Solution

ID: past-exam-of-the-mathematical-physiology-course-of-the-university-of-oxford/2023/3/f/solution

On the descending part of the graph, mark the two tangency points where
Choose the crossings that enclose the portion with . For fixed strictly between these values and sufficiently small , the unique equilibrium is a repelling equilibrium by part (e). The endpoints here are leading singular limit values; at finite the boundaries instead solve and approach these tangencies when the crossings are nondegenerate.
There is a necessary qualification: such an interval exists only when is large enough that the descending graph actually attains slopes below . If , one needs . The supplied graph has a very steep descending section, but an arbitrarily small positive need not satisfy this condition. For , numerical evaluation gives , so the leading threshold is . A graph alone does not remove this parameter restriction.
The fast motion is especially informative. In fast time ,
At leading order is constant, so fast paths are straight lines with slope . Along such a line, the fast eigenvalue at the nullcline is
It is negative on the two outer attracting segments and positive on the intervening repelling segment. The tangencies defining are consequently the fold points of a critical manifold when it is expressed in the coordinates . They are not generally the maximum and minimum of itself.
For , slow motion on an attracting segment follows
On the left attracting segment this moves towards increasing ; on the right attracting segment it moves towards decreasing . Each segment ends at a tangency and is followed by a fast jump along a line of slope to the other attracting segment. This constructs the limiting relaxation oscillation. The figure uses , for which and ; the marked endpoints change with .
Figure 1.
Calcium nullcline, slope tangencies and the limiting relaxation cycle
. Details.
To justify existence of a periodic orbit of the full planar system, rather than just draw a formal cycle, first establish a compact trapping region. The positive quadrant is forward invariant: at , , and at , . Moreover
Choose above the initial store value. Since when ,
Thus the rectangle , is a positively invariant set if and the initial point lies inside it. Every forward solution is bounded. There are no equilibria on its axes for , and its sole interior equilibrium is a hyperbolic equilibrium point with both eigenvalues having positive real parts in the strict unstable range. A sufficiently small neighborhood of this repelling equilibrium has outward flow; deleting it leaves a compact trapping region with no equilibria. The Poincaré-Bendixson theorem then supplies a periodic orbit.
Hence periodic solutions exist throughout the strict unstable interval, interpreted with the finite- correction above. This argument establishes existence, without asserting uniqueness, the criticality of possible endpoint Hopf bifurcations, or absence of coexisting periodic orbits outside the interval.

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