachial nuclei} (PBN) receives from sensory trigeminal nucleus and nucleus tractus solitarius and sends motor input to superior colliculi and motor output to hypothalamus and ventral medial basal thalamus.
Pons and thalamus nucleus {periaqueductal gray} (PAG) has opiate receptors and makes endorphins.
Brainstem nucleus {periolivary nucleus} near olive sends to superior colliculi.
Brainstem regions {posterior upper brainstem} can contain periaqueductal gray, parabrachial nucleus, monoamine nuclei, and acetylcholine nuclei. Damage to posterior upper brainstem causes coma.
Brainstem regions {quadrigeminal plate} can have superior and inferior colliculi.
Brainstem nucleus {red nucleus} {ruber nucleus} receives from amygdala and sends to hypothalamus paraventricular nucleus {stria terminalis}. It works similarly to motor cortex.
Brainstem regions {substantia nigra} {substantia nigra pars reticulata} can send to superior colliculus to control eye movement. Substantia nigra cholinergic neurons connect to sense and motor neurons in caudate nucleus and putamen in basal ganglia. Basal ganglia and other midbrain dopamine neurons inhibit caudate nucleus, corpus striatum, and thalamus to coordinate motor function and automatic movement. Alzheimer's disease degenerates substantia nigra neurons.
Mammal midbrain dorsal surface has large symmetrical bumps {superior colliculi} that mediate light accommodation, eyeball movements, body movements for vision, orientation, and attention [Aldrich et al., 1987] [Brindley et al., 1969] [Celesia et al., 1991].
anatomy
Superior colliculus has seven alternating cellular and fibrous layers with few interneurons, eight types of synaptic terminals, and broad dendrite arbors. Superficial layers I to III and deep layers IV to VII have topographic motor maps and associated visual and touch maps.
Superior colliculus removal causes failure to detect contralateral visual stimuli.
anatomy: input
Superior colliculus efferent neurons for eye movements receive input from substantia nigra.
Superior colliculus deep layers receive vision information ipsilaterally from lateral suprasylvian visual area and anterior ectosylvian visual area, not from striate visual cortex.
Deep layers receive somatosensory input from anterior ectosylvian sulcus dorsal part, contralateral sensory trigeminal complex, dorsal column nuclei, lateral cervical nucleus, and spinal cord. Contralateral sensory trigeminal complex receives C fibers and A-delta fibers.
Deep layers receive auditory input from anterior ectosylvian sulcus Field AES region, inferior colliculus contralateral brachium, inferior colliculus external nucleus, nucleus sagulum, and dorsomedial periolivary nucleus.
Deep layers receive motor input from frontal eye fields, motor cortex, zona incerta, thalamus reticular nucleus, posterior commissure nucleus, perihypoglossal nucleus, contralateral superior colliculus, locus coeruleus, raphé dorsalis, parabrachial nuclei, reticular formation, and hypothalamus.
Deep layers receive from basal ganglia through substantia nigra pars reticulata. Deep layers receive from cerebellum deep nuclei, including medial and posterior interposed nuclei.
anatomy: output
Superior colliculus deep layers send to thalamus, opposite superior colliculus, brainstem, and spinal cord. Superior colliculus deep layers connect to sense and motor cerebral cortex and to brainstem and spinal cord, to position peripheral sense organs. Deep layers also send contralaterally to tegmentum and spinal cord to reposition eyes, head, limbs, ears, and whiskers.
neurons: receptive field
Superior-colliculus neurons have central ON zones surrounded by lower sensitivity areas, not like retina and lateral-geniculate-nucleus ON-center-neuron or OFF-center-neuron receptive fields. Receptive fields are larger than in lateral geniculate or cortex neurons. Border is inhibitory {suppressive zone}. The most-effective stimulus is smaller than receptive field. Moving or flashing stimuli are more effective than stationary ones. Movement direction is more effective. Slow movements are more effective than rapid ones. Repeating same stimulus produces response habituation.
neurons: noxious
Superior colliculus neurons {nociceptive-specific neuron} (NS) can respond to noxious stimuli. Superior colliculus neurons {wide dynamic range neuron} (WDR) can respond to all mechanical stimuli, but especially to noxious mechanical or thermal stimuli.
neurons: multisensory
Superior colliculus neurons are 25% unimodal and 75% multisensory. Multisensory and unimodal neurons typically require 100 milliseconds to process information, but some multisensory neurons take 1500 milliseconds.
neurons: auditory
Superior colliculus has four auditory neuron types. Compared to cortical auditory neurons, superior colliculus auditory neurons are more insensitive to pure tones and more sensitive to spatial location, interaural time, and intensity differences. They respond better to moving stimuli, have directional selectivity, habituate to repeated stimuli, and have restricted receptive fields with maximal-response regions.
neurons: somatosensory
Superior colliculus somatosensory neurons respond to hair or skin stimulation, have well-defined receptive fields, prefer intermediate-velocity or high-velocity stimuli, habituate rapidly, are large, have best regions, have no inhibitory surrounding areas, and have no directional selectivity.
neurons: movement field
Midbrain neuron receptive fields {movement field} are like sense-neuron receptive fields. Neurons with similar movement fields are in same superior colliculus region. If neuron activity exceeds threshold, amount above threshold determines saccade movement velocity and distance.
eye movement
Mammal superior colliculi and non-mammal optic tectum process multisensory information, shift attention, and control voluntary and involuntary eye and other sense-organ movements, for orientation and attention. Stimulation shifts eyes, ears, and head to focus on stimulus location. High intensity causes withdrawal or escape.
Anteromedial superior colliculi stimulation causes contralateral, upward, and parallel conjugate eye movement.
Lateral superior colliculi stimulation causes conjugate, contralateral, and downward movement.
To initiate eye movement to periphery, caudal superior colliculus, which represents peripheral visual space, has pre-motor activity.
Visual fixation involves neurons in rostral superior colliculus.
eye movement: saccade
Superior colliculus neurons {motor error neuron} can generate low-frequency, long-duration discharge to signal difference between current eye position and target position. Superior colliculus neurons can initiate saccades and determine speed, direction, and amplitude [Corbetta, 1998] [Schall, 1991] [Schiller and Chou, 1998]. Saccade initiation and velocity, duration, and direction specification are separate processes. Saccade commands are many-neuron vector sums.
Brainstem regions {superior olive} can measure time and intensity differences to differentiate auditory-signal arrival times. Olive sides {lateral superior olive} (LSO) receive inputs from both ears for intensity-level-difference detection. Olive middle {medial superior olive} (MSO) receives inputs from both ears, for time-difference detection. Neurons have time-difference ranges.
Brainstem region {ventral tegmentum} {ventral tegmental area} (VTA) is for pleasure and motivation. Dopamine neurons inhibit nucleus accumbens, mesolimbic system, frontal cortex, and sensorimotor cortex.
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Date Modified: 2022.0225