Reticular formation

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Reticular formation
Gray701.png
Coronal section of the pons, at its upper part. [1] (Formatio reticularis labeled at left.)
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Traverse section of the medulla oblongata at about the middle of the olive. (Formatio reticularis grisea and formatio reticularis alba labeled at left.)
Details
Location Brainstem, hypothalamus and other regions
Identifiers
Latin formatio reticularis
MeSH D012154
NeuroNames 1223
NeuroLex ID nlx_143558
TA98 A14.1.00.021
A14.1.05.403
A14.1.06.327
TA2 5367
FMA 77719
Anatomical terms of neuroanatomy

The reticular formation is a set of interconnected nuclei in the brainstem that spans from the lower end of the medulla oblongata to the upper end of the midbrain. [2] The neurons of the reticular formation make up a complex set of neural networks in the core of the brainstem. [3] The reticular formation is made up of a diffuse net-like formation of reticular nuclei which is not well-defined. [4] It may be seen as being made up of all the interspersed cells in the brainstem between the more compact and named structures. [4]

Contents

The reticular formation is functionally divided into the ascending reticular activating system (ARAS), ascending pathways to the cerebral cortex, and the descending reticular system, descending pathways (reticulospinal tracts) to the spinal cord. [5] [6] [7] [8] Due to its extent along the brainstem it may be divided into different areas such as the midbrain reticular formation, the central mesencephalic reticular formation, the pontine reticular formation, the paramedian pontine reticular formation, the dorsolateral pontine reticular formation, and the medullary reticular formation. [9]

Neurons of the ARAS basically act as an on/off switch to the cerebral cortex and hence play a crucial role in regulating wakefulness; behavioral arousal and consciousness are functionally related in the reticular formation using a number of neurotransmitter arousal systems. The overall functions of the reticular formation are modulatory and premotor, [upper-alpha 1] involving somatic motor control, cardiovascular control, pain modulation, sleep and consciousness, and habituation. [10] The modulatory functions are primarily found in the rostral sector of the reticular formation and the premotor functions are localized in the neurons in more caudal regions.

The reticular formation is divided into three columns: raphe nuclei (median), gigantocellular reticular nuclei (medial zone), and parvocellular reticular nuclei (lateral zone). The raphe nuclei are the place of synthesis of the neurotransmitter serotonin, which plays an important role in mood regulation. The gigantocellular nuclei are involved in motor coordination. The parvocellular nuclei regulate exhalation. [11]

The reticular formation is essential for governing some of the basic functions of higher organisms. It is phylogenetically old and found in lower vertebrates. [2]

Structure

A cross section of the lower part of the pons showing the pontine reticular formation labeled as #9 Lower pons horizontal KB.svg
A cross section of the lower part of the pons showing the pontine reticular formation labeled as #9

The human reticular formation is composed of almost 100 nuclei and contains many projections into the forebrain, brainstem, and cerebellum, among other regions. [6] It includes the reticular nuclei, reticulothalamic projection fibers, diffuse thalamocortical projections, ascending cholinergic projections, descending non-cholinergic projections, and descending reticulospinal projections. [7] The reticular formation also contains two major neural subsystems, the ascending reticular activating system and descending reticulospinal tracts, which mediate distinct cognitive and physiological processes. [6] [7] It has been functionally cleaved both sagittally and coronally.

Traditionally the reticular nuclei are divided into three columns:[ citation needed ]

The original functional differentiation was a division of caudal and rostral. This was based upon the observation that the lesioning of the rostral reticular formation induces a hypersomnia in the cat brain. In contrast, lesioning of the more caudal portion of the reticular formation produces insomnia in cats. This study has led to the idea that the caudal portion inhibits the rostral portion of the reticular formation.[ citation needed ]

Sagittal division reveals more morphological distinctions. The raphe nuclei form a ridge in the middle of the reticular formation, and, directly to its periphery, there is a division called the medial reticular formation. The medial RF is large and has long ascending and descending fibers, and is surrounded by the lateral reticular formation. The lateral RF is close to the motor nuclei of the cranial nerves, and mostly mediates their function.[ citation needed ]

Medial and lateral reticular formation

The medial reticular formation and lateral reticular formation are two columns of nuclei with ill-defined boundaries that send projections through the medulla and into the midbrain. The nuclei can be differentiated by function, cell type, and projections of efferent or afferent nerves. Moving caudally from the rostral midbrain, at the site of the rostral pons and the midbrain, the medial RF becomes less prominent, and the lateral RF becomes more prominent.[ citation needed ]

Existing on the sides of the medial reticular formation is its lateral cousin, which is particularly pronounced in the rostral medulla and caudal pons. Out from this area spring the cranial nerves, including the very important vagus nerve.[ clarification needed ] The lateral RF is known for its ganglions and areas of interneurons around the cranial nerves, which serve to mediate their characteristic reflexes and functions.

Major subsystems

The subsystems of the reticular formation are the ascending reticular activating system, and the descending reticular system. [7]

Ascending reticular activating system

Ascending reticular activating system. Reticular formation labeled near center. Gray690.png
Ascending reticular activating system. Reticular formation labeled near center.

The ascending reticular activating system (ARAS), also known as the extrathalamic control modulatory system or simply the reticular activating system (RAS), is a set of connected nuclei in the brains of vertebrates that is responsible for regulating wakefulness and sleep-wake transitions. The ARAS is in the midbrain reticular formation. [12] It is mostly composed of various nuclei in the thalamus/hypothalamus and a number of dopaminergic, noradrenergic, serotonergic, histaminergic, cholinergic, and glutamatergic brain nuclei. [6] [13] [14] [15]

Structure

The ARAS is composed of several neural circuits connecting the dorsal part of the posterior midbrain and the ventral pons to the cerebral cortex via distinct pathways that project through the thalamus and hypothalamus. [6] [14] [15] The ARAS is a collection of different nuclei – more than 20 on each side in the upper brainstem, the pons, medulla, and posterior hypothalamus. [12] The neurotransmitters that these neurons release include dopamine, norepinephrine, serotonin, histamine, acetylcholine, and glutamate. [6] [13] [14] [15] They exert cortical influence through direct axonal projections and indirect projections through thalamic relays. [14] [15] [12]

The thalamic pathway consists primarily of cholinergic neurons in the pontine tegmentum, whereas the hypothalamic pathway is composed primarily of neurons that release monoamine neurotransmitters, namely dopamine, norepinephrine, serotonin, and histamine. [6] [13] The glutamate-releasing neurons in the ARAS were identified much more recently relative to the monoaminergic and cholinergic nuclei; [16] the glutamatergic component of the ARAS includes one nucleus in the hypothalamus and various brainstem nuclei. [14] [16] [17] The orexin neurons of the lateral hypothalamus innervate every component of the ascending reticular activating system and coordinate activity within the entire system. [15] [18] [19]

Key components of the ascending reticular activating system
Nucleus typeCorresponding nuclei that mediate arousalSources
Dopaminergic  nuclei [6] [13] [14] [15]
Noradrenergic  nuclei [6] [13] [15]
Serotonergic  nuclei [6] [13] [15]
Histaminergic  nuclei [6] [13] [20]
Cholinergic  nuclei [6] [14] [15] [16]
Glutamatergic  nuclei [14] [15] [16] [17] [20] [21]
Thalamic  nuclei [6] [14] [22]

The ARAS consists of evolutionarily ancient areas of the brain, which are crucial to the animal's survival and protected during adverse periods, such as during inhibitory periods of animal hypnosis also known as Totstellreflex. [23] The ascending reticular activating system which sends neuromodulatory projections to the cortex - mainly connects to the prefrontal cortex. [24] There seems to be low connectivity to the motor areas of the cortex. [24]

Function

Consciousness

The ascending reticular activating system is an important enabling factor for the state of consciousness. [12] The ascending system is seen to contribute to wakefulness as characterised by cortical and behavioural arousal. [8]

Regulating sleep-wake transitions

The main function of the ARAS is to modify and potentiate thalamic and cortical function such that electroencephalogram (EEG) desynchronization ensues. [upper-alpha 2] [26] [27] There are distinct differences in the brain's electrical activity during periods of wakefulness and sleep: Low voltage fast burst brain waves (EEG desynchronization) are associated with wakefulness and REM sleep (which are electrophysiologically similar); high voltage slow waves are found during non-REM sleep. Generally speaking, when thalamic relay neurons are in burst mode the EEG is synchronized and when they are in tonic mode it is desynchronized. [27] Stimulation of the ARAS produces EEG desynchronization by suppressing slow cortical waves (0.3–1 Hz), delta waves (1–4 Hz), and spindle wave oscillations (11–14 Hz) and by promoting gamma band (20–40 Hz) oscillations. [18]

The physiological change from a state of deep sleep to wakefulness is reversible and mediated by the ARAS. [28] The ventrolateral preoptic nucleus (VLPO) of the hypothalamus inhibits the neural circuits responsible for the awake state, and VLPO activation contributes to the sleep onset. [29] During sleep, neurons in the ARAS will have a much lower firing rate; conversely, they will have a higher activity level during the waking state. [30] In order that the brain may sleep, there must be a reduction in ascending afferent activity reaching the cortex by suppression of the ARAS. [28] Dysfunction of the paraventricular nucleus of the hypothalamus can lead to drowsiness for up to 20 hours per day. [31]

Attention

The ARAS also helps mediate transitions from relaxed wakefulness to periods of high attention. [22] There is increased regional blood flow (presumably indicating an increased measure of neuronal activity) in the midbrain reticular formation (MRF) and thalamic intralaminar nuclei during tasks requiring increased alertness and attention.[ citation needed ]

Clinical significance of the ARAS

Mass lesions in the ARAS nuclei can cause a loss of consciousness. [12] [32] Bilateral damage to the ARAS nuclei may lead to coma or death. [33]

Direct electrical stimulation of the ARAS produces pain responses in cats and elicits verbal reports of pain in humans.[ citation needed ] Ascending reticular activation in cats can produce mydriasis, [34] which can result from prolonged pain. These results suggest some relationship between ARAS circuits and physiological pain pathways. [34]

Some pathologies of the ARAS may be attributed to ageing, as there appears to be a general decline in reactivity of the ARAS with advancing years. [35] Changes in electrical coupling [upper-alpha 3] have been suggested to account for some changes in ARAS activity: if coupling were down-regulated, there would be a corresponding decrease in higher-frequency synchronization (gamma band). Conversely, up-regulated electrical coupling would increase synchronization of fast rhythms that could lead to increased arousal and REM sleep drive. [37] Specifically, disruption of the ARAS has been implicated in the following disorders:

Developmental influences

There are several potential factors that may adversely influence the development of the ascending reticular activating system:

Descending reticulospinal system

The reticulospinal tracts, are extrapyramidal motor tracts that descend from the reticular formation [42] in two tracts to act on the motor neurons supplying the trunk and proximal limb flexors and extensors. The reticulospinal tracts are involved mainly in locomotion and postural control, although they do have other functions as well. [43] The descending reticulospinal tracts are one of four major cortical pathways to the spinal cord for musculoskeletal activity. The reticulospinal tracts work with the other three pathways to give a coordinated control of movement, including delicate manipulations. [42] The four pathways can be grouped into two main system pathways – a medial system and a lateral system. The medial system includes the reticulospinal tract and the vestibulospinal tract, and provides control of posture. The corticospinal tract and the rubrospinal tract belong to the lateral system which provides fine control of movement. [42]

Spinal cord tracts - reticulospinal tract labeled in red, near-center at left in figure Spinal cord tracts - English.svg
Spinal cord tracts - reticulospinal tract labeled in red, near-center at left in figure

The reticulospinal tracts are the medial reticulospinal tract, and the lateral reticulospinal tract.[ citation needed ]

The ascending sensory tract conveying information in the opposite direction is the spinoreticular tract.

Function

  1. Integrates information from the motor systems to coordinate automatic movements of locomotion and posture
  2. Facilitates and inhibits voluntary movement; influences muscle tone
  3. Mediates autonomic functions
  4. Modulates pain impulses
  5. Influences blood flow to lateral geniculate nucleus of the thalamus. [44]

Clinical significance

The reticulospinal tracts provide a pathway by which the hypothalamus can control sympathetic thoracolumbar outflow and parasympathetic sacral outflow.[ citation needed ]

Two major descending systems carrying signals from the brainstem and cerebellum to the spinal cord can trigger automatic postural response for balance and orientation: vestibulospinal tracts from the vestibular nuclei and reticulospinal tracts from the pons and medulla. Lesions of these tracts result in profound ataxia and postural instability. [45]

Physical or vascular damage to the brainstem disconnecting the red nucleus (midbrain) and the vestibular nuclei (pons) may cause decerebrate rigidity, which has the neurological sign of increased muscle tone and hyperactive stretch reflexes. Responding to a startling or painful stimulus, both arms and legs extend and turn internally. The cause is the tonic activity of lateral vestibulospinal and reticulospinal tracts stimulating extensor motoneurons without the inhibitions from rubrospinal tract. [46]

Brainstem damage above the red nucleus level may cause decorticate rigidity. Responding to a startling or painful stimulus, the arms flex and the legs extend. The cause is the red nucleus, via the rubrospinal tract, counteracting the extensor motorneuron's excitation from the lateral vestibulospinal and reticulospinal tracts. Because the rubrospinal tract only extends to the cervical spinal cord, it mostly acts on the arms by exciting the flexor muscles and inhibiting the extensors, rather than the legs. [46]

Damage to the medulla below the vestibular nuclei may cause flaccid paralysis, hypotonia, loss of respiratory drive, and quadriplegia. There are no reflexes resembling early stages of spinal shock because of complete loss of activity in the motorneurons, as there is no longer any tonic activity arising from the lateral vestibulospinal and reticulospinal tracts. [46]

History

The term "reticular formation" was coined in the late 19th century by Otto Deiters, coinciding with Ramon y Cajal's neuron doctrine . Allan Hobson states in his book The Reticular Formation Revisited that the name is an etymological vestige from the fallen era of the aggregate field theory in the neural sciences. The term "reticulum" means "netlike structure", which is what the reticular formation resembles at first glance. It has been described as being either too complex to study or an undifferentiated part of the brain with no organization at all. Eric Kandel describes the reticular formation as being organized in a similar manner to the intermediate gray matter of the spinal cord. This chaotic, loose, and intricate form of organization is what has turned off many researchers from looking farther into this particular area of the brain.[ citation needed ] The cells lack clear ganglionic boundaries, but do have clear functional organization and distinct cell types. The term "reticular formation" is seldom used anymore except to speak in generalities. Modern scientists usually refer to the individual nuclei that compose the reticular formation.[ citation needed ]

Moruzzi and Magoun first investigated the neural components regulating the brain's sleep-wake mechanisms in 1949. Physiologists had proposed that some structure deep within the brain controlled mental wakefulness and alertness. [26] It had been thought that wakefulness depended only on the direct reception of afferent (sensory) stimuli at the cerebral cortex.[ citation needed ]

As direct electrical stimulation of the brain could simulate electrocortical relays, Magoun used this principle to demonstrate, on two separate areas of the brainstem of a cat, how to produce wakefulness from sleep. He first stimulated the ascending somatic and auditory paths; second, a series of "ascending relays from the reticular formation of the lower brain stem through the midbrain tegmentum, subthalamus and hypothalamus to the internal capsule." [47] The latter was of particular interest, as this series of relays did not correspond to any known anatomical pathways for the wakefulness signal transduction and was coined the ascending reticular activating system (ARAS).[ citation needed ]

Next, the significance of this newly identified relay system was evaluated by placing lesions in the medial and lateral portions of the front of the midbrain. Cats with mesencephalic interruptions to the ARAS entered into a deep sleep and displayed corresponding brain waves. In alternative fashion, cats with similarly placed interruptions to ascending auditory and somatic pathways exhibited normal sleeping and wakefulness, and could be awakened with physical stimuli. Because these external stimuli would be blocked on their way to the cortex by the interruptions, this indicated that the ascending transmission must travel through the newly discovered ARAS.[ citation needed ]

Finally, Magoun recorded potentials within the medial portion of the brain stem and discovered that auditory stimuli directly fired portions of the reticular activating system. Furthermore, single-shock stimulation of the sciatic nerve also activated the medial reticular formation, hypothalamus, and thalamus. Excitation of the ARAS did not depend on further signal propagation through the cerebellar circuits, as the same results were obtained following decerebellation and decortication. The researchers proposed that a column of cells surrounding the midbrain reticular formation received input from all the ascending tracts of the brain stem and relayed these afferents to the cortex and therefore regulated wakefulness. [47] [28]

See also

Footnotes

  1. ↑ premotor function as in integrating feedback sensory signals with commands from upper motor neurons and deep cerebellar nuclei, and organizing the efferent activities of lower visceral motor and some somatic motor neurons in the brainstem and spinal cord. [3]
  2. ↑ An EEG's electrode on the scalp measures the activity of a very large number of pyramidal neurons in the underlying brain region. Each neuron generates a small electrical field that changes over time. In the sleep state, the neurons activate at approximately the same time, and the EEG wave, representing the summation of the neurons' electrical fields, tend to be in phase and has higher amplitude, and hence it is "synchronized." In the waking state, they do not activate at the same time because of irregular or out-of-phase inputs, the EEG wave, representing the algebraic sum, will have a smaller amplitude, and hence "dysynchronized." [25]
  3. ↑ Electrical coupling is the passive flow of electric current from one cell into an adjacent cell through gap junctions, such as the cells in the heart muscle or the neurons with electrical synapses. Electrically coupled cells fire synchronously because generated currents in one cell rapidly spread to the other cells. [36]

Related Research Articles

<span class="mw-page-title-main">Brainstem</span> Posterior part of the brain, adjoining and structurally continuous

The brainstem is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is continuous with the thalamus of the diencephalon through the tentorial notch, and sometimes the diencephalon is included in the brainstem.

<span class="mw-page-title-main">Extrapyramidal system</span> Connection between brain and spinal cord

In anatomy, the extrapyramidal system is a part of the motor system network causing involuntary actions. The system is called extrapyramidal to distinguish it from the tracts of the motor cortex that reach their targets by traveling through the pyramids of the medulla. The pyramidal tracts may directly innervate motor neurons of the spinal cord or brainstem, whereas the extrapyramidal system centers on the modulation and regulation of anterior (ventral) horn cells.

<span class="mw-page-title-main">Midbrain</span> Forward-most portion of the brainstem

The midbrain or mesencephalon is the rostral-most portion of the brainstem connecting the diencephalon and cerebrum with the pons. It consists of the cerebral peduncles, tegmentum, and tectum.

<span class="mw-page-title-main">Raphe nuclei</span> Moderate-size cluster of nuclei found in brain stem

The raphe nuclei are a moderate-size cluster of nuclei found in the brain stem. They have 5-HT1 receptors which are coupled with Gi/Go-protein-inhibiting adenyl cyclase. They function as autoreceptors in the brain and decrease the release of serotonin. The anxiolytic drug Buspirone acts as partial agonist against these receptors. Selective serotonin reuptake inhibitor (SSRI) antidepressants are believed to act in these nuclei, as well as at their targets.

<span class="mw-page-title-main">Arousal</span> State of being awoken

Arousal is the physiological and psychological state of being awoken or of sense organs stimulated to a point of perception. It involves activation of the ascending reticular activating system (ARAS) in the brain, which mediates wakefulness, the autonomic nervous system, and the endocrine system, leading to increased heart rate and blood pressure and a condition of sensory alertness, desire, mobility, and reactivity.

<span class="mw-page-title-main">Spinothalamic tract</span> Sensory pathway from the skin to the thalamus

The spinothalamic tract is a nerve tract in the anterolateral system in the spinal cord. This tract is an ascending sensory pathway to the thalamus. From the ventral posterolateral nucleus in the thalamus, sensory information is relayed upward to the somatosensory cortex of the postcentral gyrus.

<span class="mw-page-title-main">Lateral lemniscus</span> Brain structure

The lateral lemniscus is a tract of axons in the brainstem that carries information about sound from the cochlear nucleus to various brainstem nuclei and ultimately the contralateral inferior colliculus of the midbrain. Three distinct, primarily inhibitory, cellular groups are located interspersed within these fibers, and are thus named the nuclei of the lateral lemniscus.

<span class="mw-page-title-main">Pretectal area</span> Structure in the midbrain which mediates responses to ambient light

In neuroanatomy, the pretectal area, or pretectum, is a midbrain structure composed of seven nuclei and comprises part of the subcortical visual system. Through reciprocal bilateral projections from the retina, it is involved primarily in mediating behavioral responses to acute changes in ambient light such as the pupillary light reflex, the optokinetic reflex, and temporary changes to the circadian rhythm. In addition to the pretectum's role in the visual system, the anterior pretectal nucleus has been found to mediate somatosensory and nociceptive information.

<span class="mw-page-title-main">Ventrolateral preoptic nucleus</span> Nucleus of the anterior hypothalamus

The ventrolateral preoptic nucleus (VLPO), also known as the intermediate nucleus of the preoptic area (IPA), is a small cluster of neurons situated in the anterior hypothalamus, sitting just above and to the side of the optic chiasm in the brain of humans and other animals. The brain's sleep-promoting nuclei, together with the ascending arousal system which includes components in the brainstem, hypothalamus and basal forebrain, are the interconnected neural systems which control states of arousal, sleep, and transitions between these two states. The VLPO is active during sleep, particularly during non-rapid eye movement sleep, and releases inhibitory neurotransmitters, mainly GABA and galanin, which inhibit neurons of the ascending arousal system that are involved in wakefulness and arousal. The VLPO is in turn innervated by neurons from several components of the ascending arousal system. The VLPO is activated by the endogenous sleep-promoting substances adenosine and prostaglandin D2. The VLPO is inhibited during wakefulness by the arousal-inducing neurotransmitters norepinephrine and acetylcholine. The role of the VLPO in sleep and wakefulness, and its association with sleep disorders – particularly insomnia and narcolepsy – is a growing area of neuroscience research.

The zona incerta (ZI) is a horizontally elongated small nucleus that separates the larger subthalamic nucleus from the thalamus. Its connections project extensively over the brain from the cerebral cortex down into the spinal cord.

The dorsal longitudinal fasciculus (DLF) is a nerve fiber tract extending from the hypothalamus rostrally to the spinal cord caudally. It containins both descending and ascending fibers. Its fibers form a distinct bundle in the midbrain.

<span class="mw-page-title-main">Tegmentum</span> Part of the brain stem

The tegmentum is a general area within the brainstem. The tegmentum is the ventral part of the midbrain and the tectum is the dorsal part of the midbrain. It is located between the ventricular system and distinctive basal or ventral structures at each level. It forms the floor of the midbrain (mesencephalon) whereas the tectum forms the ceiling. It is a multisynaptic network of neurons that is involved in many subconscious homeostatic and reflexive pathways. It is a motor center that relays inhibitory signals to the thalamus and basal nuclei preventing unwanted body movement.

The amygdalofugal pathway is one of the three major efferent pathways of the amygdala, meaning that it is one of the three principal pathways by which fibers leave the amygdala. It leads from the basolateral nucleus and central nucleus of the amygdala. The amygdala is a limbic structure in the medial temporal lobe of the brain. The other main efferent pathways from the amygdala are the stria terminalis and anterior commissure.

<span class="mw-page-title-main">Lateral hypothalamus</span>

The lateral hypothalamus (LH), also called the lateral hypothalamic area (LHA), contains the primary orexinergic nucleus within the hypothalamus that widely projects throughout the nervous system; this system of neurons mediates an array of cognitive and physical processes, such as promoting feeding behavior and arousal, reducing pain perception, and regulating body temperature, digestive functions, and blood pressure, among many others. Clinically significant disorders that involve dysfunctions of the orexinergic projection system include narcolepsy, motility disorders or functional gastrointestinal disorders involving visceral hypersensitivity, and eating disorders.

The spinoreticular tract is a partially decussating (crossed-over) four-neuron sensory pathway of the central nervous system. The tract transmits slow nociceptive/pain information from the spinal cord to reticular formation which in turn relays the information to the thalamus via reticulothalamic fibers as well as to other parts of the brain. Most (85%) second-order axons arising from sensory C first-order fibers ascend in the spinoreticular tract - it is consequently responsible for transmitting "slow", dull, poorly-localised pain. By projecting to the reticular activating system (RAS), the tract also mediates arousal/alertness in response to noxious (harmful) stimuli. The tract is phylogenetically older than the spinothalamic ("neospinothalamic") tract.

Sleep onset is the transition from wakefulness into sleep. Sleep onset usually transits into non-rapid eye movement sleep but under certain circumstances it is possible to transit from wakefulness directly into rapid eye movement sleep.

<span class="mw-page-title-main">Parabrachial nuclei</span>

The parabrachial nuclei, also known as the parabrachial complex, are a group of nuclei in the dorsolateral pons that surrounds the superior cerebellar peduncle as it enters the brainstem from the cerebellum. They are named from the Latin term for the superior cerebellar peduncle, the brachium conjunctivum. In the human brain, the expansion of the superior cerebellar peduncle expands the parabrachial nuclei, which form a thin strip of grey matter over most of the peduncle. The parabrachial nuclei are typically divided along the lines suggested by Baxter and Olszewski in humans, into a medial parabrachial nucleus and lateral parabrachial nucleus. These have in turn been subdivided into a dozen subnuclei: the superior, dorsal, ventral, internal, external and extreme lateral subnuclei; the lateral crescent and subparabrachial nucleus along the ventrolateral margin of the lateral parabrachial complex; and the medial and external medial subnuclei

The parafacial zone (PZ) is a brain structure located in the brainstem within the medulla oblongata believed to be heavily responsible for non-rapid eye movement (non-REM) sleep regulation, specifically for inducing slow-wave sleep.

The dorsal tegmental nucleus (DTN), also known as dorsal tegmental nucleus of Gudden (DTg), is a group of neurons located in the brain stem, which are involved in spatial navigation and orientation.

References

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  13. 1 2 3 4 5 6 7 Malenka RC, Nestler EJ, Hyman SE (2009). "Chapter 12: Sleep and Arousal". In Sydor A, Brown RY (eds.). Molecular Neuropharmacology: A Foundation for Clinical Neuroscience (2nd ed.). New York: McGraw-Hill Medical. p. 295. ISBN   978-0071481274. The RAS is a complex structure consisting of several different circuits including the four monoaminergic pathways ... The norepinephrine pathway originates from the locus ceruleus (LC) and related brainstem nuclei; the serotonergic neurons originate from the raphe nuclei within the brainstem as well; the dopaminergic neurons originate in ventral tegmental area (VTA); and the histaminergic pathway originates from neurons in the tuberomammillary nucleus (TMN) of the posterior hypothalamus. As discussed in Chapter 6, these neurons project widely throughout the brain from restricted collections of cell bodies. Norepinephrine, serotonin, dopamine, and histamine have complex modulatory functions and, in general, promote wakefulness. The PT in the brain stem is also an important component of the ARAS. Activity of PT cholinergic neurons (REM-on cells) promotes REM sleep. During waking, REM-on cells are inhibited by a subset of ARAS norepinephrine and serotonin neurons called REM-off cells.
  14. 1 2 3 4 5 6 7 8 9 Brudzynski SM (July 2014). "The ascending mesolimbic cholinergic system – a specific division of the reticular activating system involved in the initiation of negative emotional states". Journal of Molecular Neuroscience. 53 (3): 436–445. doi:10.1007/s12031-013-0179-1. PMID   24272957. S2CID   14615039. Understanding of arousing and wakefulness-maintaining functions of the ARAS has been further complicated by neurochemical discoveries of numerous groups of neurons with the ascending pathways originating within the brainstem reticular core, including pontomesencephalic nuclei, which synthesize different transmitters and release them in vast areas of the brain and in the entire neocortex (for review, see Jones 2003; Lin et al. 2011). They included glutamatergic, cholinergic, noradrenergic, dopaminergic, serotonergic, histaminergic, and orexinergic systems (for review, see Lin et al. 2011). ... The ARAS represented diffuse, nonspecific pathways that, working through the midline and intralaminar thalamic nuclei, could change activity of the entire neocortex, and thus, this system was suggested initially as a general arousal system to natural stimuli and the critical system underlying wakefulness (Moruzzi and Magoun 1949; Lindsley et al. 1949; Starzl et al. 1951, see stippled area in Fig. 1). ... It was found in a recent study in the rat that the state of wakefulness is mostly maintained by the ascending glutamatergic projection from the parabrachial nucleus and precoeruleus regions to the basal forebrain and then relayed to the cerebral cortex (Fuller et al. 2011). ... Anatomical studies have shown two main pathways involved in arousal and originating from the areas with cholinergic cell groups, one through the thalamus and the other, traveling ventrally through the hypothalamus and preoptic area, and reciprocally connected with the limbic system (Nauta and Kuypers 1958; Siegel 2004). ... As counted in the cholinergic connections to the thalamic reticular nucleus ...
  15. 1 2 3 4 5 6 7 8 9 10 Schwartz MD, Kilduff TS (December 2015). "The Neurobiology of Sleep and Wakefulness". The Psychiatric Clinics of North America. 38 (4): 615–644. doi:10.1016/j.psc.2015.07.002. PMC   4660253 . PMID   26600100. This ascending reticular activating system (ARAS) is comprised of cholinergic laterodorsal and pedunculopontine tegmentum (LDT/PPT), noradrenergic locus coeruleus (LC), serotonergic (5-HT) Raphe nuclei and dopaminergic ventral tegmental area (VTA), substantia nigra (SN) and periaqueductal gray projections that stimulate the cortex directly and indirectly via the thalamus, hypothalamus and BF.6, 12-18 These aminergic and catecholaminergic populations have numerous interconnections and parallel projections which likely impart functional redundancy and resilience to the system.6, 13, 19 ... More recently, the medullary parafacial zone (PZ) adjacent to the facial nerve was identified as a sleep-promoting center on the basis of anatomical, electrophysiological and chemo- and optogenetic studies.23, 24 GABAergic PZ neurons inhibit glutamatergic parabrachial (PB) neurons that project to the BF,25 thereby promoting NREM sleep at the expense of wakefulness and REM sleep. ... The Hcrt neurons project widely throughout the brain and spinal cord92, 96, 99, 100 including major projections to wake-promoting cell groups such as the HA cells of the TM,101 the 5-HT cells of the dorsal Raphe nuclei (DRN),101 the noradrenergic cells of the LC,102 and cholinergic cells in the LDT, PPT, and BF.101, 103 ... Hcrt directly excites cellular systems involved in waking and arousal including the LC,102, 106, 107 DRN,108, 109 TM,110-112 LDT,113, 114 cholinergic BF,115 and both dopamine (DA) and non-DA neurons in the VTA.116, 117
  16. 1 2 3 4 Saper CB, Fuller PM (June 2017). "Wake-sleep circuitry: an overview". Current Opinion in Neurobiology. 44: 186–192. doi:10.1016/j.conb.2017.03.021. PMC   5531075 . PMID   28577468. Parabrachial and pedunculopontine glutamatergic arousal system
    Retrograde tracers from the BF have consistently identified one brainstem site of input that is not part of the classical monoaminergic ascending arousal system: glutamatergic neurons in the parabrachial and pedunculopontine nucleus ... Juxtacellular recordings from pedunculopontine neurons have found that nearly all cholinergic neurons in this region, as well as many glutamatergic and GABAergic neurons, are most active during wake and REM sleep [25], although some of the latter neurons were maximally active during either wake or REM, but not both. ... [Parabrachial and pedunculopontine glutamatergic neurons] provide heavy innervation to the lateral hypothalamus, central nucleus of the amygdala, and BF
  17. 1 2 Pedersen NP, Ferrari L, Venner A, Wang JL, Abbott SG, Vujovic N, Arrigoni E, Saper CB, Fuller PM (November 2017). "Supramammillary glutamate neurons are a key node of the arousal system". Nature Communications. 8 (1): 1405. Bibcode:2017NatCo...8.1405P. doi:10.1038/s41467-017-01004-6. PMC   5680228 . PMID   29123082. Basic and clinical observations suggest that the caudal hypothalamus comprises a key node of the ascending arousal system, but the cell types underlying this are not fully understood. Here we report that glutamate-releasing neurons of the supramammillary region (SuMvglut2) produce sustained behavioral and EEG arousal when chemogenetically activated.
  18. 1 2 3 Burlet S, Tyler CJ, Leonard CS (April 2002). "Direct and indirect excitation of laterodorsal tegmental neurons by Hypocretin/Orexin peptides: implications for wakefulness and narcolepsy". J. Neurosci. 22 (7): 2862–2872. doi:10.1523/JNEUROSCI.22-07-02862.2002. PMC   6758338 . PMID   11923451.
  19. ↑ Malenka RC, Nestler EJ, Hyman SE (2009). "Chapter 12: Sleep and Arousal". In Sydor A, Brown RY (eds.). Molecular Neuropharmacology: A Foundation for Clinical Neuroscience (2nd ed.). New York: McGraw-Hill Medical. p. 295. ISBN   978-0071481274. Orexin neurons are located in the lateral hypothalamus. They are organized in a widely projecting manner, much like the monoamines (Chapter 6), and innervate all of the components of the ARAS. They excite the REM-off monoaminergic neurons during wakefulness and the PT cholinergic neurons during REM sleep. They are inhibited by the VLPO neurons during NREM sleep.
  20. 1 2 Cherasse Y, Urade Y (November 2017). "Dietary Zinc Acts as a Sleep Modulator". International Journal of Molecular Sciences. 18 (11): 2334. doi: 10.3390/ijms18112334 . PMC   5713303 . PMID   29113075. The regulation of sleep and wakefulness involves many regions and cellular subtypes in the brain. Indeed, the ascending arousal system promotes wakefulness through a network composed of the monaminergic neurons in the locus coeruleus (LC), histaminergic neurons in the tuberomammilary nucleus (TMN), glutamatergic neurons in the parabrachial nucleus (PB) ...
  21. ↑ Fuller PM, Fuller P, Sherman D, Pedersen NP, Saper CB, Lu J (April 2011). "Reassessment of the structural basis of the ascending arousal system". The Journal of Comparative Neurology. 519 (5): 933–956. doi:10.1002/cne.22559. PMC   3119596 . PMID   21280045.
  22. 1 2 Kinomura S, Larsson J, Gulyás B, Roland PE (January 1996). "Activation by attention of the human reticular formation and thalamic intralaminar nuclei". Science. 271 (5248): 512–515. Bibcode:1996Sci...271..512K. doi:10.1126/science.271.5248.512. PMID   8560267. S2CID   43015539. This corresponds to the centro-median and centralis lateralis nuclei of the intralaminar group
  23. ↑ Svorad D (January 1957). "Reticular activating system of brain stem and animal hypnosis". Science. 125 (3239): 156. Bibcode:1957Sci...125..156S. doi:10.1126/science.125.3239.156. PMID   13390978.
  24. 1 2 Jang SH, Kwon HG (October 2015). "The direct pathway from the brainstem reticular formation to the cerebral cortex in the ascending reticular activating system: A diffusion tensor imaging study". Neurosci. Lett. 606: 200–203. doi:10.1016/j.neulet.2015.09.004. PMID   26363340. S2CID   37083435.
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