Dorsal cochlear nucleus

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Dorsal cochlear nucleus
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Dorsal cochlear nucleus is #4, at upper left
Human caudal brainstem posterior view description.JPG
Human caudal brainstem posterior view (Dorsal cochlear nucleus is #5)
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Identifiers
Latin nucleus cochlearis posterior
NeuroNames 721
NeuroLex ID birnlex_2569
TA98 A14.1.04.248
TA2 6007
FMA 54624
Anatomical terms of neuroanatomy

The dorsal cochlear nucleus (DCN, also known as the "tuberculum acusticum") is a cortex-like structure on the dorso-lateral surface of the brainstem. Along with the ventral cochlear nucleus (VCN), it forms the cochlear nucleus (CN), where all auditory nerve fibers from the cochlea form their first synapses.

Contents

Anatomy

The DCN differs from the ventral portion of the CN as it not only projects to the central nucleus (a subdivision) of the inferior colliculus (CIC), but also receives efferent innervation from the auditory cortex, superior olivary complex and the inferior colliculus. The cytoarchitecture and neurochemistry of the DCN is similar to that of the cerebellum, an important concept in theories of DCN function. [1] Thus, the DCN is thought to be involved with more complex auditory processing, rather than merely transferring information.

The pyramidal cells or giant cells are a major cell grouping of the DCN. These cells are the target of two different input systems. The first system arises from the auditory nerve, and carries acoustic information. The second set of inputs is relayed through a set of small granule cells in the cochlear nucleus. There are also a great number of neighbouring cartwheel cells. [2] The granule cells in turn are the target of a number of different inputs, including both those involved in auditory processing and, at least in lower mammals, somatosensory inputs associated with the head, the ear, and the jaw.

Projections from DCN principal cells form the dorsal acoustic stria, which ultimately terminate in the CIC. This projection overlaps with that of the lateral superior olive (LSO) in a well-defined manner, [3] where they form the primary excitatory input for ICC type O units. [4]

Physiology

Principal cells in the DCN have very complex frequency intensity tuning curves. Classified as cochlear nucleus type IV cells, [5] the firing rate may be very rapid in response to a low intensity sound at one frequency and then fall below the spontaneous rate with only a small increment in stimulus frequency or intensity. The firing rate may then increase with another increment in intensity or frequency. Type IV cells are excited by wide band noise, and particularly excited by a noise-notch stimulus directly below the cell's best frequency (BF).

While the VCN bushy cells aid in the location of a sound stimulus on the horizontal axis via their inputs to the superior olivary complex, type IV cells may participate in localization of the sound stimulus on the vertical axis. The pinna selectively amplifies frequencies, resulting in reduced sound energy at specific frequencies in certain regions of space. The complicated firing patterns of type IV cells makes them especially suited to detecting these notches, and with the combined power of these two localization systems, an ordinary person can locate where a firework explodes without the use of their eyes.

Somatosensory inputs inhibit type IV cell activity, possibly silencing their activity during head and pinna movements. [6] While this has not been studied extensively, it may play an important role in sound source localization in elevation. A similar effect is seen in the visual system in an effect known as change blindness.

Current auditory models of the DCN employ a two-inhibitor model. Type IV cells receive excitation directly from the auditory nerve, and are inhibited by type II (vertical) cells and a wide band inhibitor (onset-c cells).

Related Research Articles

Nociception is the sensory nervous system's process of encoding noxious stimuli. It deals with a series of events and processes required for an organism to receive a painful stimulus, convert it to a molecular signal, and recognize and characterize the signal in order to trigger an appropriate defense response.

<span class="mw-page-title-main">Auditory system</span> Sensory system used for hearing

The auditory system is the sensory system for the sense of hearing. It includes both the sensory organs and the auditory parts of the sensory system.

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

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">Inferior colliculus</span> Midbrain structure involved in the auditory pathway.

The inferior colliculus (IC) is the principal midbrain nucleus of the auditory pathway and receives input from several peripheral brainstem nuclei in the auditory pathway, as well as inputs from the auditory cortex. The inferior colliculus has three subdivisions: the central nucleus, a dorsal cortex by which it is surrounded, and an external cortex which is located laterally. Its bimodal neurons are implicated in auditory-somatosensory interaction, receiving projections from somatosensory nuclei. This multisensory integration may underlie a filtering of self-effected sounds from vocalization, chewing, or respiration activities.

In physiology, tonotopy is the spatial arrangement of where sounds of different frequency are processed in the brain. Tones close to each other in terms of frequency are represented in topologically neighbouring regions in the brain. Tonotopic maps are a particular case of topographic organization, similar to retinotopy in the visual system.

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">Trapezoid body</span> Part of the auditory pathway

The trapezoid body is part of the auditory pathway where some of the axons coming from the cochlear nucleus decussate to the other side before traveling on to the superior olivary nucleus. This is believed to help with localization of sound.

<span class="mw-page-title-main">Thalamocortical radiations</span> Neural pathways between the thalamus and cerebral cortex

In neuroanatomy, thalamocortical radiations are the fibers between the thalamus and the cerebral cortex.

<span class="mw-page-title-main">Medial geniculate nucleus</span>

The medial geniculate nucleus (MGN) or medial geniculate body (MGB) is part of the auditory thalamus and represents the thalamic relay between the inferior colliculus (IC) and the auditory cortex (AC). It is made up of a number of sub-nuclei that are distinguished by their neuronal morphology and density, by their afferent and efferent connections, and by the coding properties of their neurons. It is thought that the MGN influences the direction and maintenance of attention.

<span class="mw-page-title-main">Cochlear nerve</span> Nerve carrying auditory information from the inner ear to the brain

The cochlear nerve is one of two parts of the vestibulocochlear nerve, a cranial nerve present in amniotes, the other part being the vestibular nerve. The cochlear nerve carries auditory sensory information from the cochlea of the inner ear directly to the brain. The other portion of the vestibulocochlear nerve is the vestibular nerve, which carries spatial orientation information to the brain from the semicircular canals, also known as semicircular ducts.

<span class="mw-page-title-main">Cochlear nucleus</span> Two cranial nerve nuclei of the human brainstem

The cochlear nuclear (CN) complex comprises two cranial nerve nuclei in the human brainstem, the ventral cochlear nucleus (VCN) and the dorsal cochlear nucleus (DCN). The ventral cochlear nucleus is unlayered whereas the dorsal cochlear nucleus is layered. Auditory nerve fibers, fibers that travel through the auditory nerve carry information from the inner ear, the cochlea, on the same side of the head, to the nerve root in the ventral cochlear nucleus. At the nerve root the fibers branch to innervate the ventral cochlear nucleus and the deep layer of the dorsal cochlear nucleus. All acoustic information thus enters the brain through the cochlear nuclei, where the processing of acoustic information begins. The outputs from the cochlear nuclei are received in higher regions of the auditory brainstem.

<span class="mw-page-title-main">Superior olivary complex</span> Collection of brainstem nuclei related to hearing

The superior olivary complex (SOC) or superior olive is a collection of brainstem nuclei that functions in multiple aspects of hearing and is an important component of the ascending and descending auditory pathways of the auditory system. The SOC is intimately related to the trapezoid body: most of the cell groups of the SOC are dorsal to this axon bundle while a number of cell groups are embedded in the trapezoid body. Overall, the SOC displays a significant interspecies variation, being largest in bats and rodents and smaller in primates.

<span class="mw-page-title-main">Interaural time difference</span>

The interaural time difference when concerning humans or animals, is the difference in arrival time of a sound between two ears. It is important in the localization of sounds, as it provides a cue to the direction or angle of the sound source from the head. If a signal arrives at the head from one side, the signal has further to travel to reach the far ear than the near ear. This pathlength difference results in a time difference between the sound's arrivals at the ears, which is detected and aids the process of identifying the direction of sound source.

Binaural fusion or binaural integration is a cognitive process that involves the combination of different auditory information presented binaurally, or to each ear. In humans, this process is essential in understanding speech as one ear may pick up more information about the speech stimuli than the other.

In human neuroanatomy, brainstem auditory evoked potentials (BAEPs), also called brainstem auditory evoked responses (BAERs), are very small auditory evoked potentials in response to an auditory stimulus, which are recorded by electrodes placed on the scalp. They reflect neuronal activity in the auditory nerve, cochlear nucleus, superior olive, and inferior colliculus of the brainstem. They typically have a response latency of no more than six milliseconds with an amplitude of approximately one microvolt.

<span class="mw-page-title-main">Ventral cochlear nucleus</span>

In the ventral cochlear nucleus (VCN), auditory nerve fibers enter the brain via the nerve root in the VCN. The ventral cochlear nucleus is divided into the anterior ventral (anteroventral) cochlear nucleus (AVCN) and the posterior ventral (posteroventral) cochlear nucleus (PVCN). In the VCN, auditory nerve fibers bifurcate, the ascending branch innervates the AVCN and the descending branch innervates the PVCN and then continue to the dorsal cochlear nucleus. The orderly innervation by auditory nerve fibers gives the AVCN a tonotopic organization along the dorsoventral axis. Fibers that carry information from the apex of the cochlea that are tuned to low frequencies contact neurons in the ventral part of the AVCN; those that carry information from the base of the cochlea that are tuned to high frequencies contact neurons in the dorsal part of the AVCN. Several populations of neurons populate the AVCN. Bushy cells receive input from auditory nerve fibers through particularly large endings called end bulbs of Held. They contact stellate cells through more conventional boutons.

<span class="mw-page-title-main">Calyx of Held</span>

The Calyx of Held is a particularly large synapse in the mammalian auditory central nervous system, so named after Hans Held who first described it in his 1893 article Die centrale Gehörleitung because of its resemblance to the calyx of a flower. Globular bushy cells in the anteroventral cochlear nucleus (AVCN) send axons to the contralateral medial nucleus of the trapezoid body (MNTB), where they synapse via these calyces on MNTB principal cells. These principal cells then project to the ipsilateral lateral superior olive (LSO), where they inhibit postsynaptic neurons and provide a basis for interaural level detection (ILD), required for high frequency sound localization. This synapse has been described as the largest in the brain.

The olivocochlear system is a component of the auditory system involved with the descending control of the cochlea. Its nerve fibres, the olivocochlear bundle (OCB), form part of the vestibulocochlear nerve, and project from the superior olivary complex in the brainstem (pons) to the cochlea.

<span class="mw-page-title-main">Sound localization in owls</span> Ability of owls to locate sounds in 3D space

Most owls are nocturnal or crepuscular birds of prey. Because they hunt at night, they must rely on non-visual senses. Experiments by Roger Payne have shown that owls are sensitive to the sounds made by their prey, not the heat or the smell. In fact, the sound cues are both necessary and sufficient for localization of mice from a distant location where they are perched. For this to work, the owls must be able to accurately localize both the azimuth and the elevation of the sound source.

References

  1. Bell CC, Han V et al. Cerebellum-like structures and their implications for cerebellar function. Annu.Rev.Neurosci. 2008.Vol31. PMID   18275284
  2. Wouterlood FG and Mugnaini E. Cartwheel neurons of the dorsal cochlear nucleus: a Golgi-electron microscopic study in rat. J.Comp Neurol 1984 Jul20;227(1) PMID   6088594
  3. Oliver, Douglas L.; Beckius, Gretchen E.; Bishop, Deborah C.; Kuwada, Shigeyuki (2 June 1997). "Simultaneous anterograde labeling of axonal layers from lateral superior olive and dorsal cochlear nucleus in the inferior colliculus of cat". The Journal of Comparative Neurology. 382 (2): 215–229. doi:10.1002/(SICI)1096-9861(19970602)382:2<215::AID-CNE6>3.0.CO;2-6. PMID   9183690.
  4. Davis, Kevin A. (April 2002). "Evidence of a Functionally Segregated Pathway From Dorsal Cochlear Nucleus to Inferior Colliculus". Journal of Neurophysiology. 87 (4): 1824–1835. doi:10.1152/jn.00769.2001. PMID   11929904.
  5. Shofner, WP; Young, ED (October 1985). "Excitatory/inhibitory response types in the cochlear nucleus: relationships to discharge patterns and responses to electrical stimulation of the auditory nerve". Journal of Neurophysiology. 54 (4): 917–39. doi:10.1152/jn.1985.54.4.917. PMID   4067627.
  6. Young, ED; Nelken, I; Conley, RA (February 1995). "Somatosensory effects on neurons in dorsal cochlear nucleus". Journal of Neurophysiology. 73 (2): 743–65. doi:10.1152/jn.1995.73.2.743. PMID   7760132.