La Théorie Sensorielle

By Philippe Roi⁽¹⁾, Tristan Girard⁽²⁾, Richard Dumbrill⁽³⁾, Michel Leibovici⁽⁴⁾
With the participation of Paul Avan⁽⁵⁾

⁽¹⁾Researcher in Cognitive Science, specializing in Cognitive Archaeology ; ⁽²⁾Researcher in Cognitive Science ; ⁽³⁾Professor of Archaeomusicology, Director of ICONEA, Institute of Musical Research, School of Advanced Study, Universities of Oxford and London. ⁽⁴⁾Doctor in Cellular and Molecular Biology, Paris VI University, Researcher at the CNRS, Cochin Institute, INSERM U1016. ⁽⁵⁾Director of the Laboratory of Neurosensorial Biophysic, Clermont-Ferrand.

Abstract: During the fourth millennium BCE, in southern Mesopotamia, the Urukeans invented seven remarkable tools –the ard, the normalised brick mould, writing, accounting, the harp, the vertical weaving loom and the cone image– which the foundations of our civilisation still rely upon today. These inventions, among which was the primitive harp, have been found to mirror biological mechanisms which enable our sensory organs to perceive the world in which we live, and to codify it in order to transmit its representation to the brain. With regard to the primitive harp, its inspiration came from the organ of Corti, the sensory-nervous structure of the cochlea. A question remains as to how man could have created such an instrument at a time when the anatomy and physiology of the inner ear was impossible to perceive and comprehend. In order to answer this question, Philippe Roi and Tristan Girard have combined knowledge that was fragmented and separated into various fields, such as archaeomusicology, cell biology and neuroscience. This is how they discovered that there was a logical link between the Urukean harp and the organ of Corti.

Southern Mesopotamian elites would have increased their exchanges with those in northern Mesopotamia, in order to acquire more goods which distinguished them from the lower classes. As a consequence, there would have been an escalation of conflicts, as has been hypothesized after recent discoveries of mass burials in Syria at Majnouna, Hamoukar and Tell Brak, the latter being a significant strategic crossways (Fig. 1). In these burials, marks on the skeletons of hundreds, even thousands of young adults suggest a violent death.¹ Deep test-pits in the main sites revealed evidence of northern Levantian and Anatolian raw materials which were converted locally into manufactured goods.² Silt-rich southern Mesopotamian lands were ideally suited to agriculture and cattle husbandry. However, they did not yield any semi-precious stones, metals, wood, flint, basalt or obsidian, nor foodstuffs such as olive oil, wine or honey. The land did not produce any goods which could satisfy their rulers’ ostentatious frenzy, rulers whose power, conditioned by affectation, distinguished them from their peers and neighbouring cities. Cereals, and probably exceptionally fine long-fleeced ovines, were the only bartering goods traded with northern Mesopotamian villages.

Owing to the ard, northern Urukean breeders fed their herds with generous amounts of the wheat and barley they produced. When lambs are fed with high salt content cereals and hay, they develop the finest wool, which has a diameter of less than 11.8 microns.³ North and south would have bartered exotic goods for cereals and exceptionally long-fleeced ovines (Fig. 2). These exchanges would have led to lengthy arguments carried out in various dialects. At best, the deals ended in banquets and at worst, in blood feuds. History abounds with trade arguments leading to wars. A bloody massacre such as that at Tell Brak would have led to the antagonists finding ways to appease tension during negotiations. Urukeans developed one of the most subtle and ancient arts as a possible solution. This is substantiated by the emergence of harps fitted with three or four strings, as depicted on a fourth millennium seal impression from Choga Mish, east of Uruk in Elam, in modern south-west Iran (Fig. 3). The seal depicts a four-string arched harp played by a seated person, while two others beat a drum, a bowl-drum and clappers.⁴ As early as the fourth millennium, Urukeans would have discovered that harp music could unite people, allowing them to share feelings and emotions that they could otherwise only have experienced individually. This was the primary function of the harp: the music it produced would have facilitated the exchange of goods. From the beginning it promoted equal understanding; in other words, ‘all were in tune’. From then on, owing to the fact that the harp was a remarkable mediator, music would have been included in communication procedures between different communities, changing people’s mindset and developing empathy between them. Being the first to understand that music could have a significantly smoothing effect on those that listened to it, Urukeans found that it could also affect protagonists during negotiations. If music cannot change the product itself, it can unconsciously act favourably on the customer’s mood. The smoothing effect of the harp on bad tempers, and its contribution to emotional reactions, would have meant that the instrument had high status, from dawn to dusk. The harp was found in potters’ and weavers’ workshops, on goods barges, in banquets (Fig. 6) and even during sexual intercourse (Fig. 4-B). The Urukeans’ pride in their invention was so great that they engraved it on cylinder seals; in its silent manifestation, it sounded to the inner ears of those looking at its impression on seals and tablets. On tablets it was denoted by the Sumerian words BAN.TUR, BAN meaning ‘bow’ and TUR meaning ‘small’, hence harp, and the Sumerogram BALAG, voiced as the onomatopoeic ‘dubdub’, a word echoing the sound of the object it depicts. Around 3,300 to 3,000 BCE, the pictogram with which it is associated clearly depicted a harp with three or four strings (Fig. 4). However, 500 years later the word was used to denote a drum.⁵

Harps from Uruk and Djemdet Nasr were generally monoxyle or monostructural, meaning that there was no distinction between the soundbox and the part which would become the yoke, or the neck. They would have been made from gourds or calabashes, the natural shape of which were appropriate for this. It is possible that they were domesticated through simple cultivation techniques which made them grow in the shape of musical instruments. ⁶The dried fruit was hollowed out from an oval opening, which was then covered with a soundboard made from damp sheep, pig or calf raw hide. It was stretched at the back with wet hide strands. The strings were made from fresh twisted gut or vegetal fibres. A sliver of wood was tied at the end of each string, to prevent it slipping out of the soundboard during tuning. The upper ends of the strings were tied to a strip of woven material rolled around the neck, to ensure tuning by friction (Fig. 5-left). At that time, and especially towards the end of the fourth millennium, the size of soundboxes progressively increased while necks became thinner. Gourds and calabashes would still have been used for soundboxes, but necks would now be made of wood into which the Urukeans would later have plugged tuning pegs to ensure the tension of the strings (Fig. 5-right). Fourth millennium harps would have been small, with probably no more than three strings, stretched over a plan of around 110 degrees determined by the angle of the soundbox in relation to the neck. This suggests an anhemitonic disposition with a span of no more than a musical fifth, possibly including a third. These harps were always depicted in rural scenes, surrounded by animals, but with no reference to religious rituals; practical usage was thus implied, as is clearly shown on cylinder seals. However, from the third millennium onwards, harps were always shown in scenes depicting Inanna, the guardian Goddess of Uruk; they even symbolised her. Some texts record Inanna’s animals and her attributes, which included the reed, the palm, the aster Venus, and the harp itself.

Having described the harp in its original context, it is very clear that its design rests on the fundamental principle of hearing. Mankind’s perception of sound is an outstanding phenomenon. Sounds are simple periodical variations in air pressure which travel as a wavefront, at a speed of three hundred metres per second. When the waves reach our ear they are channelled into its canal and reach a thin membrane, the eardrum, which separates the middle ear from the outer ear. The vibrations of the eardrum, which result from variations in acoustic pressure, are transmitted to a chain of four small bones located in the stony part of the temporal bone: the hammer, the anvil, the lenticular bone and the stirrup. These ossicles articulate with each other. They are connected by ligaments, and transmit vibrations from the air environment of the middle ear to the aqueous medium of the inner ear, without any loss of energy. The inner ear is a complex structure in the temporal bones, consisting of a labyrinth and several liquid-filled cavities. This system is made up of canals, cavities and a spiralling structure called the cochlea. It is home to two very distinct sensory organs: the vestibular system, which detects and adapts to spatial body position, and the cochlea, which is the auditory receptor organ.

The cochlea, from the Latin meaning ‘snail’, is a small spiralling structure measuring 1.2 mm in diameter by 35 mm in length. It begins at the anterior part of the vestibule and rotates two and three quarter times around a bony pillar, the modiolus. At its lower end are the oval and round windows, which separate the middle ear from the inner ear. The cochlea is divided lengthwise into three chambers. On each side are the scala tympani and the vestibuli, both of which are filled with a liquid called perilymph. A cochlear duct is located between the two scalae; its lower part ends in the basiliar membrane, its upper part in Reissner’s membrane. The cochlear canal contains the organ of Corti, which is a complex system lying on the basilar membrane and extending right along the cochlea. This organ transforms vibrational energy into electric signals which the brain can interpret (Fig. 8). There are up to 15,000 outer hair cells spread over three or four rows, and 3,500 inner hair cells in one row (Fig. 9). The cellular bodies of the hair cells float in perilymph liquid, while hair bundles stand in the endolymph-filled cochlear canal. The ionic sealing of the cochlear canal is ensured both at its base by the reticular lamina, resulting from the tightly joined apical surfaces of hair cells and the supporting cells of the organ of Corti, and at its apex by Reissner’s membrane, which forms the ‘roof’ of the cochlear canal. When the stirrup moves under eardrum vibration pressure, it initiates motion of the incompressible scala vestibuli liquid. This motion reaches the top of the cochlea, reverses at the helicotrema, and finally runs down the scala tympani where it activates the round window placed at the other end of the system. The liquid movements initiate undulation of the basilar membrane, each part of which vibrates to a given frequency (Fig. 8-D). In turn these vibrations initiate displacements of the outer hair cells, which are rooted between the basilar membrane (by their basal pole, via Deiters cells), and the tectorial membrane (via their hair bundle) (Fig. 10). A relative alternating movement of the outer hair cells facing the tectorial membrane follows, provoking stereocilia deflection. Thus, the mechanical opening of the ionic channels results from a cellular depolarisation which leads to a rapid contraction of the outer hair cells, or electromotility, which then increases the amplitude of the basilar membrane vibrations. Consequently, this non-linear amplification phenomenon increases the weaker stimuli, which might not otherwise be perceived without interfering with high-intensity stimuli which would damage the inner hair cells, as a result of amplification. Because of this amplification, the stereocilia of the inner hair cells – which are the genuine sensory receptors of the auditory organ – are dragged in by the tectorial membrane and are also deflected. It follows that inner hair cell depolarisation releases neurotransmitter glutamate at its basal pole. This generates a spike train, sent to the brain by the auditory nerve.

An initial comparison can now be made between the neck of the harp and the basilar membrane (BM). The tuning pegs, around which the strings are wound and then driven into the neck, are comparable to the basilar membrane where Deiters cells are attached, and to which one of the ends of the outer hair cells (OHCs) is affixed. The neck is an essential part, because all the components of a harp rest on it, directly or indirectly. The functional importance of the basilar membrane is equally critical, because the various elements constituting the organ of Corti, again directly or indirectly, are attached to it. The harpist plucks the strings with greater or lesser intensity, his fingers complementing the work of the neck which the musician holds against his chest with his palm. Similarly, the sound vibrations transmitted by the perilymph generate upward and downward movements of the basilar membrane, at a precise location, resulting in the outer hair cells vibrating more or less intensely. As with the neck, the basilar membrane remains motionless right along the organ of Corti (Fig. 11-left).

A second comparison can be made between the tuning pegs inserted into the thick neck and Deiters cells (DCs), where the base is solidly affixed to the basilar membrane. The function of the tuning pegs is to secure the tension of the strings between the neck and the soundboard. Likewise, Deiters cells firmly grip the outer hair cells (OHCs) stretched between the basilar membrane (BM) and the tectorial membrane (TM). As for the clockwise or anti-clockwise rotation of the tuning pegs to tighten or relax the strings for tuning, the deformation of Deiters cells can modulate vibrations of the OHCs to which they are linked, as has recently been shown.⁷ They modify their membrane tension. This localised modification contributes to the refinement of the acoustic stimuli response (Fig. 11-right).

A third comparison may be drawn between the harp strings and the outer hair cells of the cochlea. One end of each string is wound around a tuning peg; the other is tied under the soundboard. This two-point attachment gives sufficient tension for their vibration, when plucked. This generates radial movement of the string as well as a light oscillation, as it is made of twisted material. A longitudinal wave propogation follows, starting from peg to soundboard. Similarly, the base of each outer hair cell is inserted in a Deiters cell, and its hair bundle is attached to the tectorial membrane. Thus secured, the OHCs generate rapid tension/relaxation cellular body motions – cellular electromotility – coupled with periodical oscillations of the hair bundle. Linked to electromotility, oscillations from the base of the OHC have been shown as leading to a radial displacement of the cell. It is important to add that the harp strings are only activated by plucking. Indeed, their arrangement does not allow for shortening of their speaking lengths, as is the case with the violin, for example. Each free string produces only one sound. In a similar way each transverse row of outer hair cells, in respect of its position along the organ of Corti and its structural characteristics, only responds to the stimulation of a single frequency (Fig. 12). Finally, in a manner similar to the work of the harpist’s fingers, which can dampen the vibrations of certain strings and allow others to sustain their sound – sometimes even sympathetically – numerous efferent fibres are projected from the central nervous system of the OHCs via the cholinergic synapses, in order to modulate their response intensity.⁸

A fourth comparison can be established between the soundboard and the tectorial membrane. The soundboard is a membrane made of fibrous/gelatinous animal hide stretched over the soundbox. One end of each string is driven through the soundboard, as can be seen from the holes when the strings are taken off. As the surface of the strings is very limited, the sound produced will have little intensity. However, if the string vibrations are transmitted to a larger surface, with better conduction, the sounds produced will be shorter but more intense. Not only does the soundboard amplify the signal originating from the vibration of the strings, it also transmits it to the soundbox. Similarly the tectorial membrane is also fibrous/gelatinous, and coats the organ of Corti. The ends of the outer hair cells’ longest ciliae are set in this membrane. Their surface shows hollow marks which are clearly visible with an electron microscope. When the OHCs vibrate, their hair bundles bend and amplify the relative motion between the basilar and the tectorial membranes. The signal which is thus amplified is transmitted to the inner hair cells. Therefore the tectorial membrane is paramount to the transmission of the sound stimulus, as is the soundboard (Fig. 13).

A fifth comparison can be made between the anchoring slivers and the tectorial membrane ties. String ties are thin slivers of wood placed horizontally under the soundboard. They contribute to the transmission of vibrations and prevent loosening of the knots at the lower extremities of the strings, without which they would slip under tension. Similarly, there are attachment ties under the thick tectorial membrane layer. Ties are needed to anchor the tip of the upper row of stereocilia of the OHCs’ hair bundles to the tectorial membrane (Fig. 14). This connection increases the vibration efficiency of the OHCs and, to a lesser degree, protects them from excessive tension.⁹

Finally, a sixth comparison can be observed between the soundbox (and its sound hole(s)) and the inner hair cell (and its afferent fibres). In organological terms, the soundbox is the part of a harp, the function of which is to gather and render audible the vibrations of strings and soundboard. When sound waves enter the soundbox, they reach its walls. The walls absorb some of the waves, while others bounce off them. The waves which are reflected converge and add to each other. This results in more perceptible sounds, most of which escape from the openings, or sound holes, placed at the base or sides of the soundbox. A third acoustic dimension adds to the two aforementioned ones generated by the strings and transmitted by the soundboard. This third acoustical element, generated by the soundbox, comes from the shape of the acoustical volume. It is the structure of the soundbox which shapes the sound. Additionally, however, the timbre is enriched by soundboard bars glued under it: they guide waves and enrich them euphonically (Fig. 15). Thus, depending on the complexity and quality of the bars, the timbre will result in more or less complex forms. This explains why the timbre and volume of instruments depend on their morphology. The inner hair cell (IHC) is the real sensory cell of the organ of Corti, since it perceives external stimulation and sends it to the brain. The IHC is mechanically stimulated by vibrations transmitted through the tectorial membrane deflecting the apical hair bundle. This deflection triggers a calcium influx at the base of the cell, which in turn generates a correlated neurotransmitter stimulation. This results in action potentials or spikes within the afferent nervous fibres. These are channelled via the auditory nerve to the superior nervous centres. It has therefore been shown that the pre- and postsynaptic components have a morpho-functional distribution which allows the IHC to shape its electric response.¹⁰ Thus, in a comparable manner, the soundbox and its sound holes, and the IHC and its afferent nervous fibres, both convert a simple entry signal (oscillation of the hair bundle) into a complex output signal.

NOTES
¹ McMahon, A. (2007) Soltysiak, A. (2007).
² Oates, J. (2007).
³ Lupton, C.J. et al. (2007).
⁴ Delougaz, P.P.; Kantor, H.J. (1972).
⁵ Dumbrill, R. (1998). Période au cours de laquelle les cités-États sont en guerre.
⁶ Dumbrill, R. (2012).
⁷ Yu, N.; Zhao, H.-B. (2009).
⁸ Frolenkov, G.I. (2006) Maison, S.F. et al. (2007) Richard, C. (2010).
⁹ Richardson, G.P. et al. (2008) Verpy, E.; Leibovici, M. et al. (2011).
¹⁰ Safieddine, S.; El-Amraoui, A.; Petit, C. (2012).

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In book (for the original French version), La Théorie Sensorielle. I – Les Analogies Sensorielles (2013) Roi, P. & Girard, T. Published by First Edition Design Publishing. Library of Congress Cataloging in Publication Data. Dépôt légal à la Bibliothèque royale de Belgique.

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