Primary auditory cortex
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| Brain: Primary auditory cortex | ||
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| Brodmann areas 41 & 42 of the human brain. | ||
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| The Primary Auditory Cortex is highlighted in magenta, and has been known to interact with all areas highlighted on this neural map. | ||
| NeuroNames | ancil-428 | |
The primary auditory cortex is the region of the brain that is responsible for processing of auditory (sound) information.
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[edit] Function of the Primary Auditory Cortex
As with other primary sensory cortical areas, auditory sensations reach perception only if received and processed by a cortical area. Evidence for this comes from lesion studies in human patients who have sustained damage to cortical areas through tumors or strokes, or from animal experiments in which cortical areas were deactivated by cooling or locally applied drug treatment. Damage to the Primary Auditory Cortex in humans leads to a loss of any 'awareness' of sound, but an ability to react reflexively to sounds remains as there is a great deal of subcortical processing in the auditory brainstem and midbrain.
Neurons in the auditory cortex are organised according to the frequency of sound to which they respond best. Neurons at one end of the auditory cortex respond best to low frequencies; neurons at the other respond best to high frequencies. There are multiple auditory areas (much like the multiple areas in the visual cortex), which can be distinguished anatomically and on the basis that they contain a complete "frequency map." The purpose of this frequency map (known as a tonotopic map) is unknown and is likely to reflect the fact that the sensory epithelium of the auditory system, the cochlea, is arranged according to sound frequency. The auditory cortex is involved in tasks such as identifying and segregating auditory "objects" and identifying the location of a sound in space.
Human brain scans have indicated that a peripheral bit of this brain region is active when trying to identify pitch. Individual cells consistently get excited by sounds at specific frequencies, or multiples of that frequency.
The primary auditory cortex is about the same as Brodmann areas 41 and 42. It lies in the posterior half of the superior temporal gyrus and also dives into the lateral sulcus as the transverse temporal gyri (also called Heschl's gyri).
The primary auditory cortex is located in the temporal lobe. There are additional areas of the human cerebral cortex that are involved in processing sound, in the frontal and parietal lobes. Animal studies indicate that auditory fields of the cerebral cortex receive ascending input from the auditory thalamus, and that they are interconnected on the same and on the opposite cerebral hemispheres.
The auditory cortex is composed of fields, which differ from each other in both structure and function.[1]
The number of fields varies in different species, from as few as 2 in rodents to as many as 15 in the rhesus monkey. The number, location, and organization of fields in the human auditory cortex are not known at this time. What is known about the human auditory cortex comes from a base of knowledge gained from studies in mammals, including primates, used to interpret electrophysiologic tests and functional imaging studies of the brain in humans.
When each instrument of the symphony orchestra, and the jazz band, plays the same note, the quality of each sound is different — but the musician perceives each note as having the same pitch. The neurons of the auditory cortex of the brain are able to respond to pitch. Studies in the marmoset monkey have shown that pitch-selective neurons are located in a cortical region near the anterolateral border of the primary auditory cortex. This location of a pitch-selective area has also been identified in recent functional imaging studies in humans.[2][3]
The auditory cortex does not just receive input from lower centers and the ear, but also provides it.
[edit] Brodmann area 41
This area is also known as anterior transverse temporal area 41 (H). It is a subdivision of the cytoarchitecturally-defined temporal region of cerebral cortex, occupying the anterior transverse temporal gyrus (H) in the bank of the lateral sulcus on the dorsal surface of the temporal lobe. Brodmann area 41 is bounded medially by the parainsular area 52 (H) and laterally by the posterior transverse temporal area 42 (H) (Brodmann-1909).
[edit] Brodmann area 42
This area is also known as posterior transverse temporal area 42 (H). It is a subdivision of the cytoarchitecturally-defined temporal region of cerebral cortex, located in the bank of the lateral sulcus on the dorsal surface of the temporal lobe. Brodmann area 42 is bounded medially by the anterior transverse temporal area 41 (H) and laterally by the superior temporal area 22 (Brodmann-1909).
[edit] Relationship to auditory system
The auditory cortex is the most highly organized processing unit of sound in the brain. This cortex area is the neural crux of hearing, and, in humans, language and music.
The auditory cortex is divided into three separate parts, the primary, secondary and tertiary auditory cortex. These structures are formed concentrically around one another, with the primary AC in the middle and the tertiary AC on the outside.
The primary auditory cortex is tonotopically organized, which means that certain cells in the auditory cortex are sensitive to specific frequencies. This is a fascinating function which has been preserved throughout most of the audition circuit. This area of the brain “is thought to identify the fundamental elements of music, such as pitch and loudness.”[4] This makes sense as this is the area which receives direct input from the medial geniculate nucleus of the thalamus. The secondary auditory cortex has been indicated in the processing of “harmonic, melodic and rhythmic patterns.”[4] The tertiary auditory cortex supposedly integrates everything into the overall experience of music.[4]
An evoked response study of congenitally deaf kittens by Klinke et al. utilized field potentials to measure cortical plasticity in the auditory cortex. These kittens were stimulated and measured against a control or un-stimulated CDC and normal hearing cats. The field potentials measured for artificially stimulated CDC was eventually much stronger than that of a normal hearing cat.[5] This is in concordance with Eckart Altenmuller’s study where it was observed that students who received musical instruction had greater cortical activation than those who did not.[6]
The auditory cortex exhibits some strange behavior pertaining to the gamma wave frequency. When subjects are exposed to three or four cycles of a 40 hertz click, an abnormal spike appears in the EEG data, which is not present for other stimuli. The spike in neuronal activity correlating to this frequency is not restrained to the tonotopic organization of the auditory cortex. It has been theorized that this is a “resonant frequency” of certain areas of the brain, and appears to affect the visual cortex as well.[7]
Gamma band activation (20 to 40 Hz) has been shown to be present during the perception of sensory events and the process of recognition. Kneif et al, in their 2000 study, presented subjects with eight musical notes to well known tunes, such as Yankee Doodle and Frere Jacques. Randomly, the sixth and seventh notes were omitted and an electroencephalogram, as well as a magnetoencephalogram were each employed to measure the neural results. Specifically, the presence of gamma waves, induced by the auditory task at hand, were measured from the temples of the subjects. The OSP response, or omitted stimulus response, was located in a slightly different position; 7 mm more anterior, 13 mm more medial and 13 mm more superior in respect to the complete sets. The OSP recordings were also characteristically lower in gamma waves, as compared to the complete musical set. The evoked responses during the sixth and seventh omitted notes are assumed to be imagined, and were characteristically different, especially in the right hemisphere.[8] The right auditory cortex has long been shown to be more sensitive to tonality, while the left auditory cortex has been shown to be more sensitive to minute sequential differences in sound specifically speech.
Hallucinations have been shown to produce oscillations which are parallel (although not exactly the same as) the gamma frequency range. Sperling showed in his 2004 study that auditory hallucinations produce band wavelengths in the range of 12.5-30 Hz. The bands occurred in the left auditory cortex of a schizophrenic and were controlled against 13 controls (18) . This aligns with the studies of people remembering a song in their minds; they do not perceive any sound, but experience the melody, rhythm and overall experience of sound. When schizophrenics experience hallucinations, it is the primary auditory cortex which becomes active. This is characteristically different from remembering a sound stimulus, which only faintly activates the tertiary auditory cortex.[9] By deduction, an artificial stimulation of the primary auditory cortex should elicit an incredibly real auditory hallucination. The termination of all audition and music into the tertiary auditory cortex creates a fascinating nexus of aural information. If this theory is true, it would be interesting to study a subject with a damaged, TAC or one with artificially suppressed function. This would be very difficult to do as the tertiary cortex is simply a ring around the secondary, which is a ring around the primary AC.
Tone is perceived in more places than just the auditory cortex; one specifically fascinating area is the rostromedial prefrontal cortex.[10] Janata et al, in their 2002 study, used an fMRI machine to study the areas of the brain which were active during tonality processing. The result of which displayed several areas which are not normally considered to be part of the audition process. The rostromedial prefrontal cortex is a subsection of the medial prefrontal cortex, which projects to the amygdala, and is thought to aid in the inhibition of negative emotion.[11] The medial prefrontal cortex is thought to be the core developmental difference between the impulsive teenager and the calm adult. The rostromedial prefrontal cortex is tonality sensitive, meaning it is activated by the tones and frequencies of resonant sounds and music. It could be hypothesized that this is the mechanism by which music ameliorates the soul (or, if one prefers, the limbic system).
[edit] References
- ^ Cant, NB (Jun 15, 2003). "Parallel auditory pathways: projection patterns of the different neuronal populations in the dorsal and ventral cochlear nuclei". Brain Res Bull. 60 (5-6): 457-74. DOI:10.1016/S0361-9230(03)00050-9.
- ^ Bendor, D; Wang, X (2005). "The neuronal representation of pitch in primate auditory cortex.". Nature 436 (7054): 1161-5. DOI:10.1038/nature03867.
- ^ Zatorre, RJ (2005). "Neuroscience: finding the missing fundamental". Nature 436 (7054): 1093-4. DOI:10.1038/4361093a.
- ^ a b c Ferragamo, M. J. (2002). "Octopus cells of the mammalian ventral cochlear nucleus sense the rate of depolarization". J Neurophysiol 87 (5): 2262-70.
- ^ Klinke, Rainer; Kral, Andrej; Heid, Silvia ; Tillein, Jochen ; Hartmann , Rainer (Sep. 10, 1999). "Recruitment of the auditory cortex in congenitally deaf cats by long-term cochlear electrostimulation". Science 285 (5434): 1729-33. DOI:10.1126/science.285.5434.1729.
- ^ Strickland (Winter 2001). "Music and the brain in childhood development". Childhood Education 78 (2): 100-4.
- ^ Bertrand, O.; Tallon-Baudry, C.; Fischer, C.; and Pernier, J.. "Object representation and gamma oscillations".
- ^ Knief, A.; Schulte, M.; Fujiki, N.; and Pantev, C.. "Oscillatory Gamma band and Slow brain Activity Evoked by Real and Imaginary Musical Stimuli".
- ^ Abbott, Alison Music, maestro, please! Nature v. 416 no. 6876 (March 7 2002)
- ^ Petr Janata et al. The Cortical Topography of Tonal Structures Underlying Western Music. Science, Vol 298, Issue 5601, 2167-2170 , 13 December 2002
- ^ Cassel, Topography of projections from the medial prefrontal cortex to the amygdala in the rat. Brain Res Bull. 1986 Sep;17(3):321-33
[edit] External links
- NeuroNames ancil-77: area 41
- NeuroNames ancil-78: area 42
- BrainMaps at UCDavis primary auditory cortex
