A model of the hair bundle's spontaneous oscillation.
(A) The channel open probability equilibrates with the
tension in the gating springs on a time scale much faster than that of
adaptation. Under this quasistatic condition, a free bundle must reside
at a point of zero force. The region of negative slope stiffness in the
initial displacement-force relation (dashed black curve) implies that
the bundle is bistable: there are two such points (green stars).
Assuming that the bundle first occupies the negative stable point, the
transduction channels' low open probability causes adaptation to shift
the displacement-force relation in the negative direction. The shift
proceeds with a slope set by the stiffness of the stereociliary pivots
(ref. 15; dotted green lines). When the relation's left-sided local
maximum becomes tangent to the abscissa (at point 1 on the blue curve),
however, the negative stable point vanishes, and the bundle must leap
to the positive stable point (point 2) to maintain the zero-force
condition. For the parameter values of Fig. 1C, this
transition corresponds to an abrupt increase in the channel's open
probability from 0.15 to 1.0. At the bundle's new position, the
channels' high open probability promotes adaptation in the opposite
direction until the bundle's position corresponds to the right-sided
local minimum of the displacement-force relation (point 3 on the red
curve) and the bundle jumps in the negative direction (to point 4). The
channel's open probability correspondingly plummets from 0.85 to 0.0.
Oscillation ensues from repetition of this sequence. (B)
Trajectory (1 → 2 → 3 → 4) of the hair bundle along the
displacement-force relation as this relation undergoes the adaptive
shift depicted in A. The double arrows indicate the fast
transitions across the unstable region of negative stiffness (1 → 2
and 3 → 4), whereas the single arrows demarcate the slow adaptive
movements along the stable branches of the relation (2 → 3 and 4 →
1). C, Hair-bundle oscillation produced by a model of
bundle mechanics that incorporates the parameter values for the cell
whose response is depicted in Fig. 1A. The
numbers relate phases of the oscillation to points on the
displacement-force relations of A and
B.