Cochlear fluids

KUNNAMPALLIL GEJO JOHN, BASLP,MASLP AUDIOLOGIST

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To be Discussed

Origin Composition Absorption Dynamics Functions

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Inner ear
1.

Bony labyrinth: Intricate series of interconnecting fluid filled tubes.

2. Membranous labyrinth: series of membranous structures suspended within the chambers of bony labyrinth.

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The fluid within the membranous labyrinth is termed as Endolymph. The fluid that surrounds the membranous structures is termed as Perilymph.

The endolymph and perilymph Differ in composition Play a vital role in physiology of the inner ear.
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Cochlea

Scala Tympani (ST) Scala Media (SM) Scala Vestibuli (SV)

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Points to be noted
1.ST and SV both contain perilymph and they have openings to the middle ear cavity which are closed by the round window membrane and the footplate of the stapes. 2. In the apical turn, the SV and ST are joined through an opening called the Helicotrema.

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3. In the basal turn, the SV has a wide connection with the perilymphatic space of the vestibule. 4. The perilymph of ST is connected to the CSF of the sub arachnoid space by the cochlear aqueduct. 5. The SM containing endolymph, is present between the perilymph scalae.

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6. The cells surrounding the endolymphatic compartment constitute an endolymphperilymph barrier. 7. In the basal turn,SM is joined by a narrow duct, the ductus reuniens to the endolymphatic compartment in the saccule.

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Scala Media
Three major structures bounding endolymph: 1.Reissners membrane - avascular structure - composed of two cell layers, forms boundary between SM and SV. 2.Stria vascularis - highly vascular structure, multilayered tissue - forms the lateral wall of scala media
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3. Organ of Corti - complex structure contains sensory hair

cells, supporting cells and basilar membrane. - the hair cells with their apical surfaces are in contact with endolymph and their baso lateral membrane are in contact with fluid of perilymph like composition. - the sensitivity of transduction process depends on the maintenance of this condition.
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Composition
Fluid sampling and analysis techniques: Glass micropipette (suction) method Techniques for estimation of ionic concentration Micro flame photometers or helium glow photometers Electrometric titration technique - for chloride concentration

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Ion selective electrodes by choice of ion exchange resin, the electrode can be made sensitive to K+, Na+, Cl-, H+ or Ca2+. X- ray microanalysis in the scanning electron microscope- estimates of elemental composition of cochlear fluids. Laser microprobe mass spectrography- to compare intracellular ion concentrations in different cell types.

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Perilymph
The ionic composition of the perilymph is similar to that of other extra cellular fluids like CSF. It has a high Na+ content and low K+ content (Na+ is the predominant cation) The Na+ content of perilymph of SV(mean 140.6 mM) is slightly lower than that of ST( mean 147.3 mM). The K+ content of perilymph in SV (6.7mM) is higher than that of ST (mean 3.4 mM). Therefore the perilymph composition is not homogenous throughout the cochlea. The osmolarity is similar to that of blood plasma i.e perilymph normally close to osmotic equilibrium with blood.

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ST PL SV PL

EL

CSF BICARBONATE

CHLORIDE

POTASSIUM

SODIUM

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Endolymph
It has unique ionic composition. Has high K+ and low Na+ content ( predominant cation is K+) Na+ concentration in the endolymph is between 0.5 and 2.0 mM. K+ concentration is between 150-165mM. Cl- concentration is in the region of 125140mM.

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It is positively polarized by approximately +80mV,the Endocochlear Potential(EP)

(Smith, Lowry& Wu 67)

The EP and the endolymph K+ concentration are lower in higher (apical) turns of the cochlea than in the basal turn.

(Sterkers et al 69)

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Cortilymph (Perilymph like fluid)

It is found in tunnel of corti (organ of corti). Is rich in Na+ and poor in K+ but its composition is different from that of perilymph Potential of cortilymph is 0mV Composition is similar to the composition of perilymph in ST.

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ST Pl.

SV Pl.

CSF

Coc.El. Sac. El. En. Sac El.

Na+
K+ Ca2+ ClpH Elec. pot

149
3.7 0.7 127 7.28 0

140
8 0.6 125 7.26 5

146
3.2 1.2 131 7.28 0
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1
158 0.02 136 7.37 85

3
150 0.09 119 5

108
14 0.47 98 13

Importance of composition

Provides the charge carrier and the ionic milieu for the process of transduction. Acts as a reservoir of metabolic substrates for the surrounding tissues.

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ORIGIN AND ABSORPTION

Three main principles

Longitudinal flow: The fluids are secreted at a site spatially separate from the site of resorption creating a volume flow along the cochlea.
Radial flow: Fluids are secreted and absorbed in all turns of the cochlea resulting in a radial flow. (secreted by RM and absorbed by stria vascularis)
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Homeostasis: It is the physiological process in an organism that maintain relative stability of its internal environment. Here its the composition of the fluids with out any volume production or volume flow taking place. That means there is no production or flow, it is homeostatic state.

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Perilymph
Hypothesis 1 1.The perilymph is derived from CSF by a longitudinal flow through the cochlear aqueduct. This was speculated due to To the existence of the cochlear aqueduct The similarity of the ionic composition of perilymph and CSF.

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Kaupp and Giebel : Perilymph marked with

fluorescent rhodamine rapidly appears in the subarachnoid space i.e chemical entered CSF after it was introduced to perilymph.

Calborg and Farmer: Mixing of CSF and perilymph

as a result of small cyclical volume movements accompanying pressure change during respiration. Under normal physiologic conditions, there was no significant pressure difference between perilymph and CSF.
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Hypothesis 2: 1. Perilymph origins in the cochlea by an ultra filtration mechanism. 2. Hawkins described capillaries in the SV of the spiral ligament close to the attachment of RM, which is believed could be site of ultra filtration. It could also involve longitudinal perilymph flow. 3. If perilymph was resorbed in the lower spiral ligament of ST near the basilar membrane, then a volume flow from SV to ST through the helicotrema could be expected.
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Hypothesis 3: Perilymph appears to be maintained by local mechanisms that do not necessarily involve perilymph secretion at all. John et al gave alternate mechanisms: a) Active diffusion b) Passive diffusion b) Facilitated transport d) Exchange across a blood labyrinth barrier comprised of the pericytes, fibrocytes and endothelial cells associated with the capillaries of the spiral ligament. Through this barrier Na+, Cl- and Ca2+ enter into the perilymph.
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Endolymph
Hypothesis I:
1.Stria vascularis is responsible for the active transport mechanism, which helps in maintaining EP and high endolymphatic K+ concentration. It is the source of EP. 2. Sellick et al :Showed that EP originates, active secretions of K+ into the endolymph. Speculated EP could be generated by conventional Na+/K+ ATPase. 3.If cochlea becomes anoxic or is treated with K+ transport inhibitors, the EP falls from its normal value of approximate +85mv, endolymph K+ begins to fall and Na begins to rise. endolymph equilibrates passively with perilymphatic compartments.
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Hypothesis II: The existence of endolymph flow was proposed by

Guild.
1.The existence of the ductus reunions connecting the cochlear and saccular endolymph, suggests the potential for longitudinal flow between these compartments. 2.He suggested endolymph was secreted in cochlea,longitudinally flowed through the ductus reunions and the saccule and absorbed in the endolymphatic sac. 3.Variety of traces substance that was injected in the cochlea, chemical reached endolymphatic sac.
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Hypothesis III: Endolymphatic homeostasis may occur.If water equilibrates passively, then endolymph composition would be maintained without volume flow necessary taking place. This process is called local homeostasis. Hypothesis IV: The difference in osmolarity between endolymph and perilymph leads to bulk flow. Suggests that the resulting influx of water into endolymph could drive its movements towards sac.
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Hypothesis V:

Naftalin & Harison (54) : The endolymph flowed

radially, secreted by Reissners membrane and resorbed by stria vascularis.

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Barrier between fluids

1. Anatomical barrier between endolymph and perilymph consists of the epithelial lining the SM, the saccule, the utricle, the three semi circular canals, endolymphatic duct and endolymphatic sac. 2. Functional barrier for ions and organic compounds but not for water. It is a difference in ionic composition that requires active transport for maintenance. Glucose, proteins and most amino acids in endolymph in much lower concentration than in perilymph.
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It is compatible with selective transport processes operating between these two fluids.

traces between blood, CSF, perilymph and endolymph support this concept.

Sterkers et al (82): The kinetics of distribution of

Thalmann,salt & de mott(88): In contrast the

apparent water permeability of the endolymphperilymph barrier remains unsolved since several different approaches did not yield unambiguous and quantitative results.
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Existence of both endolymph-perilymph barrier and blood perilymph barrier suggest: Perilymph as a source of endolymph. 1.Konishi,Hamrick et al `82: The compartmental distribution of radio active tracers is consistent with a K+ and Cl- exchange occurring between blood and perilymph and subsequent between perilymph and endolymph rather than directly between blood and endolymph. 2. Marcus & Thalmann 81: K+ free perfusion of both perilymphatic scales causes a rapid decline of the EP. Whereas K+ free perfusion of the vasculature is ineffective.
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Volume of the inner ear fluids

Data for this estimation have been generated by techniques like: Serial histological sections 3-D magnetic resonance microscopy. The total volume of all inner ear fluids spaces including cochlear and vestibular portions is approximately 204-228ml in humans and 21ml in guinea pigs.
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Cochlear endolymph volume (L)

Cochlear perilymph volume (L)

Endolymphatic sac & duct volume(L)

Humans

7.7

75.9

3.92

Guinea pig

1.6

12.1

0.12

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Pressure of Inner ear fluids

Compartments that influences fluid pressure include:
1.

2.
3.

The cranium, via CSF pressure Middle ear cavity Arterial and venous blood pressure transmitted by vasculature of the ear.

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The cochlear aqueduct plays a central role in regulation of perilymph pressure. Two conditions present: In animals where the cochlear aqueduct is patent, the pressure of CSF that is transmitted through the cochlear aqueduct dominates perilymphatic pressure.

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This includes both the static pressure and the pressure fluctuation in CSF associated with respiration, heartbeat, posture changes, coughing and sneezing. The CSF pressure averages to 11mm Hg in humans. Characteristics of pressure transfer from CSF to perilymph across the cochlear aqueduct vary in frequency dependent manner.
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It has been calculated that aqueduct acts as a low pass filter, attenuating CSF pressure fluctuations entering the perilymph for frequency above 20Hz. Pressure fluctuations below this frequency and sustained pressures are thought not to be attenuated by the aqueduct.

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When the cochlear aqueduct is occluded respiration induced pressure fluctuation in perilymph are highly attenuated. Experimentally induced CSF pressure changes result in smaller, delayed pressure response that is believed to be mediated by endolymphatic sac.

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Perilymph pressure- correlated with the arterial blood pressure changes during manipulation of systemic blood pressure.

Manipulation of middle pressure- results in larger perilymphatic pressure changes that recover slowly because of the diminished capacity of aqueduct to shunt pressure changes to the CSF.

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The endolymphatic space (ELS)- incompressible, flexible walled compartment suspended within the perilymphatic space. No hydrostatic pressure difference in the normal state. Similar pressure fluctuation associated with respiration and induced pressure changes of CSF in both endolymph and perilymph. - The static pressure differential across the basilar membrane, would decrease its compliance. The minimization of pressure differences between endolymph and perilymph is likely to contribute to the maintenance of high cochlear sensitivity to mechanical stimuli.
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Both endolymph and perilymph pressure fell to zero when the round window was perforated. - These studies confirm that perilymphatic pressure changes are transmitted to endolymph via mechanically compliant boundary membranes. Such pressure changes occur without endolymph volume change or movements of the membranous walls because endolymph is a fluid and it is thus incompressible.
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There are two viewpoints I. Mass action mechanism: The movement of stapes is transmitted directly to the fluid column in the cochlea, which responds as a whole. Inward movement of the stapes causes the perilymph to flow up the SV through the helicotrema and then down the ST. The round window is pushed outward by an amount directly proportional to the inward movement of the stapes. During the outward movement the direction of flow of the fluid column is reversed. Sound energy transmitted by the vibrating fluid column is selectively abroad by the B.M.
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Dynamics

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II. Alternative point is that the pressure generated in the SV is transmitted across the SM to the ST. Such a transmission of pressure result in distortion of vestibular membrane (Reissners membrane) and in turn the BM. There will be displacements of round window that are out of phase with the direction of movement of stapes. The fluid movement may be distinctive for a particular frequency- this produces the distortion of BM at the specific frequency.
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Functions of cochlear fluids

1. Transport of dissolved gases and nutrients between blood and many cell types of cochlea. 2. Transmission of acoustic vibrations from stapes to sensory structures. 3. Provision of suitable ionic environment for sensory hair cells, thus helping in physiological process of cells. 4. Removal of waste products. 5. Provision of chemical environment needed for transfer of energy from vibration (mechanical) to neural (electrical) signals.
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ROLE OF FLUID IN TRANSDUCTION.

The Standing Current: Von Bekesy (1950) : Described a +ve potential in the endolymphatic space and a ve potential inside the organ of corti. The presence of such potentials would drive a circulating current. This is the basis for Daviss Mechano-electrical Battery theory of cochlear transduction (1957).

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This theory postulates that acoustic stimulus itself does not need to generate the energy for transduction. The energy provided by a standing current flowing through the hair cells. Transduction will only change the resistance of the hair cells and there by the current flow

(Dallos `73).
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The Stria vascularis is believed to be solely responsible for the generation of the standing current. 1. It secretes K+ ions into endolymph and generates the large potential across the epithelial lining the scala media. 2. This trans epithelial potential of about +80mv is the EP and drives the standing current in conjunction with the ve membrane potential of the hair cells and steep gradient of K+ between endolymph and perilymph.
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The standing current generated by Stria vascularis flows radially through the SM towards 2 current sinks. One part of current flows through the organ of corti containing the sensory hair cells where it enters the ST Other part flows through the RM where the current crosses the SV.

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Zidanic and Brownel`90: Both branches of the

current return radially via the spiral ligament to the stria vascularis of the stria vascularis. The spatial secretion of the current source in the stria vascularis from the sensory cells by almost half a millimeter It may be pre requisite for the high sensitivity of the auditory system. This arrangement attenuates the noise originating from blood flow in the highly vascularised stria.

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Physiologic significance of fluid composition:

The specific ion concentration of endolymph and perilymph maintain structure and function in the cochlea. Variations in endolymphatic K+, Na+ or Ca2+ affect the conformation of the tectorial membrane (Kron ester frel,1979).

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