In this delayed carbon-dioxide ventilation feedback model, is a representative carbon-dioxide partial pressure, an effective storage capacitance, metabolic production, and removal by pulmonary ventilation. The delay represents transport and sensing between the gas-exchange compartment and the ventilatory response. The Hill equation response rises sigmoidally with pressure, reaches half its maximum at , and saturates at ; permits a steep threshold-like response.
For positive parameters, increases strictly from zero to infinity because its derivative is . Its graph therefore intersects the horizontal line exactly once: . The figure shows this argument and the first imaginary-root crossing.
Figure 1.
A unique respiratory equilibrium can become unstable through delayed feedback
. Illustrative dimensionless Hill response with , , , and equilibrium . The second panel separates the instantaneous-damping and delayed-gain vectors at the first imaginary root. Details.
For a small perturbation , linearization gives
The sigmoidal delayed ventilation threshold follows from the characteristic equation of a delay differential equation, . At an imaginary root , separate real and imaginary parts:
For , the first positive crossing has
At zero delay the scalar equation is stable. No root can cross the imaginary axis before this first crossing, and zero itself is not a root. To check the crossing direction, implicit differentiation gives
All subsequent imaginary crossings have the same direction. Thus the equilibrium is linearly stable below and unstable above it. This establishes the oscillatory linear threshold; the full nonlinear Hopf bifurcation additionally requires the usual nondegeneracy conditions. If , no finite imaginary-root crossing occurs and the equilibrium is stable for every finite delay. For the stated response, , showing explicitly when the gain can exceed instantaneous damping.
If , then and . Consequently the small-amplitude period near onset is . The comparison is to the critical delay near the first instability, rather than an arbitrary delay far beyond onset. A longer circulatory transit time can move the physiological system across this delayed-feedback threshold, providing a mechanism for periodic breathing associated with impaired cardiac output.
In delayed carbon-dioxide ventilation feedback, is the effective carbon-dioxide partial pressure of the lumped body compartment. Its stored carbon-dioxide amount is proportional to , with effective capacitance , so is its accumulation rate. Metabolism produces carbon dioxide at rate . Ventilation removes it at a rate proportional to partial pressure times ventilation, , with gas-conversion constants absorbed into the model coefficients. The equation is therefore a one-compartment carbon-dioxide balance: production minus respiratory removal equals accumulation. It idealizes mixing and treats metabolic production as constant over the time scale of interest.