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The Brain’s Processing of Music
Romesh Senewiratne (MD)
What happens in the brain and nervous system when we listen to music and
when we create musical sounds? What parts of the brain are activated by
music, and how are these connected with each other? Which aspects of music
perception are processed by the right side of the brain and which by the left
side? What is the relationship between neural processing of verbal language
and that of music?
This is a puzzle that has unraveled considerably in recent years, during which
neuroscientists have investigated neural activity in people with intact brains
(including variously trained musicians) using new imaging techniques, such as
functional magnetic resonance imaging (fMRI) and positron emission
technology (PET) scanning. Such research has followed a longer tradition of
inferring the function of parts of the brain by studying people with acquired
musical deficits (amusia), and the use of increasingly sophisticated cognitive
evaluations of the musicality of subjects with and without known brain damage
and with and without musical training. Attempts to integrate these and other
findings have led to both consensus and controversy about the neural
processing of music.
The fact that music perception is 'subjective' is sometimes put forward as a
reason to support the argument that music cannot be studied 'scientifically' at
all. It is true that one person's 'music' is another's 'noise'. At times, what one
listener loves may induce boredom or displeasure in another. The experience
of music is subjective in many ways, and the development of taste is
idiosyncratic, since everyone’s musical experience is, to some degree, unique.
At the same time, certain aspects of music are based on the objective
properties of sound waves – their frequency, amplitude, duration and form.
These have psychological equivalents: pitch, loudness, timing and timbre,
respectively. The elements of music that are commonly discussed are derived,
subjectively and with certain distortions, from these physical properties of
sound waves. These elements unfold to the listener with each new note –
change in pitch leads to melody, simultaneous pitches to harmony. Rhythm is
derived from the temporal arrangement of notes which also determine the
meter and tempo of the music. Timbre, or tonal quality, is determined by the
form of the sound wave. These elements are perceived as distinct attributes of
music, providing a means of sensibly discussing the neural and mental
processing of the sounds we identify, differently, as 'music'. It is common to
hear people describing music in terms of these elements; not so the physical
properties of the sound waves that give rise to them. These music parameters
or elements have been vigorously researched in universities around the world
over the past 40 years or more, with most of the global publications emanating
from the USA, Canada and Europe.
Several of these elements have equivalents in spoken language. Prosody is the
melody of spoken language. The rhythm of spoken language is found in syntax
and phrasing. The timbre of the spoken voice may be as important for verbal
communication and the attractiveness of spoken words as it is for music.
Harmony is a more exclusively musical phenomenon; though one may speak of
a harmonious discussion, when people talk at once the result is cacophony, not
music. (Except the curious exception when people speak in union). We also
speak of harmonious colours and forms, not to mention harmonious flavours.
Discovering the neurobiology of language and music
A longstanding debate has centred on the neural processing of language versus
music. Attribution of speech to the dominant left hemisphere, and localisation
of speech expression and comprehension to separate localised areas of cortex
has been part of medical orthodoxy since the famous French neurologist Paul
Broca (1824-1880) described selective loss of speech capacity in patients with
damage to a small area in the left, inferior frontal lobe, as confirmed at postmortem. The discovery of what is now called ‘Broca’s Area’, occurred in the
1860s and rekindled a waning enthusiasm for phrenology – the often
misguided science of localising aspects of the mind to specific parts of the
brain. Though the first phrenological ‘maps’ in the early nineteenth century
included an ‘organ of tune’ separate from an ‘organ of language’ its proposed
location was more whimsical than scientific.
Sylvian (transverse) fissure
Primary Auditory Cortex (PAC)
Fig 1. Location of Wernicke’s and
Broca’s areas relative to the PAC.
In 1874, not long after Broca’s discovery of expressive aphasia following
localised damage to an area of cortex in the left frontal lobe, the German
neurologist Carl Wernicke (1848-1905) described a specific deficit in
understanding speech (receptive aphasia) following damage to another part of
the cortex, again on the left side, but posterior to the auditory cortex. This
area, located in and around the angular gyrus that curves around the tip of the
Sylvian fissure, is now known as Wernicke’s Area.
The basic model of Wernicke, that auditory information is transmitted from
the medial geniculate nucleus of the thalamus to the primary auditory cortex
(in the temporal lobes), and from there to Wernicke’s area has stood the test
of time well, although minor adjustments to the model have been necessary to
accommodate subsequent clinical and neurological findings. His hypothesis
that words which are read are also comprehended in Wernicke’s area has been
subsequently modified: visual information does pass through the lateral
geniculate nucleus (LGN) of the thalamus as Wernicke theorized in the 19th
century, but is referred to an area of cortex adjacent to the part involved with
comprehension of spoken and signed language (Wernicke’s Area). There also
appears to be different processing, involving different parts of the brain, for
words that make sense and ‘nonsense words’ and, like many neuroscience
models that followed, Wernicke’s model neglects the emotional reactions
elicited by words, speech and language.
The identification of Broca’s and Wernicke’s speech areas in the left-sided
cortex led to a resurgence of ‘phrenological’ thinking, with neuroscientists
agreeing that many, or most, mental functions are localised to parts of the
cortex. There were exceptions to this, however. Some mental functions, such
as memory, and consciousness itself, were regarded by many neuroscientists
as emergent properties from the organisation of the whole brain. It became an
accepted dogma that speech sounds are processed on the left side of the brain
while non-verbal sounds (including music) are processed on the right. Though
this generalisation has some merits it is an oversimplification in some ways and
frankly inaccurate in others.
The complex mental processes that occur in comprehending speech and
communicating verbally involve many areas of the brain simultaneously,
including areas associated with recognition of words, comprehension of
phrases, memory formation and recall, creation of ideas and language,
simultaneous visual imagery, and emotional response. Though verbal language
is mainly a left sided function the neural mechanisms and pathways underlying
language appear to be widely distributed in the brain, as they are in music
Following the discovery of receptive and expressive speech centres in the left
side of the brain in all right-handed people and most left-handed and
ambidextrous ones as well, the non-linguistic aspects of music as well as other
non-verbal environmental sounds have been commonly ascribed to the right
side of the brain. In fact, music can truly be said to provide exercise for the
whole brain (and with practice, the whole body). It is true, however, that
particular aspects of musical thought involve different parts of the brain, some
of these being lateralised, as is the comprehension and generation of speech.
The right temporal lobe, in particular, has been associated with specific musical
functions, especially in fine discrimination of pitch, melody and harmony.
However, both right and left temporal auditory cortices as well as frontal,
parietal and occipital lobes appear to be involved in various aspects of the
perception and creation of music. In addition to these (outer) cortical areas,
other areas in the core of the brain are also involved in musical activity. These
will be further explored in the following analysis.
Such amusia has been reported following damage to the right side of the brain and right temporal lobes. . suggesting a degree of independence between musical elements in the perceptual analysis of music. including deficits in perception.6 Hemispheric lateralisation of speech and music Much knowledge of the neural processing of music has been acquired over the years by the study of people who have lost musical ability after localised brain injuries. however the authors found no correspondence between amusia and aphasia in those with left hemisphere damage. In 2000. Although rare (especially because musical deficits are rarely tested for) isolated loss of various aspects of musical ability can result from strokes. Sometimes. These can affect perception of. sometimes not. in particular.5% of patients) with right hemisphere damage and 9 out of 12 (75% of patients) with left hemisphere damage. following damage to the left temporal lobe. melody or timbre but not rhythm or tempo. Of those with left hemisphere damage half (6 out of 12) had evidence of aphasia. brain surgery or head injuries. It has been observed that people may develop different forms of amusia after strokes. This included 5 out of 8 (62. Echart Altenmuller and colleagues tested patients who were still in hospital following known left and right sided strokes. verbal deficits accompany the musical deficit. expression or both. Musical deficits acquired in this way may specifically affect one or more aspects of music cognition. while others are spared. though. Amusia has also been reported. Though none reported amusia spontaneously (partly due to the amusical hospital environment) when tested with a battery of neuropsychological tests for musicality a surprising number had subtle music reception deficits. a study from Germany’s Institute of Music Physiology and Performing Arts Medicine by Maria Schuppert. say. after such injury.
it may be possible to use such testing to chart the recovery of patients after stroke. a neuropsychological test battery for the evaluation of music processing. even in the long term. suggesting a much lower incidence of amusia. these earlier studies had tested patients several weeks or months after the stroke. The authors also approach the question of whether melodic and temporal structures are processed by autonomous . They concluded that listening to music involves an initial right hemisphere recognition of contour and metre followed by identification of interval and rhythm by subsystems in the left hemisphere (Schuppert et al. Closer analysis of the musical deficits in these stroke patients by the German researchers considered the relative contributions of the left and right hemisphere as inferred by observable damage to the brain on MRI scanning. even though there may be coexistence of language and musical impairments. This suggests that music testing can be used to monitor recovery from strokes. Since symptoms of amusia often resolve with time. until they developed one. except for active musicians. The authors make the observation that these deficits often remain unnoticed since.7 This study used simple tests of musical ability that could be quickly and easily used in a hospital setting. On the matter of evolution of the music instincts. they point out. and maybe to predict prognosis (a vexing problem for neurologists at present). In their analysis Schuppert and colleagues compare their results with earlier studies by the Canadian neuroscientists Robert Zatorre and Isabel Peretz who have also done extensive research on patients with strokes. 2000). and help provide a guide to prognosis. the German authors of this study support the hypothesis that musical stimuli are predominantly processed independently of language functions. has not been available. Also. providing sufficient sensitivity yet being short enough to be carried out as a bedside test. Significantly. the patients may not be aware of altered music perception.
read. melody and pitch discrimination do appear to be lateralised to the right side (to some degree. The occurrence of musical auditory hallucinations in temporal lobe epilepsy provides further support for the view that the temporal lobes. The relative importance of the left and right hemispheres of the brain in processing music have been the subject of scientific study since the mid-19th century and the first description. read or write in words. are involved in perceiving. in particular. Since then it has been established that injuries affecting speech may leave musical ability intact and vice versa. The fact that amusia was more likely to be noted with right-sided brain injuries and aphasia with left-sided ones strengthened the view that music is a ‘right brain activity’ while language is ‘left brained’.8 neural networks. write and compose music after a brain injury left him unable to speak. Studies since the 1940s have revealed much greater complexity. in . analysing and remembering music. Such testing began in the 1950s in Canada with the work of Brenda Milner and has continued in Montreal with the work of Isabel Peretz. While timbre. In their sample 10 out of 14 symptomatic subjects had combined deficits in melodic and temporal perception. suggesting that different areas of the brain are necessary for speech and music. and right temporal lobe in particular. while 3 had dissociated deficits in either melodic or temporal discrimination tasks. Other evidence localising the cognition of at least some elements of music to the right temporal lobe has come from studies of people who have had surgical excision of the temporal lobes (usually for the treatment of temporal lobe epilepsy). of a musician who continued to be able to play. pointing out that it is still controversial due to contradictory findings in brain-damaged patients. by Jean-Baptiste Bouillard in 1865. Most often brain injuries causing musical deficits affect language as well (and vice versa) though there have been many detailed reports of specific loss of various musical functions with linguistic ability spared.
melodic contour perception.9 most people) other elements of music. In terms of temporal relations. do not show the same degree of lateralisation. maintaining a steady tempo. brain injuries can selectively affect pitch discrimination. emotional reactions from music or other aspects of ‘subprocessing’. melody recognition. musical memory. Such findings have contributed to the wide acceptance of a ‘modular processing model’ – the view that different elements of music and language employ both shared and distinct neural substrates. Certain aspects of timbre discrimination may be affected and not others. especially rhythm. there is evidence that meter and rhythm (patterns of strong/loud and weak/soft beats) have opposite lateralisation – the ‘grouping’ aspect of rhythm is more associated with the left hemisphere while maintaining and synchronising with a regular meter in music appears more right-sided. Cortical localisation Frontal lobes Primary Auditory Cortex (PAC) in the first transverse gyrus of the temporal lobe (Heschl’s gyrus) Thalamus Medial Geniculate Nucleus (MGN) of the thalamus Fig. or are lateralised to the left hemisphere. Furthermore.2: Location of PAC and MGN .
does cause ‘sensory deafness’. auditory information is distributed to both sides of the brain from each ear. Studies have shown that people (and other mammals) who are deaf due to bilateral damage to the primary auditory cortex do retain some response to. Bilateral PAC damage. for example. more is transmitted to the opposite (contralateral) side than the side of the stimulated ear (ipsilateral). Though information is distributed bilaterally. while language is a left-brain activity. and involves several 'music centres'. along with the knowledge that bilateral damage to this area causes deafness – at least conscious deafness. an important sub-cortical structure that will be discussed shortly. which receives auditory data from the Medial Geniculate Nucleus (MGN) of the thalamus. Progress in identifying these music centres has accelerated since the older view. Brain localisation in the 20th century was focussed on cortical function. which is far less common. was challenged. that music and other ‘environmental sounds’ were exclusively the domain of the right hemisphere. It has become increasingly clear that the processing of auditory information once it reaches the forebrain is extremely complex. (though asymmetrical distribution) is that damage to the PAC on one side of the brain only causes a 20 percent or so decrease in auditory acuity.10 The first site of cortical processing of all sounds is the primary auditory cortex (PAC). The result of this bilateral. Some of these have been elucidated in recent decades with the help of functional Magnetic Resonance Imaging (fMRI) scans. with widespread use of electrical stimulation of the cortex of humans and other animals using fine electrodes. due to sub-cortical processing (this is not seen if the deafness is due to damage to the auditory nerves and the lower connections in the auditory pathways). The location of the primary auditory cortex in the temporal lobes has been known for more than a hundred years. alarming sounds. Many of the orthodoxies of neuropsychology can . Unlike visual data.
who contributed immensely to the science of brain localization until his death in 1976. This work contributed to cortical maps indicating ‘secondary auditory’ cortex in the adjacent temporal lobe gyri (notably the superior and middle temporal gyri). . Robert Zatorre. These gyri and the superior temporal sulcus. Isabel Peretz and Daniel Levitin have been pioneers in developing detailed models of music processing based on integration of recent research findings with older studies. were found to occasionally elicit complex auditory sensations. which separates them. in which Montreal played a leading role with the work of Wilder Penfield at the Montreal Neurological Institute. including Anne Blood.11 be credited to the pioneering work of the US-trained Canadian neurosurgeon Wilder Penfield (1891-1976) who reported in a series of widely-cited papers the results of systematically stimulating the surface of the brain with small electrical shocks using (then) state-of-the-art needle electrodes. It was during his preoperative investigations that he famously mapped the cortex functionally by stimulating it with a fine electrode. Since the surface of the brain has no pain sensors. More recently. also pioneered a technique for brain scar excision (when such scars were causing epileptic foci). cognitive psychologists in Canada. the neurosurgeon was able to do this in patients who were awake and therefore able to tell him what they were experiencing. Penfield. including longforgotten songs and conversations.
In an article published shortly after his death in 1976. As Milner explains. One of the icons of music neuroscience. Brenda Milner. mainly men. Milner includes diagrams. worked under Wilder Penfield at the Montreal Neurological Institute he had founded some years earlier. were conducted in the years following the Second World War on patients. Wilder Penfield had unprecedented (and subsequently unrivalled) access to patients with focal epilepsy resulting from war-related head injuries. Penfield’s studies. who had developed localised (focal) epilepsy following brain trauma. Points evoking complex auditory images in Penfield’s brain stimulation experiments. including the one reproduced above. part of the neurosurgeon’s preoperative evaluation before excising (or advising against excision of) part of the cortex. which were also used as the basis of the motor and sensory ‘homunculi’ that are essential inclusions in every introductory neuroscience text (despite criticism that they can be misleading) led to auditory cortex maps .12 Fig 3. These studies. as Canada’s most famous neurosurgeon. indicating the parts of the cortex that evoked complex auditory experiences when Penfield stimulated the area with an electrode.
the findings of Gordon (1970) and Bever and Chiarello (1974) suggested. Milner’s studies compared musical ability before and after surgery using the Seashore Measure of Musical Talents. as Werner had postulated in the 1940s. crying. working with students (again with intact brains). that differences may exist in the neural activity of trained musicians. right hemisphere) dominance for non-speech vocal sounds like laughing. This area (coloured yellow in this reproduction of the original image) only evoked simple ringing or buzzing sounds under the neurosurgeon’s stimulating electrode.13 indicating the ‘secondary’ or ‘association’ cortex as a ring around the primary auditory area in Heschl’s gyrus. Gordon found . Milner stresses that the complex auditory experiences reported by Penfield and herself were never evoked by stimulation of the primary auditory cortex. moaning and coughing. for their interpretation. who studied hemispheric dominance in people with intact brains using dichotic listening tasks. were supported by the experimental results of Kimura and others. on the assumption that most sound from the right ear is transmitted to the left hemisphere and most from the left ear to the right hemisphere. Kimura’s pioneering studies were significantly different from Milner’s because they involved healthy volunteers. and relied. also in the 1960s. In 1972 Kimura and King. a standardised test developed by the psychologist Carl Seashore in the 1930s. Again using dichotic listening tasks. Her findings. these being students and postgraduate nurses rather than people with known brain damage. which suggested a specific role of the right temporal lobe in at least some aspects of musical perception. These involved presenting different musical stimuli to the left and right ears. demonstrated left ear (and by inference. Thus Kimura’s discovery that melodies are more easily recognised when played to the left ear and speech sounds when played to the right have been interpreted as supporting Milner’s findings. Also.
large individual variations in brain activity in response to music have been demonstrated. some are more left-brained than right-brained. such that we are able to recognize the voice of a loved one as soon as we hear the first syllable they utter over the phone. most people are better at recognizing melodies with their left ear and speech in their right ear. As suspected in the pre-MRI age. that the right hemisphere is involved in ‘holistic appraisals’. To explain such findings subsequent researchers have concluded. though. There is no reason to believe that this is a unique human capacity. studies have consistently demonstrated that musical training results in a shift from the right to the left hemisphere in terms of greatest neurological activity. and fMRI and similar new technologies may cast more light on one of the most sensitive of human abilities – our capacity to discriminate between tiny differences in timbre. Numerous studies have confirmed that music training consistently increases left hemisphere activity relative to activity in the right side of the brain. though not necessarily of speech. in line with Werner’s views. . In addition. However.14 left ear preference for chords but no asymmetry in perception of melodies in professional musicians. This is an interesting area of ongoing research. ‘piecemeal’ analysis. Other animals have extremely sensitive hearing and recognition of different timbres provides obvious survival benefits (in addition to musical benefits). in both musically trained and untrained subjects. While studies continue to show that certain aspects of music cognition are lateralised. whereas the left performs the detailed. Functional MRIs have also shown greater activation in the right temporal lobe when subjects are focusing on aspects of harmony and when discriminating between different timbres of musical instruments. The ability of a trained saxophone or guitar listener to identify by their tone (meaning timbre) numerous musicians they have never seen or heard live is truly extraordinary.
. These recent studies have also demonstrated the important role of other ‘subcortical motor areas’ of the brain in appreciating and creating music.15 MRI studies have also revealed the surprising finding that that previously learning to play an instrument results in motor cortex activation in areas responsible for movements required to play the particular instrument one is familiar with. especially the basal ganglia and the cerebellum. requires a re-evaluation of conventional ideas about the brain’s motor function generally. and the role of the motor cortex in ‘generating’ voluntary movements. which has been confirmed by several researchers. even when one is not physically moving or consciously imagining playing the instrument. This finding.
including adjacent gyri in the temporal lobes that constitute the ‘secondary auditory cortex’ and areas of ‘association cortex’ that exist between the temporal lobes and occipital lobes (containing the visual cortex) and parietal lobes (containing the somato-sensory cortex). all the processing of sound between the ear and the primary auditory cortex (PAC) is ‘subcortical’. a body of grey matter at the caudal end of the thalamus. which are known to play an important role in short term memory and emotional reactions (in addition to many other mental functions). the ‘cortical processing’ begins. as is the thalamus.16 Overview of the auditory system The neural processing of information arriving through the senses is traditionally divided between ‘subcortical’ and ‘cortical’ stages. These deeper structures. to which electrical impulses return. The following model illustrates this general sequence of auditory data processing: . including those traditionally described as comprising the ‘limbic system’ and the basal ganglia are subcortical. once the sound (which has been transduced into electrochemical impulses in the inner ear) reaches the cortex. the insular cortex (located between the temporal lobes and frontal lobes) and to deeper structures. having been sent to the PAC from the Medial Geniculate Nucleus. In the auditory sense. Auditory information is also distributed to the frontal lobes. From the PAC the auditory-evoked signals are distributed to other parts of the brain.
since additional metabolic energy is contributed by the nerve cells. so it should not be seen as a pure process of electrical transduction. but first I should point out some flaws and over-simplifications in this diagram. which I drew in 1999. Transduction of the kinetic energy of the air vibrations we perceive as ‘sound’ into electrical impulses in the organ of Corti (in the cochlea of the inner ear) requires metabolic energy in the sensory neurones of the inner ear. The electrical impulses that travel along the auditory nerve and subsequent transmission along the chain of neurones that carry the auditory information through the brainstem and midbrain to the thalamus likewise require metabolic activity in the these cells.17 Fig 4. since they are created by waves of chemical ions passing in and out of cells through the . These electrical impulses are better described as electrochemical. Transduction of sound energy (original integrative theory) The flow diagram (Figure 4) above is a simplified model of how auditory impulses are transmitted through the brain. and how what we hear may affect diverse aspects of physiology through the known effects of the hypothalamus on the secretions of the pituitary and pineal glands. These will be explored later.
It is important to note that most of the synaptic activity in the brain is inhibitory rather than stimulatory.to vibrate at various frequencies to exist at all.atoms and molecules . In the human ear (and those of other animals and birds) the vibrations in the air generate movement in the eardrum (tympanic membrane) that are transmitted through the air-filled middle ear to the inner ear. . Here they result in stimulation (or inhibition) of the electrochemical activity in neurons (nerve cells) which carry the raw data along the auditory nerves to the brain. where they cause corresponding vibrations in the fluid-filled cochlea. Sound requires matter .18 semi-permeable cell walls of neurones and their axonal and dendritic processes. There are no sounds in empty space. This inhibition is essential for selective attention and to stop the brain from being flooded by sensory information.
which is shaped like a snail-shell. Anatomy of the middle and inner ear The perception of (external) music requires analysis of the information coded as electrical signals. . These electrical impulses are generated by sensory cells in the organ of Corti in the cochlea of the inner ear in response to vibrations of cilia on specialised sensory cells. It has been known for many years that the base of the cochlea. which are decoded the brain.19 Fig 5.
The reticular activating system (RAS) is known to be involved in attention and concentration (as well as conscious state) and has been extensively studied in the medical specialty of anaesthetics. This is a skill which can be developed with practice and presumably involves activity of the widely distributed network of reticular activating system neurons (amongst other things). This is obvious from everyday experience. however. These pathways have been partly identified by neurological study over the past 150 years combined with modern data from MRI. It is not as clear as to what the effect of subliminal lyrical content is. Aeration of the chamber of the middle ear is maintained by the eustacian tube.20 responds to high frequencies. which connects the middle ear with the pharynx. Background music can be irritating. although some influence on thought and behaviour would be expected. This tonotopic arrangement is retained through the hindbrain and midbrain to the thalamus and the primary auditory cortex. in addition to various mental associations. and more specifically to music. Actively listening to music requires mental focus (attention and concentration). music can have significant effects on the mind. PET and MEG scanning. the cingulate gyrus (involved . while the apex (in the centre) responds to low frequencies. Neuroimaging studies have indicated that listening to music also activates. Sound vibrations are transmitted and selectively amplified from the eardrum to the inner ear via three small connected bones (ossicles) which cross the airfilled middle ear. Listening to sound. is a more active mental process than just hearing it. in addition to the auditory cortex. probably depending on such things as the volume at which it is played and the clarity of the words. calming or energising. Even without actively listening to it. All listening obviously involves the pathways which conduct auditory signals around the brain.
the limbic structures (involved in emotional reactions) and the insular cortex. . There is also activation of the cerebellum. structures mentioned. memory (and anticipation) and movement (including reflex movement of the head and eyes towards sounds). from the cortex and from neural structures involved in such things as attention and concentration. with some nerve fibres continuing on the same (ipsilateral) side. frontal lobes (which include areas involved in working memory). also in the brainstem. basal ganglia. functionally precortical. It is possible that this subcortical. The main auditory structures in the midbrain are the lateral lemniscus and inferior colliculus. thalamus and other subcortical structures that will be discussed shortly. There are several stages of relay and neural processing between the auditory nerves and the medial geniculate nucleus (MGN) of the thalamus. This diagram and the one on the next page show the location of the subcortical.21 in decisions. pre-cortical neural activity contributes to the distinctly (and almost uniquely) human tendency to move rhythmically to the beat of music – what may be called our dance instinct. including the decision to listen). These relay the auditory impulses to the MGN. These include the cochlear nuclei in the brainstem which project to the superior olivary nuclei. while others cross to the opposite (contralateral) side. These subcortical structures do not function as mere relay stations – they are involved in processing the ‘sound’ in various ways and are subject to feedback from each other. The cochlear nuclei and olivary nuclei project to neural structures in the midbrain in a complex way.
This structure is involved in reflex movements of the head. they are thought to be more widely distributed to the outermost (Layer I) of the cortex. . There are also smaller numbers of auditory fibres outside the lateral lemniscus.22 Fig 6. These extra-lemniscal fibres synapse in other parts of the thalamus (other than the MGN) and rather than projecting to Layer IV of the PAC. Some of the neurons and axons in the lateral lemniscus (which contains grey and white matter) synapse in the inferior colliculus. Layer I projects to deeper and to adjacent areas of cortex. Other axons continue past the colliculus to synapse in the core of the MGN. Precortical auditory structures Primary Auditory Cortex (PAC) medial geniculate nucleus (MGN) inferior colliculus lateral lemniscus superior olivary nucleus cochlear nucleus The medial geniculate nucleus is divided into an inner core and an outer shell. especially alarming ones. the eyes and attention towards sounds. which in turn project back to the thalamus and to other parts of the brain.
others make output connections with the basal ganglia. The vast majority though. One such tract. These can be distinguished under the microscope by the relative density of different cell types. Most of these connections are with adjacent gyri. limbic structures and input ganglia of the cerebellum. is composed of six interconnected layers. Layers III and V contain pyramidal cells. Layer V. medial geniculate nucleus inferior colliculus lateral lemniscus cochlear nucleus superior olivary nucleus The auditory cortex. though long tracts of white matter conduct neural impulses between the various lobes of the brain.23 Fig 7. is the ‘output’ layer of the cortex – long cell processes (axons) emerge from this layer to make connections with other cortical areas and with deeper structures. the inner pyramidal layer. Some leave the brain to synapse in the spinal cord. Location of important precortical neural structures in the auditory system. carry information to other areas of the cortex. like other neocortical areas. connecting the temporal and frontal lobes with particular relevance to . while layers II and IV contain smaller stellate cells that appear granular when stained with methylene blue.
This is the case in primary somatosensory cortex (in the parietal lobes). 8. The location of these other primary sensory areas and their main connections is relevant to understanding the neural processing of music and the cross-modal associations integral to evaluation. primary visual cortex (in the occipital lobes) and the primary auditory cortex. The relative thickness of the different layers gives an indication of what a particular area of cortex does. .24 music perception. in which the overlying parts of the frontal. Primary sensory areas of cortex are characterised by a thicker layer IV populated with numerous small granular neurones. appreciation and verbal description of music. is shown in Fig. parietal and temporal lobes have been removed: Fig.8 White matter tract connecting frontal and temporal lobes.
This deep location may protect the important senses of vision and hearing from trauma to the outside of the head. . the egg-shaped structure outlined in the middle of the brain shows the location of the thalamus. Areas of primary sensory cortex. It can be seen that most of the primary visual cortex is on the medial side of the brain (around the calcarine sulcus). being buried in the sylvian fissure. Like the primary visual cortex.25 Fig 9. Somato-sensory cortex in parietal lobe Auditory cortex in temporal lobe Sylvian fissure Visual cortex in occipital lobe Calcarine sulcus In figure 9. most of the primary auditory cortex is not on the outer surface of the brain.
cats. which shows progressive increase in size and convolutions across increasingly intelligent primates.26 Fig 10. The secondary auditory cortex extends to the middle temporal gyrus and the angular gyrus. this mode of research contributed little to understanding the complexity of human emotions and less to an understanding of the neural processing of music (not least because rats. Cats and monkeys bore the brunt of this horribly cruel neurophysiology and neuroanatomy research. Those with an inclination towards physiology spent many decades engaged in grotesque ‘brain reduction’ experiments in an effort to identify what the deeper structures did. . angular gyrus Primary Auditory Cortex (PAC) superior temporal gyrus middle temporal gyrus superior temporal sulcus Cortical and Subcortical Processing of Music Because the size of the cortex was thought to be closely allied with intelligence (with some reason) the outer layer of the brain. While confirming that deep brain structures were essential to emotions and other mental faculties including memory. The primary auditory cortex extends deep into the transverse (Sylvian) fissure that separates the temporal lobe from the frontal lobe. dogs and monkeys do not respond to music remotely like humans). has long been the primary focus of neuroscientists with an inclination towards psychology.
like the rest of the neocortex. Directly under the cortical layer are dense tracts of white matter – the axons of nerve cells projecting to neurons in the cortex and those emerging (descending) from the outer cortical layer of the brain. in the sense that they are well-developed in ‘subhuman’ species. motivation and emotions become possible in humans as much as in other animals. Cortical processing of music The auditory cortex. movement. It is through the constant interaction of the cortex with these structures that perception. basal ganglia and thalamus are more primitive. about half a centimetre thick and consists of six thoroughly studied layers. For this reason it may make more sense to refer to ‘precortical’ and ‘post-cortical’ processing when discussing ‘sub-cortical’ brain activity. within the first transverse temporal gyrus (Heschl’s gyrus). such as the cerebellum. to ‘reason’. It is true that these structures. directly or indirectly. These layers have complex connections with each other and with all parts of the brain and the body. This reflects a long-held prejudice in the neurosciences that emotions are antithetical. is.27 One unfortunate consequence of the division between cortical and ‘subcortical’ processing is a spurious idea that cortical function is where ‘intelligence’ resides and that the ‘subcortical structures’ are both less important and more primitive. It has been known for a century that the primary auditory cortex (PAC) – the first cortical area to receive auditory data from the thalamus – is located in the upper temporal lobes. It is certainly not true that they are less important than the cortical layers of the brain. and inferior. In terms of music perception the auditory cortex is an early processing area – it is what happens after the auditory signals reach the PAC that determines whether we perceive the sound as music or noise. and whether we are motivated to listen to (or escape) the music. From the PAC sound signals are transmitted laterally to the secondary .
and subcortical structures including the basal ganglia. This granularity is greatest in the external and internal ‘granular layers’ (layers II and IV) of cortex. Fig 11. . (reproduced from Morosan et al 2001) This image is from a 2001 paper by German researchers from the Vogt Institute of Brain Research published in the journal Neuroimage. Also.2 areas (blue and light green). layers III and V – the external and internal pyramidal layers – are characterised by larger. Te 1. Their study (on postmortem human brains) showed that though the primary auditory cortex is contained within Heschl’s gyrus as has long been known. arrive at the cortex from relay neurons in the thalamus. midbrain and brain stem. thalamus.0 (dark green) is highly granular. An MRI showing the primary auditory cortex (PAC) in horizontal section.28 auditory cortex which extends through the superior temporal gyrus to the middle temporal gyrus. The granularity of cortical layers is an indicator of sensory inputs. other than smell (olfaction). indicating this area receives the auditory signals before they spread to the less granular Te 1.1 and Te 1. examination of the cell types in the cortical layers suggests that considerable variation exists between people in the extent of what is assumed to be primary auditory cortex due to its high concentration of granular neurons (koniocortex). and data from all the senses. agranular ‘pyramidal’ neurons with long axons that carry information from a particular area of cortex to other cortical areas on both sides of the brain.
Te 1. Functional MRIs in living people have confirmed the location of auditory areas in the temporal lobes bilaterally and that listening to (or just hearing) sounds increases blood flow in Heschl’s gyrus. also known as Heschl’s gyrus. though.2) do not conform precisely to the visible landmarks on the surface of the brain. as might be expected. Broca’s area (speech generation) Insular cortex Heschl’s gyrus Insular cortex Heschl’s gyrus (location of PAC) (PAC) Basal ganglia Thalamus Wernicke’s area (speech comprehension) Fig 12. The study did confirm. The precise areas have come as something of a surprise for . that the primary auditory cortex is located where it has been thought to be for a century – the first transverse temporal gyrus. Listening to music also increases blood flow (which is what is measured in fMRI) in many other parts of the brain.1 and Te 1.29 these areas (Te 1.0.
30 those who underestimated the importance of ‘subcortical brain structures’ when it comes to what many would regard as the pinnacles of human cultural achievement – our music. human activities driven by aesthetic and a sense of delight in beauty. Research in recent years has shown the insula to play an important role in music . Location of the PAC relative to the insular cortex (insula) Insula Primary Auditory Cortex (Heschl’s gyrus) The primary auditory cortex (PAC) occupies the first transverse gyrus of the temporal lobes bilaterally. literature and art. also known as Heschl’s gyrus. Fig 13. in the diagram above parts of the (left) parietal and frontal lobe that form the operculum (‘lid’) have been removed. is buried in the sylvian fissure that separates the temporal and frontal lobes. Most of this gyrus. exposing the first transverse gyrus and the intriguing area of cortex immediately anterior to it – the insula.
essential to the complex auditory perceptions involved in listening to music. The fact that there is separation of aesthetic appreciation in different elements of music can be seen in the common experience of liking some aspects of a piece of music and not others – one can like the groove but not like the timbre of particular instruments or the singing. . This is the case with the basal ganglia. This mirrors the tonotopic arrangement of sensory fibres in the cochlea of the inner ear and their connections through the lower part of the brain through to the thalamus. One may like the melody.31 perception. of course. including complex emotional reactions to various aspects of music. These higher functions are. thalamus and the ‘limbic structures’ which appear to play important roles in post-cortical music processing. or parts of it. They are also of vital importance when considering appreciating music – something that requires emotional reactions. It has been known for many years that the nerve cells in the PAC are tonotopically arranged – neurons of the medial part respond to high frequencies (high sounds – shown in blue) while the lateral part of the gyrus responds more to lower frequency sounds (shown in red). Brain structures that are subcortical are not necessarily functionally precortical. How this tonotopic arrangement relates to higher auditory functions – to secondary and subsequent processing of sound – remains the subject of considerable uncertainty and debate. but find the rhythm boring – and any number of permutations and combinations of likes and dislikes that affect overall (or gestalt) aesthetic judgement of the music. something that was not suspected in the past. Here the limitation of thinking about music neuroscience in terms of ‘cortical’ and ‘subcortical’ processing becomes clear. appreciation and creation. This observation is congruent with what is now known about the neural processing of various elements of music.
The input to the PAC originates in the Medial Geniculate Nucleus (MGN) of the thalamus. These include cortico-cortical connections and postcortical connections with subcortical structures including the basal ganglia and thalamus. Outputs from the PAC project to the secondary auditory cortex (located in the superior temporal gyrus.32 Post-cortical processing of music Pre-cortical Primary auditory cortex Auditory signals Secondary auditory cortex Temporo-parietal association cortex Temporo-frontal association cortex Insular cortex Basal ganglia – including amygdalae and nuclei accumbens Motor cortex Temporo-occipital association cortex Thalamus Cerebellum Fig 13. superior temporal sulcus and middle temporal gyrus). but other cortical areas also project to these structures (via the . Cortical and post-cortical processing The diagram (figure 13) above shows some of the main connections of the primary auditory cortex. The diagram (figure 13) only shows the motor cortex’s connection with the basal ganglia. which projects to layer IV of the PAC. For simplicity the important connections with limbic structures (involved in emotional reactions) are not shown. The secondary auditory cortex projects to other areas of cortex and back to the thalamus.
These association areas are likely to play a key role in the perception of music and their connections with the basal ganglia. cerebellum and limbic system appear to be involved in various emotional reactions to music. including pitch.33 caudate nucleus and putamen. melody. somatosensory and visual areas. with and without words. the main input nuclei of the basal ganglia) as well as to the thalamus. melody and rhythm suggest cross-modal mental associations. It seems to reasonable to infer that mental associations in perception and memory of timbre. The thalamus also has strong reciprocal connections with the association cortices – the areas of cortex between the primary and secondary auditory. Mental associations with Music and their Neural Substrate Mental associations are fundamental to appreciation of music. temporal and parietal lobes between the primary sensory areas that occupy much smaller areas of these lobes. Several (English language) terms used to describe musical characteristics. harmony. including those of the sensory association cortex – the large areas of cortex in the occipital. Some of these associations suggest connections (functionally and anatomically) between the temporal lobe auditory cortex and other sensory cortices – the visual cortex in the occipital lobe and somato- . however the vital nature of mental associations in perceiving. rhythm and harmony reflect neural connections. That they are fundamental to understanding the lyrical content of music is self-evident. timbre. appreciating. These outputs from the cortex to the thalamus project to a number of nuclei within the complex structure which will be explored in more detail later. understanding and creating music is less widely appreciated.
In addition. Visual associations in music are obvious and easily identified: High notes/low notes (pitch) Bright/dull tone (timbre) Bright melody Pretty melody (we don’t say something tastes or feels ‘pretty’) Brilliant musicianship/solo/performance The visual cortex is also necessarily involved in reading music notation and chord charts. visual information augments somatosensory (including kinaesthetic and proprioceptive) information relevant to movements of the limbs when playing music and dancing. Bass) Hot solo Warm sound Cool organ/guitar (e.g. Others suggest associations with more abstract and ‘intellectual’ concepts that are more suggestive of connections between the temporal and frontal lobes. Somato-sensory (parietal) association examples include: Sharp (pitch) – can also have visual connotations Flat (pitch) – several meanings – visual connotations also Smooth sound/tone (timbre/gestalt) Steady rhythm Hard sound regular Soft Light (also visual) Heavy bass Thick sound (esp. chorus pedal) Contour of melody (also visual associations) .34 sensory cortex in the parietal lobes.
constantly influencing the emotional state of the person listening to the music. such as tight. loose. Medium and long term memories are also involved in the moment-to-moment evaluation of music that is listened to. These include more abstract concepts.35 Other mental associations are suggestive of connections between the auditory temporal lobes and the frontal lobes. fear and anger) and other areas known to be involved in emotional reactions. melancholy. the amygdala (deep within the temporal lobes. aesthetic/emotional terms used to describe music are also used to describe other sensory modalities: beautiful. etc. parallel processing of different aspects of timbre. It may be more accurate to say that the gestalt occurs simultaneously with sub-processing of different aspects of music. ugly. harmony (the ‘elements’ of music) occurs before they are brought together as a gestalt. sad. Any time music is described in words. pretty. melody. music is devoid of melody and rhythm. bold and other adjectives. The fact that music seems to be processed in piecemeal fashion though producing a gestalt – the piece of music can be appreciated as a whole – has led to a multi-modal view of music neuropsychology. happy. activity in Broca’s area in the left frontal lobe is necessary. These will be discussed shortly. PET and MEG studies have shown activation in deep structures including the nucleus accumbens (part of the basal ganglia associated with feelings of pleasure). associated with excitement. modern MRI. Often. Satisfying and skilfully manipulating the listener’s search for tonal resolution based on their musical memories is the hallmark of good musicians everywhere – a subconscious rather than a conscious quest in most. The frontal lobes are also involved in immediate (‘working’) memory. tempo. rhythm. which is essential to enjoying and appreciating music – if you can’t remember the notes and beats that just finished. Memoric associations caused by prior exposure to music shape expectations of the sounds to come. Separate. These infer connections between the auditory sense and limbic structures. .
The insular cortex has been known for many years to be involved in taste – as in the gustatory sensation produced by various chemicals stimulating sensory receptors on the surface of the tongue. from the 1960s onwards more and more evidence pointed to it being a highly developed area of cortex with several interesting functions. including some related to music. taste in literature. could the insula be one of these structures? . or Island of Reil. taste in art. But the word ‘taste’ is also used in reference to various aesthetic preferences – taste in music. Then. Could this reflect shared neural circuitry between brain structures involved in the different senses? If this is the case.36 Gestalt Listening to music Emotional reaction Focus on lyric Focus on melody Focus on individual instruments Focus on harmony Focus on timbre Focus on rhythm Fig 15. was considered to be of little importance. Model of selective processing of musical elements Is the Insula involved in musical and other aesthetic taste? For many years the insula. The diminution of taste when smell is prevented (by blocking the nose) indicates the role that olfaction plays in augmenting taste.
Anterior Insula Posterior Insula Primary Auditory Cortex Fig.37 Central Sulcus separating Frontal and Parietal lobes.18: Left hemisphere with frontal and parietal parts of operculum removed to show the insula and primary auditory cortex (PAC) The anterior insula (including the insular cortex and underlying white matter) is located behind (caudal to) the primary motor area for speech (Broca’s Area) which occupies the frontal lobe section of the operculum and the gyrus in front of (rostral to) it: .
These gyri were identified as home to the primary auditory cortex in the early twentieth century.38 Fig. Recent discoveries about the many functions of the insula may fill some of the gaps in our knowledge about how we perceive and respond to music (and other .19: horizontal section of brain at the level of Heschl’s gyrus Broca’s Area – primary speech motor area Anterior Insula Posterior Insula Heschl’s gyrus – location of primary auditory cortex The posterior insula is located adjacent to the primary auditory cortex located in the first transverse gyri. however what happens to the auditory signals when and after they arrive in Heschl’s gyri remains subject to debate. also known as ‘Heschl’s gyri’ after the German anatomist Richard Heschl (1824-1881) who is credited with their first scientific description.
3-4 short sulci in the anterior insula and 2-3 long sulci in the posterior insula. ‘Opposite hemispheric lateralization effects during speaking and singing at motor cortex. including the nuclei accumbens and amygdalae are of particular relevance to the insula and its possible role in what we call ‘taste’ in music. with the left insula more activated by verbal activity. Studies have also shown other signs of functional asymmetry between left and right sides. The insulae show left-right asymmetry but usually have 5 to 7 sulci. 2000). because of their extensive connections with this multimodal area of cortex.39 sounds) and how the primary auditory cortex works in concert with neighbouring parts of the brain. with the right insula activated with singing out loud but not silently (Riecker. The reported study. while the right is involved in auditory sequencing and sound movement detection. Recent neuroimaging studies have shown the insulae to play key roles in music cognition. insula and cerebellum’ was published in the journal Cognitive Neuroscience in June 2000. Of these. According to these authors. A 2003 literature review by Bamiou. including that of rhythm. by Axel Riecker and colleagues at the University of Stuttgart in Germany is one of hundreds of scientific papers revealing an extraordinary range of activities that are accompanied by increased blood flow and metabolic activity in the insula. the left insula is thought to be involved in musical rhythm processing. Other more distant parts of the brain have also been implicated in the neural processing of music by fMRI. Musiek and Luxon (of the National Hospital for Neurology and Neurosurgery in London and the University of Connecticut) refers to several findings indicating the importance of the insulae in music cognition on the basis of PET and fMRI scans. PET and magnetoencephalographic (MEG) studies in recent years. the basal ganglia. These showed differences between the left and right insula in auditory processing. This study used .
indeed mundane. task revealed fascinating inter-hemispheric differences in blood flow between silent and overt singing and silent and overt speaking. The left anterior insula was activated during the verbal task of reciting the months of the year. changing. their motor function may have obscured their important role in mediating emotions. This simple. their relative contributions to voluntary movement (including vocalisations) may have been exaggerated relative to that of the insulae.40 functional MRI to compare changes in cerebral blood flow in eighteen healthy right-handed German men and women aged between 22 and 63. meaning the motor cortex. Silent (covert) speech resulted in an opposite hemispheric localisation – activation of the left motor cortex and right cerebellum. also. basal ganglia and cerebellum. and singing with or without words. The activation of the insulae with overt but not covert singing and speaking raises doubts about the classical ‘motor circuits’ in the brain. This cortical area is the right-sided equivalent of Broca’s area. and left cortex and right cerebellum with speech) but also hemispherically asymmetrical activation of the insulae. given that Broca’s speech area lies in the adjoining frontal cortex extending into the frontal part of the operculum (the ‘lid’ covering the insula). The musical task of singing the notes of a familiar German tune was compared to the verbal task of reciting the months of the year from January to December. Importantly the researchers compared the blood flow when these tasks were performed overtly and silently. Silent (covert) singing revealed exclusive activation of the right motor cortex/posterior inferior frontal gyrus as well as the left cerebellar hemisphere. quietly or at the top of one’s voice. Though these structures are certainly involved in initiating. correcting and learning movements. . silently. including the plethora of feelings evoked by listening to and playing musical instruments. The role of the left anterior insula in spoken language is not surprising. the right insula with singing the tune (without words). Overt singing and speaking resulted in bilateral activations (though greater in the right cortex and left cerebellum with singing.
The entire debate may be based on a spurious dichotomy. the theory was put forward by Heinz Werner (1890-1964) that a shift in activity occurs in musically experienced listeners to involve the whole of the brain (or towards the left side) from being a mainly right-sided function in inexperienced listeners. These findings further refine the long-held view that the right hemisphere is more involved with music while the left is more involved with words. since the ‘line’ between music.41 Another study of the insula in 2004 by Patrick Wong and colleagues at the University of Texas used Positron Emission Tomography (PET) scanning. the homologous area in the right hemisphere activated. Even more recently . His explanation was that musically experienced listeners have learned to perceive a melody as an articulated set of relations among components rather than as a whole. and may reveal profound insights into the development of the brain and its capacity for what has been termed ‘plasticity’. comparing native speakers of Mandarin Chinese versus English. but when they discriminated pitch patterns embedded in English words. The left anterior insula showed increased activity when ‘Mandarin listeners discriminated pitch embedded in Mandarin lexical tones’. who show more right hemisphere activity. Neural plasticity and development of the auditory system In the 1940s. to show that the left anterior insula is implicated in ‘pitch pattern perception’ with a linguistic context. as it did in Englishspeaking listeners discriminating pitch patterns embedded in either Mandarin or English words. Werner’s theory gained support from subsequent neuroimaging and neuropsychological studies suggesting greater left hemisphere activity in trained professional musicians than in amateurs. singing and speaking may not be as definite as has been traditionally supposed. This observation is interesting.
While new synaptic connections are forming through the growth of axons and dendrons. Loss of brain cells was thought to be irreversible and loss of function due to death of cells was regarded as permanent. the cells that survive can continue to grow throughout life. Our potential to create new synaptic connections throughout life and to improve the efficiency of those already established have profound implications for learning and the development of strategies to prevent dementia. Though there is an overall loss of neurons from early childhood. however. mental ability. other synapses and cells are being lost. parietal or temporal lobes developing vision-processing capability. For many years it was assumed that once the brain has grown to adult size no fundamental changes occur in its structure and function other than gradual (or sudden) loss of brain cells. greater plasticity is likely than in vision. . Plasticity of the brain (and neurons) allows for cells previously involved in one type of activity to assume a different function in the event of damage to the cells that usually perform this function. with them.42 studies on children with a rare genetic syndrome called William’s Syndrome (WS) has thrown Werner’s conclusions about holistic processing in the right hemisphere into question. similar limitations doubtless exist. They continue to make new axonal and dendritic connections that link the electrically active cells in a dynamic (and potentially expanding) network. The key factor determining overall gain or loss appears to be use: the principle of ‘use it or lose it’ applies to the brain as well as other parts of the body. No cases have been recorded of people who are blind because of damage to their occipital lobes (visual cortex) gaining their sight by their frontal. and. however. In the instance of musical perception and creation. This is now known not to be entirely true. This phenomenon is a limited one.
aspects of neural plasticity exist. secreted by specialised cells that wrap the fatty material around the axons of neurons. including fMRI. and it has been noted that loss of white matter precedes loss of grey matter in age-related loss of brain tissue. insulates the nerve fibres and also greatly speeds up the transmission of electricity along the nerve. affecting how easily they conduct impulses. but equally important. Myelination of the white matter of the brain continues through life. Recent studies by Peter Schneider and co-workers at Heidelberg University in Germany of several children with a . have confirmed an increase in left sided brain activity in trained musicians. The myelin of the white matter of the brain and nerves. This is thought to be a result of musical training rather than because people with large corpus callosa are more likely to become musicians. Focal demyelination of nerves and white matter of the brain is also the characteristic neuropathological finding in multiple sclerosis (MS). the observation does suggest that learning to play musical instruments may protect against loss of brain function later in life. is larger in trained musicians. This loss has been correlated with declining mental powers with age. Another area of the brain that has been shown to be larger in trained musicians and talented music students in the left auditory cortex – specifically Heschl’s gyrus. This involves a form of plasticity of the synapses. It has been debated as to whether this is a reflection of different brain structure due to innate giftedness or the result of growth of this area due to musical training. along with structural changes in the brain. Recent research. Myelination of nerves.43 Other less dramatic. One consistent finding is that the corpus callosum. which has been shown to be up to twice as large in trained musicians as in non-musicians. the large tract of white matter that connects the two hemispheres. The fact that efficient communication between motor functions of the hemispheres is important when playing instruments with both hands makes this finding less surprising. though not generally called ‘plasticity’. affects the speed and efficiency of neural transmission. Neural pathways that are used frequently become more easily activated.
unlike the sample of matched controls. In an interesting 2010 study. The missing fundamental is an interesting mental phenomenon whereby the fundamental note (pitch) is perceived as present due to the presence of the spectrum of natural harmonics above the note. even when the actual note is missing. It is one of several interesting auditory illusions relevant to music. and furthermore have suggested that the left hemisphere is more involved in holistic appraisals. which is caused by a defect in Chromosome 7 and is a rare cause of intellectual and motor disability in children. Perceiving the missing fundamental is the key factor in determining whether subjects are classified as ‘holistic’ or ‘spectral’ listeners according to the classification system used by Schneider and co-workers. who wrote an extensive thesis on music and sound perception in the 1860s. This is a comparable increase to the upper range of changes seen in trained musicians – earlier . Schneider and co-workers used fMRI. in terms of blood flow (on MRI) and magnetic field generated (on MEG) in the left temporal lobe. Despite their intellectual and motor deficits.44 rare developmental problem known as William’s Syndrome have pointed to nature rather than nurture. This study showed that. Schneider writes that what he calls ‘holistic’ used to be called a ‘fundamental mode’ while the ‘spectral mode’ used to be called the ‘analytical mode’ by earlier authors on the subject. children with William’s Syndrome showed extremes of a ‘holistic’ (versus ‘spectral’) mode of music perception. What is going on here? Could it be a matter of terminology? Possibly. including the famous German physician and physicist Hermann von Helmholtz (1821-1894). which are accompanied by diffuse loss of neurons and white matter especially in the parietal and occipital lobes. while the right is more involved in perceiving the harmonic spectrum. accompanied by increase in activity. children with Williams Syndrome are typically very sociable and musical. MEG and psychoacoustic tests to evaluate the exceptional musicality of children with Williams Syndrome. specifically in Heschl’s gyrus and the areas of cortex adjoining it.
including structures involved with emotions. and continues through life. Plasticity of this sort is the basis of auditory learning generally. including musical experiences. . such as the amygdalae. with an increased number of neurons in this part of the brain. This forms the basic structure of the brain and determines how various collections of cell bodies (grey matter) are connected with each other. It is a common observation that the experience of listening to a lot of music changes what and how you hear. to shape the developing brain.45 studies by Peter Schneider at Heidelberg have demonstrated increased blood flow and electrical activity (on EEG and MEG) in the left auditory cortex of trained musicians and gifted music students. Neural plasticity is also of fundamental importance in memory and learning during the ontological development of the brain. Schneider leans toward nature over nurture and mentions postmortem studies demonstrating that individuals with Williams Syndrome have an enlarged primary auditory cortex. the latter is supported by introspective analysis of changes in musicality with increased experience. and provide an obvious mechanism for pre-natal experiences. despite having less in others. The connections established by the growth of axons from the developing nerve cells in the embryonic brain become the tracts of white matter. which is more convoluted than is usual. studies have indicated that musical training causes progressive changes in the structure and function of the brain. He also mentions that postmortem studies have also shown relative preservation of other areas of the brain with Williams Syndrome. is established by the migration of cells from the inner wall of the developing neural tube. and the way the layers of cortex are connected to subcortical brain structures. In the developing foetus the structure of the brain. At the same time.
A shift from right to left sided activity would be consistent with the view that the right hemisphere is involved with more ‘holistic’ or ‘gestalt’ perceptions than the left. At this stage. according to contemporary thinking.46 The auditory system. Perhaps the apparent contradiction can be explained by the authors’ different use of the term ‘holistic’. Schneider and co-workers seem to be saying the opposite – that trained musicians use more of the holistic mode of the left side of the brain. timbre and other qualities of music when listening to it. and this stands to reason. mean what is not needed. continues to develop rapidly in the first two years of life. with most people somewhere in the middle. Back in the 1860s Helmholtz observed that it is possible to focus on individual elements and aspects of music than rather than the melody as defined by movement of the fundamental tone alone. however. What is not used does not. rhythm. with different results in different cultures. It is generally accepted that those connections that are not used are pruned. This is a significantly different mental activity from listening to a piece of music as a whole. though functional in a third trimester foetus. as has long been said. Werner identified a possible cause of this in terms of a changed focus to parts of the melody rather than the melody as a whole. It may be that music education and training lead people in different directions. It is possible to focus one’s attention on elements of harmony. as the Heidelberg research suggests. Pruning results in loss of nerve cells and the processes (axons and dendrons) that grow out of the cell bodies. It may be that the ‘natural’ way of listening varies between two polarities. . A shift from right to left sided brain activity with musical training suggests that change in the way people think about music can result in physiological and structural changes in the brain. This is an interesting field for future research. the process of ‘pruning’ begins.
People may learn to read written music. equivalent to the grammar and syntax of written language. plasticity in the brain and nervous system emphasizes the benefits of experiential learning in music – practicing listening and playing . as in other matters. This may have a direct bearing on the described shift from right to left hemispheric activity in trained musicians. The left hemisphere activation with music in children with William’s syndrome may be due to use of shared neural structures in music and language. have particular meanings. Some people who have great musical virtuosity in playing instruments or singing are poor readers or completely unable to read musical notation. decided on and defined by convention and common use. develop. but be unable to ‘play by ear’ or improvise. reading music and thinking about music in terms of musical symbols and theory are fundamentally different from listening to and thinking about music without theoretical knowledge. Music has linguistic elements and spoken language has musical elements. Education in music.47 Another aspect of musical thinking that may explain a shift towards the left hemisphere relates to language. They may develop virtuosity on one instrument but be unable to play another with which they are unfamiliar. even among those who play musical instruments. Training in music is a complex subject and people can be trained in very different ways. and feel great love for music without playing any musical instruments themselves. Particular symbols. Reading and writing music involves learning a language (or languages) as specific and complex as verbal language. in order to fluently read and write musical notation (script). It may be that as people are trained in music ‘linguistic’ aspects of musical thought become more pronounced. People can learn to be extremely discriminating listeners. Maybe it merely suggests that for these children music is just another means of communication – another language. may be broad or narrow. On a practical note. One has to learn the specific rules of a particular musical language. It is a skill few people. In this way.
2006). more so than how long it had gone on for or how much daily exposure to music the dreamer had. when they are young – preferably before they are 5 or 6 years old. until recently. especially emotional reactions to rhythms within music. nurture and the exercise of volition to engage in musical activities. however. It is well known that with practice what was once difficult becomes easy.48 musical instruments fundamentally changes the structure and function of the brain for the better. This is partly to do with traditions embedded in European classical music and a view that ‘serious’ (and therefore ‘good’) music should be food for the intellect rather than the emotions (a reflection of the old dichotomy between ‘thinking’ and ‘feeling’). On the neurobiology of rhythm Of the main elements of music (rhythm. An interesting paper by Valeria Uga and co-workers on music in dreams emphasizes the importance of early music training. of emotional reactions to music. harmony and timbre) rhythm has been relatively neglected by psychologists and neuroscientists over the years. These authors suggest that early training in music also increases ambidexterity if introduced before the age of 7 and absolute (‘perfect’) pitch before the age of 9 (Uga et al. There has also been neglect. Their study. comparable to the importance of spoken language in early childhood development. This makes it imperative that children be introduced to good quality musical instruments of a sufficient variety that their particular taste and aptitude for music can be nurtured. the development of musicality is a combination of nature. The younger a person is when they develop their musicality the easier it is. or at least easier. Though musicians and their audiences were certainly meant to feel . melody. showed that reporting musical dreams correlated closely with the age at which musical training began. comparing professional musicians and non-musicians in Italy. Of course.
and each discipline or faculty provides different ladders. each with its own ‘classical’ musical heritage to honour and proclaim. since the scientific study of the brain and mind is centralised in the global university system. This is not necessarily a bad thing – the academic tradition has a lot to be proud of. or opinions and interests of particular academics who have established empires at that university. The highest echelons of these faculties. This is dominated by the oldest universities in the biggest cities of the world’s most vociferous nations. Until fairly recently these men were among the minority of the world’s population that listened largely. especially in science. however. This is in stark contrast to the medical profession. like old men. Like other hierarchical institutions. but also in the arts. It can. the oldest universities tend to be the most conservative. especially if they threaten the existing curriculum. It should be remembered that until relatively recently many of the world’s more famous orchestras were composed entirely of men (with the interesting exception of harpists). Universities are conservative institutions. lead to a narrowness of view and reluctance to embrace new ideas. The influence of Western classical music and this particular tradition’s great composers on research into music perception and creation and on the neurosciences generally is to be expected. Since the 1960s and the ground-breaking work of Brenda Milner at McGill University. meaning tradition-bound (rather than any economic or political connotation). there is a ladder to climb in universities. and. including the specialties of neurology and psychiatry (the two specialities that might be expected to have . is dominated by men.49 emotions at a classical music concert. The research projects they approved were only those they could defend as studying ‘serious’ music. and to conserve. it was not good form to jump around with excitement – that was for the ‘common folk’ with their ‘folk music’. or exclusively. women have been at the forefront of music neuropsychology. to ‘classical music’.
which has been especially concerned with those attributes that have been regarded as uniquely human – language. is used to fund music neuropsychology or music therapy research. biochemistry and biophysics (physiology). and moving our bodies in time to the music. and to the sounds that make us feel like tapping our and feet. and its propensity to induce people to gyrate their bodies in sexually suggestive ways that rhythm has long . the use of psychological tests developed for other reasons being adapted for research into the neuroscience of music has resulted in confusing and contradictory findings. Harmony and melody may make us smile. which are dominated by men. The fossil record has underpinned the development of the modern neuroscience paradigm. bobbing our heads. Chauvinism may be a relevant factor in the shameful neglect of music as a therapy and as a subject worthy of serious scientific study (and funding) by the medical profession over the past half century. Unfortunately. evolutionary biology. The paradigm of modern neuroscience has emerged from a sometimes uncomfortable integration of psychology. especially when it comes to the emotional reactions people have to music. It is precisely because of its ‘primitive’ appeal. and the other major sources of medical research funding – the drug companies – are hardly likely to sponsor research that might establish music as a serious competitor in the therapeutic market. Human sensitivity to timbre and rhythm are fundamental to our musicality. Though music has been regarded as one of a relatively small number of ‘uniquely human’ abilities. and rhythm within music.50 an interest in music and the brain). but timbre and rhythm make us dance. and ‘intelligence’ as measured by various ‘behavioural’ tests in animals and ‘psychological’ tests in humans. if any. especially in terms of senior academic and research positions. in Australia and many other countries little public money. psychiatry and neurology with pharmacology.
never drums. the popularity of the guitar. much as several previous generations had disparaged jazz. followed by the fanatic enthusiasm for Elvis Presley and later the Beatles and Rolling Stones brought Berry’s ‘rock’ version of the ‘rhythm and blues’ to a vast new global audience. When these pop musicians were first featured on television variety shows and the radio was playing their recordings there was predictable opposition from many conservative members of the older generation who complained that the new music was loud noise. It’s not that great grooves began with Chuck Berry and Rock and Roll. black magic. or at worst trumpets. not the first bastions to crack under the sonic onslaught of Rock. more so than any other instrument. in particular. Though music schools were slow to add jazz studies and its focus on improvisation to their curricula. But the popularity of Berry. prior to the international explosion of Rock and Roll. Drumming was associated with voodoo. bass and drums and form their own bands in towns and cities around the world. It was only in the 1980s that the Western world’s universities evolved musically to a point where the senior echelons of the relevant faculties grew up on pop. Rock changed everything when it came to what music and musicians were venerated – but the neuroscience establishments were. guaranteed that even academia had to yield to the rhythm oriented music that has since transformed human culture – in the north. predictably. rock and jazz. all about rhythm. and is. Rock music was. The angels in religious paintings over the centuries were usually depicted playing harps. inspired huge numbers of young people to take up the guitar. black people and the dangerous allure of hypnotic trances. which is distinguished by complex. the south. syncopated rhythms in the melodies and in the ‘rhythm section’ (bass and drums).51 been regarded as the least respectable of the elements of music. The Beatles. the east and . Darkskinned people dancing semi-naked around an open fire in a frenzy inspired by sweat-covered men beating drums was a familiar image Hollywood reserved to depict the ‘savages’ in ‘Darkest Africa’ through the 1950s and 1960s. and slower to add rock and roll.
later electric) and a drum kit. all of which contributed to the synthesis Chuck Berry popularised as Rock and Roll) are bass (first acoustic. All around the world musicians have been incorporating guitars into their music. In the past two decades several facts have been established about the neural processing of pitch. (non-musical) movement and the complex circuits and neural structures that subserve our emotions. One thing that has become clear is that though small areas of the brain are primarily involved with music-related functions (mainly in the right temporal lobe).52 the west. Unlike older investigations. such as surface (scalp) electrical readings (EEGs). including language. to people with localised brain . The new generation of neuroscientists. using a combination of new and older radiological techniques and a human experimental population ranging from professional musicians through students and members of the general public. most music cognition and creation involves parts of the organ and networks that are used for other. brought up on a diet of beat-oriented music has belatedly begun a serious. these new (and relatively expensive) scans allow imaging of increased blood flow to various parts of the brain. nonmusical functions. systematic study of human perception of the temporal characteristics of music using the most modern imaging techniques available – functional magnetic resonance imaging (fMRI) and positron emission tomography (PET) being favourites. melody and rhythm by neuroscientists in the USA. though much remains unknown in this area (despite brain localisation of musical functions being a popular research objective). These instruments have also been incorporated into popular music sung in hundreds of languages around the world. With careful experimental design it has been possible to localise many of the brain’s musical functions. Canada and Europe. The other ‘standard’ instruments of a rock/pop band (originating from a format shared between country. folk and blues.
at least in my case). rhythm and meter. More is known about our processing of pitch and melody than about rhythm. in ‘The Neural Correlates of Temporal Structures in Music’ that regions of the brain involved with temporal structure in music are bilateral. partly for the reasons mentioned. Daniel Levitin summarises. . activity extends to the ventral tegmental area.53 damage (from disease and surgery). whereas grouping would rely essentially on the left”. Temporal structure. and partly because of the deep location of structures that appear to be involved in our mental processing of time. In a 2005 review of the literature Robert Zatorre of McGill University and Isabelle Peretz of the University of Montreal explained that creating a rhythmic pattern appears to be processed more by the left side of the brain. are less accessible to testing by sticking electrodes onto the scalp or directly into the brain. which includes tempo. superior temporal poles and cerebellum. This theory is supported by the fact that it is easier to tap a steady beat with the left hand and a syncopated rhythm with the right than the other way around (I tried it myself. inferior frontal cortex. while beat perception causes more right-sided activity. the function ascribed more to the left hemisphere. The authors argue that this supports the hypothesis that the “right hemisphere handles meter. nucleus accumbens and hypothalamus. Grouping. These structures. When the music is emotionally meaningful. is “the segmentation of an ongoing sequence into temporal groups of events based on their durational value”. recruits networks in the prefrontal cortex. timing and rhythm. until recently. unlike those used in speech which are left-lateralised. been the mainstay of ‘neuropsychology’ research as far as brain localisation was concerned. including the cerebellum and basal ganglia. and found it to be true. Levitin reports. The argument is that extraction of the beat from a piece of music (meter) results in the “metrical organization corresponding to periodic alternation between strong and weak beats”. which has.
There is an accumulating body of evidence that the cerebellum is also involved in emotional reactions. The motor cortex. The modern fMRI and MEG scanning research also points to the close connection between emotions and movement. though the details of how this function is achieved are uncertain. If rhythmic music can modulate the activity of the hypothalamus this can surely be used to control blood pressure. and is thought to function as a ‘timekeeper’. Interesting research by Daniel Levitin and colleagues at McGill has shown that cerebellar activity (on fMRI scans) increases when people listen to pleasurable rhythmic music they have heard before. specifically a lobule on the medial side of the posterior parietal lobe (the precuneus) is involved in maintenance of a steady beat . The hypothalamus regulates the autonomic nervous system as well as the endocrine system. is only one component of the primate brain’s motor system.54 This research is interesting and provides a rationale for music therapy in a range of medical problems in which modulating the activity of the hypothalamus by judiciously using listening and sound is an obvious objective. since the negative effects of stress on the hypothalamus are well established. heal ulcers and remedy the host of medical problems known to be aggravated by stress. including the basal ganglia and cerebellum. The motor strip is intimately connected with the other major components. meaning voluntary and automated movements. even if this music is experienced as pleasurable. reduce heart disease. The fact remains that even .a patient with localised damage to the right precuneus lost his ability to clap in time to music. mentioned earlier. The cerebellum is known to be centrally involved in both correcting and learning movements. Emerging evidence also suggests that the parietal lobes. which play key roles in regulating movements and learning new motor skills. and especially pleasurable responses to rhythm. but not when they listen to novel music.
especially the caudate nuclei. and the structures have been extensively researched. when animals were thought not to have much in the way of emotions. and how similar they are to our own. a mental faculty with great relevance to music cognition. Other parts of the basal ganglia.55 such simple testing is rarely done in hospitals. and psychology experiments were obsessively focussed on measuring stimulus and response. there has been a relatively rapid recognition of the complex emotions and thinking of other mammals. The neurobiology of musical emotions Since the dark age of behaviourism. though. not just that of music therapists. in fact. The structures commonly known as the basal ‘ganglia’ are actually nuclei rather than ganglia (ganglia are defined as collections of nerve cells outside the central nervous system. This structure. have been implicated in organisational memory. but the basal ‘ganglia’ are located deep inside the brain). appears to be specifically involved with the experience of pleasure. Their role in controlling movement has been recognised for many decades. a rediscovery of observations made a hundred and fifty years ago by Charles Darwin. which rarely employ music therapists in acute neurology wards. Many of the neural structures implicated in evaluation and appreciation of rhythm have long been known to be involved in gait. Using music to evaluate the function of the brain should be part of general medical training. the nucleus accumbens. This is. though now we have a much clearer idea of the brain structures – the neural substrate . it is only in recent decades that a particular structure within the basal ganglia system has been identified as playing a key role in emotional reactions. balance and coordination. Such testing could be part of a resident medical officer or nurses evaluation skills. since dysfunction in the basal ganglia is known to cause the movement disorder known as Parkinson’s Disease. Though Parkinson’s Disease is known to be associated with depression.
emotions. affect and emotions. let alone conflicting opinions on the distinction between moods. fewer have looked at excitement and very few have studied what music makes people tap their hands and feet and why. In music cognition research. thalamus. the various emotions that music elicits in people across cultures and age-groups must be considered. birds and mammals. since there is neither popular nor scientific consensus on what emotions humans are capable of. but avoids the pitfalls of psychiatry. whose essential components are shared by reptiles. Some studies have looked at anger and fear (usually with classical music and soundtracks). The deep structures of the limbic system. most attention has gone into studying what areas of the brain are active when experimental subjects perceive passages of music as happy versus sad. where emotions tend to be classified in a negative way. the question of how the emotional aspects of human musicality evolved becomes a thorny one. once denigrated as the ‘primitive lizard brain’. or what parts of the brain (and body) are involved in feeling different emotions. Structures like the hippocampus. and the difficulty in measuring emotions. or pleasant versus unpleasant. Adding to this the confusing use of the term ‘affect’ by the medical profession to describe emotions. amygdala and nucleus accumbens have been studied in considerable detail for decades. This is a controversial area. is belatedly being recognised as being essential for the most ‘human’ of mental attributes . but only recently have we seen meaningful syntheses of the complex and sometimes contradictory findings of researchers around the world. or feelings. The classification of emotions used by music cognition researchers tends to be simple. cingulum. how to classify them.56 – of emotions. To understand the neurobiology of music. there has been a steady . Though the terms ‘limbic system’ and ‘limbic structures’ are widely used to refer to the neural circuit that underpins emotions. using various techniques. and the difficulty in defining different emotions.
the counter-intuitive theory by William James (1842-1910) that our emotions follow our physiological arousal.57 stream of evidence. were not included in Papez’s model. were key structures in the ‘Papez Circuit’. Attempts to study emotions scientifically have been hampered by the paradigm of ‘experimental physiology’ propounded by Harvard’s Walter Cannon during and after the First World War (1914-1918). for example. structures now recognised to play a vital role in emotions. Cannon was responsible for developing the theory of emotion commonly referred to as the Cannon-Bard theory. His errors were understandable – injecting rabies into the brains of cats is neither insightful nor the most accurate way of studying emotions scientifically. while the hippocampus and mammillary bodies. commonsense view differed from his own insights: “Our natural way of thinking about these standard emotions is that the mental perception of some fact excites the mental affection called the . as the dominant paradigm of American psychology. Cannon (18711945). The amygdala and nucleus accumbens. from the 1960s onwards. was credited by Papez (1883-1958). now known to be more important to memory than to emotional reactions. that the original ‘emotional circuit’ proposed by James Papez in 1937 and elaborated by Paul McLean in the 1950s (McLean popularised the term ‘limbic system’) is inaccurate and incomplete. This theory had replaced. a structure at the base of the brain on which the neuroanatomist Papez placed great emphasis within the ‘emotional circuit’ he proposed. who coined the phrase ‘fight or flight’ to describe the activity of the sympathetic nervous system. in the latter’s landmark 1937 paper (‘A Proposed Mechanism of Emotion’) as one of a handful of scientists who “greatly advanced knowledge of the functions of the hypothalamus”. In his 1884 paper titled ‘What is an Emotion?’ James had explained how the less enlightened.
but spent his whole career as an academic at Harvard since 1873. America’s oldest and most esteemed university. and that this latter state of mind gives rise to the bodily expression. it may turn out to be useful in presence of other features of the environment that had originally nothing to do with either producing it or preserving it”. as seems intuitively to be the case. we feel scared because we tremble. what really happens. rather than seeing a bear. William James was trained in medicine. feeling afraid and running away. ‘What is an Emotion?’ William James refers to Darwin’s work and “the well-known evolutionary principle that when a certain power has once been fixed in an animal by virtue of its utility in presence of certain features of the environment. In his 1884 paper. How William James’ theory was disproved by Cannon says a lot about the direction ‘experimental physiology’ went in once personal introspection as a means of understanding the mind was renounced in favour of ‘hard facts’. is we see the bear. run away and therefore feel afraid. My thesis on the contrary is that the bodily changes follow directly the PERCEPTION of the exciting fact. 1884. Initially employed as instructor in physiology and anatomy.” (James. . Likewise. and that our feeling of the same changes as they occur IS the emotion. his emphasis) James explained than in his opinion.58 emotion. disproved it that the ‘mainstream’ abandoned the ‘JamesLange theory’ for the ‘Cannon-Bard theory’. shortly after Darwin published The Descent of Man (1871) followed by The Expression of the Emotions in Man and Animals (1872). Though many others doubtless voiced doubts about the veracity of William James’ theory of emotions. Walter Cannon. he claimed. James regarded music as such a power. it was only when another Harvard professor. angry because we strike and sorry because we weep. he rose to the position of professor of psychology (then a fledgling science) and professor of philosophy at Harvard.
be seen to have had two phases. more often than not. which used the computer as a metaphor for mental functions. They argue. also. Emotions have been studied intensively in more recent years. Still today. Cannon proved him wrong by showing that dogs in which he had severed the nerves carrying visceral information to the brain still expressed emotions. memory. . that emotions are not confined to ‘subcortical’ brain structures. They tend to ignore emotions. pioneered by such neuroscientists as Antonio Damasio. 2010) The ‘cognitive revolution’ in psychology can. has only just begun. A few years ago. Firstly. the term ‘cognition’ was limited to perception. when the new discipline of cognitive neuroscience has increasingly included emotions as an essential part of ‘cognition’. the majority of experimental psychologists are ‘cognitivists’ by default. Joseph Le Doux. The second phase of the revolution. the brain is a machine. Emotion was specifically excluded from the theories and research of ‘cognitivists’ who were antagonistic to the views of ‘emotivists’. These scientists recognise that emotions are an integral part of decision-making and that reason is not a unique human faculty to control ‘irrational’ emotions. ‘thinking’ had to be re-included from a narrow focus on ‘behaviour’. It has a long history starting with Descartes early separation between emotion and reason. Daniel Levitin and Peretz herself. and other aspects of what was broadly called ‘thinking’ (or ‘intellect’). Isabelle Peretz explains: “The antagonism between ‘cognitivists’ and ‘emotivists’ is not limited to music psychology. (Peretz. then ‘emotions’ had to be included within the domain of ‘cognition’.59 James came to his contentious conclusions by analysing his own emotions. This neglect partly reflects the information-processing approach that started in the early sixties. then. According to this view. devoid of emotions”. as synonymous with ‘emotional centres’ in the brain. although the rather misleading terms ‘limbic system’ and ‘limbic structures’ continue to be used.
The neurobiology of music emotions. including music. Others have observed relative activity in parts of the brain when experimental subjects have listened to music they liked.60 Since auditory stimuli. compared with music they disliked. might be expected to provide a useful model for understanding emotional reactions to other auditory stimuli. These . having been studied more thoroughly than that of music. sadness and fear as ‘basic emotions’. As in other areas of experimental psychology. From what is known. derived from electrical stimulation of these structures. More recent imaging studies have confirmed the view. while the nucleus accumbens appears to be specifically associated with feelings of pleasure. though cortical areas specifically involved in speech comprehension and generation were identified a hundred and fifty years ago. has remained obscure. Other areas of the brain (including areas of cortex) have been associated with displeasure and disgust. can affect our emotions it is obvious that information reaching the PAC must be transmitted to parts of the brain involved in emotional reactions. Unfortunately. music neuroscientists have tended to concentrate on happiness. Such emotions are easily recognised cross-culturally and several studies have tested ability to differentiate ‘happy’ and ‘sad’ music in people with damaged and intact brains. these are mostly ‘subcortical’ structures – notably the amygdaloid complex (amygdala) and nucleus accumbens. though investigated with state-of-the-art imaging techniques. anger and sexual excitement. that the amygdalae are activated with fear. the common observation that understanding speech and expressing oneself verbally can induce pleasure has not yet been explained in terms of neuropsychology. What happens between the Primary Auditory Cortex and the brain structures involved in subjective ‘feelings’? And how adequate are existing models of the neurobiology of emotions to explain the agonies and ecstasies of music? The neural processing of speech.
at least in part. a world leader in music neurobiology at Canada’s University of Montreal concluded recently: “Although many questions about the neurobiological basis of musical emotions remain unsolved. most of the neural pathway underlying emotional responses to dissonance has been delineated and involves a complex and distributed system in the brain. For instance. Many happy tunes in non-European music are played in minor keys with mainly minor chords. anger and joy in Western and Hindustani music. Isabelle Peretz. 2010) In Towards a neurobiology of musical emotions Peretz discusses complex and sometimes contradictory findings about the cortical and sub-cortical processing of even the ‘basic emotions’. This emotional pathway is not simple.61 have generally confirmed the view that music emotions remain mysterious. while Japanese listeners can recognise sadness. there is evidence that musical emotions depend on a specialized emotional pathway that may recruit various subcortical and cortical structures that might be shared. fear and anger. . The other well known characteristic of happiness and sadness – major and minor chords and scales – appears to be largely due to acculturation. A cross-cultural recognition of basic emotions in music is suggested by the observation that Western listeners can easily recognise happiness. sadness and anger in Hindustani ragas. other researchers have suggested that the main variable tested in happiness/sadness evaluations of music is tempo. Her discussion is limited to these basic emotions – happiness. unitary emotional system underlying all emotional responses to music.” (Peretz. However. There is not a single. with other biologically important systems. The same piece played faster may make a sad piece sound happy. sadness.
since it is involved in processing of visual and auditory signals. Listening to music with the eyes closed is a different experience to listening with the eyes open. especially relating to their synchronisation and the way we turn our attention to visual or auditory signals – or both at the same time. with reciprocal connections between specific cortical layers and the thalamus. 20. The large nucleus adjacent to the MGN and LGN is the pulvinar. and listening with headphones and the eyes closed different again. It is necessary not just for music cognition but for any cognition and for consciousness – indeed for life. Medial Geniculate Nucleus (MGN) Fig. Destruction of the thalamus is fatal. The thalamus pulvinar Lateral Geniculate Nucleus (LGN) Auditory signals first enter the thalamus at the Medial Geniculate Nucleus (MGN) a lump of grey matter at the caudal end of the structure. which may have relevance to music perception and creation. This may reflect activity in the thalamus.62 Post-cortical processing of music in the thalamus The thalamus is one of the most complex and important structures in the brain. and the entire cortex is closely related to the structure. . Adjacent to the MGN is the lateral geniculate nucleus (LGN) which receives visual signals before projecting them to the visual cortex in the occipital lobes.
When listening to music or playing it with undivided attention one needs to ignore other distractions until the piece is finished. We are constantly subject to a hierarchy of distractions. especially somatic ones. recurring thoughts and ruminations. This provides a means of synchronising auditory signals internally and externally. mentioned earlier. The parietal lobes process information from the muscles. If one had to confirm each note by sight. not so. worries and the vagaries of our individual thought patterns. our concentration is usually divided between competing subjects of interest. there are also populations of neurons throughout the thalamus that project more diffusely to the outer layer of the brain (layer I). This information is essential for the development of skill on any instrument. The thalamus is also involved in succumbing to or resisting distractions. but we can return quickly to the subject of our attention. . An interesting recent finding is that though most of the thalamic neurons synapse in layer IV of the cortex in the same region as inputs received from layer V. In reality. ranging from bodily functions to intruding worries and concerns. This structure appears to be one component of an integrated network of cortical and subcortical structures involved in keeping one’s mind on the matter at hand.63 The pulvinar of the thalamus also has extensive reciprocal connections with the parietal lobes and provide a neural mechanism by which the mental metronome in the precuneus. such that the ‘imagined’ beat is synchronised with the external beat of the music. The thalamus appears to be involved in this splitting and unification of mental focus at different times and in different circumstances. The need to cough or sneeze is hard to resist. The connections of the pulvinar may also explain the ability to play instruments by touch. joints and skin which allow a person to play an instrument without looking at it. tendons. music-making would be a slow and laborious process. sends its signals to the rest of the brain.
however. maybe not. Much of the research on the thalamus has concerned its role in mediating pain. is an important function of music when used therapeutically. This is a network of nerve fibres that originates in the . Fortunately music rarely causes pain – not physical pain. In this function the thalamus is assisted by the reticular activating system (RAS). though. woodwork. an extraordinary slice of the Australian psyche is tuned in to commercial TV every evening. tidying. gardening. anyway. gaining the attention of people at the expense of the television distraction is. important. drawing. It may be that associations involving the thalamus are involved in a judgement of a piece of music being ‘painful’. television watching is an unhealthily sedentary pastime. jingles in TV ads and much that is chosen by radio stations. and in the important quest for using music therapeutically. The same time spent listening to music enables one to relax while doing housework. This includes music in shopping centres. If the statistics are to be believed. Distraction from the disease-creating audio-visual influence of television is an important first step. Though the thalamus is an interesting neural structure I will leave my discussion of it without exploring it much further at this stage. talking or dancing – and many other healthier ways of spending ones time than watching TV. There is always a large minority who spent much time and effort seeking out music and who choose to listen to music rather than watch TV in the evenings. In practice. physical and mental. Apart from implanting and reinforcing negative views of the world. I expect that most of the people affixed to television sets do not have the time and energy to read about my explorations of music and the brain.64 Nowadays. The role of the structure in enabling selective focus is less uncertain. Trying to wrest the thalami of such people from the cathode ray tube or plasma screen which is the sole focus of their vision and hearing for several hours a day is a task I will not even attempt. The thalamus is generally involved with how we direct our attention. most of the music people hear is probably not music they choose themselves. writing. Distracting people from pain. and this has relevance to music therapy in many ways. painting. others and ourselves.
including the thalamus. 21. Somatosensory cortex Fig. Primary auditory cortex (PAC) Medial geniculate nucleus (MGN) Lateral geniculate nucleus (LGN) precuneus Visual cortex . The thalamus and some of its cortical projections relevant to music perception.65 brain stem delivering the neurotransmitter noradrenalin to all parts of the brain.
Specifically. a nucleus known as the nucleus accumbens is now known to be activated with the positive emotion of pleasure. They are known to be the main site of pathology in Parkinson’s Disease and iatrogenic (treatment induced) Parkinsonism. slowness and difficulty with movement. Though the role of the basal ganglia in movement has been extensively studied for several decades. especially initiating movement and. emotional flatness. Fig 23 Basal ganglia showing location of nucleus accumbens and amygdala Caudate nucleus putamen Nucleus accumbens Amygdala . importantly. with symptoms and signs similar to Parkinson’s Disease – stiffness. The latter is a common problem with antipsychotic drugs. it is only recently that their function in emotional reactions has been focused on. This has obvious relevance to music.66 Post-cortical processing of music in the basal ganglia The basal ganglia are bilateral collections of nuclei deep within the cerebral hemispheres that are involved in motor function.
which some experts class as part of the basal ganglia and others as an extra. too. . that the insula is separated from the striatum by two sheets of white matter divided by a narrow plane of grey matter. deep. The inputs to the nucleus accumbens arrive from other parts of the striatum and from many parts of the cortex (including the insulae. including its possible role in music perception. promises to be an interesting area for future research. rather than specialised role in synchronising sensory data. All parts the cortex and not just the motor areas provide inputs to the basal ganglia. according to some authors. and is characterised by a relatively homogenous population of neurons. The claustrum appears to be involved in integrating various mental percepts into a single. which together form the ‘striatum’ which is characterised by large numbers of medium spiny neurons. Much of the function of this structure remains conjectural. structurally and functionally.67 The basal ganglia develop in the embryonic brain from the anterior-most part of the nervous system (telencephalon). It can be seen. This. of the striatum. This is the claustrum. with which the basal ganglia have a close relationship). along with the cerebral cortex to which it is extensively connected. Their lack of specialisation and extensive connections suggest. The claustrum is known to be involved in synchronising neural inputs. The nucleus accumbens forms part of the ventral striatum and acts as a central point. seventh layer of the insula. unitary percept. These synapse with the interestinglyshaped caudate nucleus and the putamen. a generalised. in the following diagram.
The neurons that produce dopamine (from the amino acid tyrosine) are pigmented with neuromelanin. a variant of the dark pigment in the skin. The selective stimulation of the ganglia is from dopaminergic neurons that originate in the ventral tegmental area and substantia nigra of the midbrain. Most of the cortical inputs to the striatum (the caudate nucleus and putamen) have GABAergic inhibitory synapses. . Location of the striatum. The main neurotransmitters known to be active in the basal ganglia are the inhibitory transmitter GABA and the stimulatory transmitter dopamine. having relevance to motivational states and learning as well as being involved in Parkinson’s Disease and drug-induced Parkinsonism.68 Striatum (caudate nucleus and putamen) Insula claustrum Fig 24. The neurotransmitters known to be active in these nuclei have been studied for several decades. claustrum and insula The biochemistry of the basal ganglia is interesting. hair and eyes.
is not amenable to treatment. Huntington’s Disease is characterised by early onset dementia and unusual writhing movements of the trunk and limbs (chorea). Tardive Dyskinesia (TD) is a crippling disease caused by long term use of antipsychotic drugs. notably Huntington’s Disease and Tardive Dyskinesia. 25 Damage to the basal ganglia is also associated with other diseases. pigmented neurons. Fig. trunk and face go involuntarily into grimaces. The condition. where the limbs. In both cases music is an obvious way of stimulating dopamine production. spasms. Though music is again a potential therapy for Huntington’s Disease and Tardive Dyskinesia.69 It is known that Parkinson’s Disease is characterised by selective degeneration of these dopamine-producing. In drug-induced Parkinsonism the symptoms are caused by blockade of dopamine receptors by the drugs. tics and contortions. musical strategies have inevitably been ignored before . which may be permanent even if the causative drugs are stopped.
Specifically.70 these terrible conditions have been declared untreatable and potentially permanent. including the motor areas of the cortex. the cerebellum and other parts of the basal ganglia. at the junction of the caudate nucleus and putamen is thought to be an important pleasure centre in the brain and part of the hedonic ‘motivational system’ that generates pleasure which acts as a motivator. Making pleasurable music activates even more of the brain. putamen Body of caudate nucleus Head of caudate nucleus Thalamus Fig 26. evidence is emerging that the basal ganglia are also deeply involved in learning. emotion and general motivational states. Since humans derive much pleasure from music it comes as no surprise that the nucleus accumbens has been shown to be activated by listening to pleasurable music. Location of basal ganglia relative to thalamus (left lateral view) Tail of caudate nucleus Nucleus Accumbens amygdala The main output of the basal ganglia goes to the thalamus and the subthalamic nuclei. In addition to their role in the fine control of movement and in preventing unwanted movements. These pathways form a modulatory cortico-striatal-thalamic-cortical loop for the . the nucleus accumbens. The latter project to the thalamus modulating the signals going from the thalamus back to the cortex and to other parts of the brain.
functionally (though not anatomically) parallel to the large tract of nerve fibres connecting the cortex with the cerebellum and the tracts that lead from the cortex to the spinal cord and skeletal musculature. There are therefore three parallel. integrated motor circuits that function synergistically when working well. All three movement systems can be modified by music. especially rhythmic music. . basal ganglia (red) and thalamus (yellow).71 motor system. Precortical auditory structures are shown in green. 27: Angled view of the left sided cerebellum. Cerebellum Fig.
including the amygdala and nucleus accumbens are structurally connected to the basal ganglia. the cingulate gyrus forming a border around the corpus callosum (the tract of white matter that connects the two hemispheres) when the brain is medially transected.72 Pre and post-cortical processing of music in the limbic system The term “limbic” is derived from the Latin limbus meaning “border”. The diagram below shows the position of the structures included in the limbic system in a 1990 neuroanatomy book Clinical Neuroanatomy by Professor Richard Snell (professor of anatomy at the school of medicine and health sciences at the George Washington University in Washington. since the 1960s there has been a steady stream of additions to the list of structures various authors have included in the ‘limbic system’. stria terminalis. This ring of tissue is involved in emotions. Broca had attributed emotions to this ring of grey matter and Papez thought he had confirmed it when he injected rabies into the brains of cats and examined them after the disease had progressed through the brain. and the structures they included in their emotional circuits included some that are more involved with memory and decision-making than emotions. USA). named the gyrus that includes the cingulate gyrus above the corpus callosum and the hippocampal formation below it. hippocampus. but so are many other parts of the brain. Papez and Broca were only partly right. the ‘great limbic lobe’. These include the amygdala. the . As it turns out. and Broca was very much into dissecting brains (he even weighed and dissected that of the great French naturalist Georges Cuvier (who had an exceptionally large brain to match his famously large head). back in 1878. This grey matter forms a ring around the medial side of a transected brain. and the model I have been developing explores what this could mean functionally. The famous French neurologist Paul Broca had. Some of these. fimbria. as well as the anterior nucleus of the thalamus. indusium griseum and fornix. Consequently.
included only the cingulate gyrus. hippocampus. mamillary bodies and anterior thalamic nuclei in his hypothesis.73 mammillary bodies. however. cingulate gyrus. . first coined by the neurologist Paul McLean in the 1950s. dentate gyrus and the parahippocampal gyrus. Papez. the anterior commissure. There is. who postulated an ‘emotional circuit’ in the human (and mammalian) brain following various experiments on animals. is of use at all. based on the neuroanatomist James Papez’s 1930s model. some dispute as to which structures should be included as components of the ‘limbic system’ or even whether the term.
Pituitary secretions are controlled by the hypothalamus.74 Figure 28: the limbic system In the diagram above the effect of the limbic structures on the secretions of the pituitary gland are illustrated. which is closely connected with both the amygdalae and the nuclei accumbens. since the hypothalamus. in addition to controlling the hormones secreted by the pituitary gland and pineal organ. These connections provide a neural mechanism by which emotions can affect diverse aspects of physiology. regulates the function of both the sympathetic and parasympathetic .
blood pressure. . thyroid stimulating hormone (TSH) and adrenocorticotropic hormone (ACTH) by the anterior lobe of the pituitary and oxytocin and vasopressin (antidiuretic hormone) by the posterior lobe.75 branches of the autonomic nervous system (which controls such things as heart rate. by a more complex model developed in 2000. centred on love (positive) versus fear (negative) that I was exploring in 1996. growth hormone (GH). contractility of the digestive tract and blood flow to the muscles and brain). A simple model of positive and negative emotions. vasoconstriction and vasodilation. prolactin (Prl). these physiological processes and connections provide a mechanism by which music can affect organs and tissues throughout the body. when I drew this diagram has been replaced. secretory activity of exocrine and endocrine glands. in my theory. luteinizing hormone (LH). This model of how emotions affect our physiology is shown in the next diagram (figure 29). The pituitary hormones under the control of the hypothalamus include follicle stimulating hormone (FSH). Since music is a powerful stimulator of emotions.
This model is expanded in the next diagram (figure 30).77 The diagram above (figure 29) presents a general model of the psychology of music and speech. . looking at some of the mental processes involved in listening to music.
Fig.78 which proposes a sequence of primary. secondary and tertiary emotional responses to music.31: Integrated model of musical emotions .
looking particularly at the role of the basal ganglia. This remains a work in progress.79 In the following diagram (figure 32) an attempt is made to integrate the structures discussed previously. These include. in addition to the auditory cortices. amygdalae and hypothalamus in perception of and response to music. the basal ganglia. . Figure 32: Integrated model of music neurobiology In the diagrams that follow (figures 33 and 34) some of the neural structures that underpin the dance instinct are explored. cerebellum and various limbic structures. Research over coming years will doubtless add further complexity to what is already a complex picture of how music and other auditory stimuli affect our physiology.
80 Obviously the urge to dance and physical dancing also includes the motor cortex. Figure 33: model of neural control of dance . located in the pre-central gyrus of the frontal lobes (the ‘motor strip’) and the pre-motor cortex anterior to this gyrus.
81 Figure 34: model of neural response to musical rhythms .
partly hidden under the occipital lobe. Research by Daniel Levitin and co-workers at McGill University in Canada has suggested that the cerebellum is also involved in emotional reactions to music (in addition to the known function of learning movements necessary to play musical instruments). like the cerebral hemispheres. though enthusiasm for singing out of tune often increases dramatically (this may have more to do with reducing inhibitions when the alcohol affects the frontal lobes). the cerebellum is dense with nerve cells on its outer areas. A wide-based gait is typical of people with damage to the cerebellum. These balance centres are obviously important for dancing (and drumming). with other congregations of cells in its core. such as playing instruments. though. These include musical movements. and the cerebellum is closely connected with the vestibular centres of the inner ear. resembling in appearance (though not function. Gait disturbance. has more nerve cells than the rest of the brain combined. In This is Your Brain on Music Levitin summaries the role of the cerebellum: . and lack of coordination and balance when the cerebellum is not functioning well point towards the important role of this part of the brain in control of movement. located at the rear of the brain. The cells and structure of the cerebellum are unlike those of the cerebral cortex. the lack of coordination that results from drunkenness is associated with cerebellar dysfunction. in most people) a ball of wool attached to the brainstem. With drunkenness both pitch and rhythm discrimination and performance become impaired. These include centres for balance. This ancient structure is finely folded.82 Pre and post-cortical processing of music in the cerebellum The cerebellum.
The various biological rhythms are synchronised to variable degrees. All these rhythms come under the influence of the brain. p. and an increased concentration of gray matter. Specifically. Interestingly the cerebellum was also activated when people listened to music they liked and had heard before. which may contribute to an explanation of the amazing abilities of Snowball the dancing cockatoo (his performances on You Tube are worth watching) and other dancing birds. such as are acquired by musicians. but not to music they disliked – or music they liked but were hearing for the first time. mainly via the hypothalamus. breathing and circadian rhythms. the important and complex structure at the base of the brain to which the pituitary is attached. A connection between a bipedal gait and music may also explain a natural preference for 2. including an increased number and density of synapses.” (Levitin.83 “Several studies have found microstructural changes in the cerebellum after the acquisition of motor skills.227) In addition to its role in motor function. Even the peristalsis of the bowels is rhythmic. Interestingly. Levitin found strong activations in the cerebellum when people listened to music but not to noise. 2006. parrots and other birds have well-developed cerebellums. There are other biological rhythms influenced by the brain. 3 and 4 and rhythms that can be divided into the beat of a bipedal rhythm. including heart beat and other cardiac rhythms. Activation of the cerebellum with rhythmic music supports the hypothesis that the dance instinct is connected with our bipedal gait. Schlaug [Harvard neuroscientist Gottfried Schlaug] found that musicians tended to have larger cerebellums than nonmusicians. Could music be used to synchronise all the biorhythms? . the cerebellum appeared to be involved in tracking the beat of music. It seems reasonable to infer that better health would be achieved if all these rhythms are well-synchronised.
Memory is fundamental to intelligence. From the very first note. especially various types of dementia. Decision-making processes centred on card games played on computers by psychology students seem to have had an inordinate amount of funding compared to that spent on musical memories. gambling games using cards. every piece of music produces expectations of the notes to follow. but there is general agreement that listening to music utilises every type of memory.84 Could music be specifically composed for such a purpose? Future research may cast light on these questions. many neurological problems. memory for playing cards. if duration is the aspect being classified. The neural substrate of musical memories Memory is one psychological attribute that has been studied in great detail. But duration is not the only way memory is classified – there are also different types of memory. but many are. and much is to be gained from improving memory. computer games and most importantly. long term memory and so-called sensory memory. Also. There are many ways that memories have been categorised and debate about how long each type of memory operates for. for obvious reasons. A therapist cannot control the mental associations that spring to the mind to someone who is listening to or . Musical memories are all-important in the evaluation and appreciation of music. This has meant video games. memory for faces and numbers and names. such as autobiographical memories. Psychologists and physiologists have studied memory in the usual ways but with all the new technology as it came on the market. are characterised by memory loss (partly because progressive memory loss is routinely diagnosed as dementia). and these are based on memories – short term memory. Not all types of memory are relevant to music and the brain.
This structure loops around in the wall of the third ventricle. and damage to the structure causes a specific inability to form long-term memories. including the hippocampus. Yet these memories are essential to the therapeutic potential of music when listened to or created. receive input from the hippocampus via tracts of white matter (masses of axons) that form the fornix. especially the anterior nucleus of the thalamus which receives input from the mammillary bodies via the mamillothalamic tract.85 making music. so named because they looked like breasts to the early dissectors of the primate brain. Figure 35: the hippocampal formation fornix mamillary bodies hippocampus The hippocampal formation is thought to be involved in long term and medium term memory. the lateral walls of which are the medial sides of the thalami. the mammillary bodies and the thalamus. deep in the temporal lobes. damage to the hippocampus. Several structures deep in the brain are known to be involved in memory. mammillary . Interestingly. The mamillary bodies.
This suggests that these long-term memories are not stored in the structures. meaning immediate recall. language and music) are widely distributed through the brain. but also other parts of the brain. This is a key function in the perception of music. which plays a central role in learning and remembering movements. The imagery we create when remembering involves the primary and secondary sensory areas. Rather. the hippocampal formation is clearly important in musical memories.86 bodies and anterior thalamic nuclei does not prevent access to long term memories established before the injury. and involve novel connections and uses of various brain structures. morality. . Penfield’s research. How the brain is wired. there is an emphasis on the synthetic aspect of memory – memory is a creative process in which the individual doing the remembering is the central player. as the phrenologists of the nineteenth century thought. faces and motor memory are not thought to involve the hippocampus. including the frontal lobes and insula. Nevertheless. and how components of the circuitry are activated and inhibited appears to be more responsible for unique human abilities than evolution in our brains of new ‘organs’ of language. vocabulary. aesthetics or music. where musical memories were evoked by electrodes stimulating the secondary auditory cortex in the temporal lobes led the famous neurosurgeon to believe that everything we have attended to is retained somewhere in the brain. This is not accepted these days. These are especially autobiographical memories – personal memories of the individual’s life. rather than the development of new structures. they involve areas of cortex as well as the cerebellum. Memory for facts. The frontal lobes are known to be centrally involved in ‘working memory’. rather wiring through the hippocampus and its connections is necessary for depositing the memory – and only specific types of memory. instead. Many mental functions (including memory.
Creatures such as ourselves that are social and have evolved a dependence on the security of a social settlement are likely to have evolved complex place maps including all our senses. Studies on monkeys and university students have suggested that such place cells also exist in primate brains. is likely to involve the hippocampus and its extensive connections with the temporal. Though muffled by amniotic fluid. while that of an adult is 60-90 beats per minute at rest.would be expected in any animal that has a homing instinct. The perception of music and the mental imagery of sounds involves activation of the primary auditory cortex. The auditory place map begins forming with a background of the mother’s heartbeat and the pulse of an unborn babies own heart. An auditory place map. by analogy . The development of this auditory place map and its relationship with other mental place maps (including maps of locations. which is known to be active during auditory hallucinations too. Certain neurons (place cells) in the hippocampus fire only when the experimental rat is in a certain location. A neonatal heart rate in the third trimester is about 140-160 beats per minute. possibly including a magnetic sensitivity as well (including sense of direction by orienting according to the earth’s geomagnetic fields). faces and personalities) may be considered a factor in the development of aesthetic responses to music and the complexities of musical memory. where they are activated by looking at familiar locations as well as being in them. the developing baby can hear the mother’s voice and those of the company the . parietal and occipital lobes. one component of the total place map in every mind. Such place maps – internal GPS systems. Studies on rats and monkeys have suggested that the hippocampus is a key structure in the creation of place maps. regardless of whether this is a maze or an open living environment.87 where the notes that have just been played dictate expectations of the notes that will follow.
safety and a sense of belonging. Basal ganglia thalamus Figure 36: coronal section of the brain through the hippocampus. After birth the infant is exposed to a multitude of unfamiliar sounds. some of which are alarming. insula and basal ganglia.88 pregnant mother keeps. Your place maps belong to you but you belong to your place maps. As in all alarming situations the helpless infant is responding in such as way as to get protection from the mother – the most important feature in his or her auditory place map. despite them being figments of your imagination. These induce a ‘startle reflex’ where the arms jerk open and the eyes widen in response to sudden sounds (I was trained to ‘elicit the startle reflex’ when I worked as a paediatric registrar in Brisbane’s Royal Children’s Hospital in the 1980s). Of course. People love their place maps – the place maps are deeply connected with concepts of home. too. The sounds that the mother listens to (except via head phones) the unborn child listens to as well. These are the sounds that our auditory place maps are connected with temporally. this auditory place map is superimposed on a visual place map and deeply entwined with emotional reactions. insula parahippocampal gyrus hippocampus . thalamus.
Most of the inputs to the structure come via the adjacent parahippocampal gyrus. rather than in the ‘limbus’ – the ring of cortex around the transected brain. especially those . runs the tail of the caudate nucleus of the basal ganglia. and several nuclei have been identified within it (hence the term amygdaloid complex). It can be seen from the cross-section above that the hippocampus is a coiled layer of cortex that folds under the temporal lobes. The structure of the amygdala is complex. Interest in the amygdala increased when the structure was found to be activated with sexual excitement as well. The end of the caudate nucleus is expanded to form the amygdaloid complex (amygdala) which has been the focus of much attention since the 1960s when it was shown to be of central importance to emotional-physiological reactions of fear. The parahippocampal gyrus is also closely associated with areas involved in smell. which is filled with cerebrospinal fluid. though Papez and McLean did not include it in the emotional circuits they proposed in the 1930s and 1950s.89 The hippocampus is an archaic part of the brain and. Also. unlike the neocortex. terror and rage. including the visual cortex in the occipital lobes and the auditory cortex in the temporal lobes. The amygdaloid complex also provides a mechanism by which our autobiographical memories and place maps can have a profound effect on our emotions. Above the ventricle. The hippocampus runs along the floor of the lateral ventricle along the length of the inferior horn of the ventricle. explaining why odours have such a strong effect on memory. This area of cortex on the underside of the temporal lobes is extensively connected with the secondary sensory areas. Most of these functions relate to emotional reactions and the amygdala provides a mechanism by which particular emotional reactions cause immediate and profound physiological changes through the close connection between the amygdala and the hypothalamus (which is also complex and composed of several nuclei with different. is composed of only three layers. attached like an Ankylosaurus’ club to the tail of the caudate nucleus. the amygdala is located deep in the anterior end of the temporal lobes. functions). though integrated. The amygdala is regarded as an essential part of the limbic system.
90 of fear. Journal of Psychiatry and Clinical Neuroscience. References: Bamiou. and a dramatic increase in the amount of research being done in the area. T. I have indicated some of the areas in which future research can fill in the many holes in our current knowledge about music and the brain – I am sure that that there are many others. This is an exciting time to be researching music and the brain. This is despite music being a ubiquitous feature of human society and culture. pp 143-154 Bever. Most neurosciences books contain few if any references to music. These models are still works-in-progress. (1974) Cerebral Dominance in Musicians and Nonmusicians. and the neural structures that underpin these emotions. Vol 21. It seems reasonable to conclude that the amygdala plays a key role in our enjoyment of exciting music. Musiek. In this paper I have explored some of the research that has contributed to a greater understanding of how music affects the brain and tried to bring some of the data together to formulate integrated models of how music affects the brain. and Chiarello. anger and excitement. Ignoring the neural processing of music means that the enormous potential of music as a healing agent is also neglected by the medical profession and the health sciences. Conclusion Scientific study of the neural processing of music is still in its infancy. R. pp 94-97 . The topic is truly at the cutting edge of medical science. much is still uncertain about how music affects emotions. Vol 42. In particular. Luxon (2003) The Insula and its Role in Auditory Processing – literature review. Brain Research Reviews.
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