Resting cells enter proliferation at rate . A proliferating cell takes a fixed cycle time to divide and survives that cycle with probability under a constant death hazard . Thus exit from the proliferating compartment at time is , and it returns twice as many daughter cells to the resting population. Resting-cell loss at rate is separate.
More explicitly, the number currently in the proliferating stage is
for a consistent initial history. Differentiation gives the stated equation. Counting resting-cell entry, loss and daughter-cell return gives its equation. Negative feedback in suppresses recruitment when the resting population is high; its Hill exponent gives a steep response around the concentration scale .
Set and define
Then , so the resting-cell equation becomes exactly
The nonzero steady state solves , or for positive . The latter right side decreases strictly from to zero. Thus when there is exactly one positive equilibrium,
Zero is also an equilibrium; negative populations are outside the biological domain. The same result is visible by intersecting the line with the hump-shaped .
Figure 1.
Resting-cell recruitment feedback has one positive equilibrium when the small-population reproduction factor exceeds one
. Details.
Put . Linearization and a mode give
Here is the coefficient called in the source's characteristic equation; it is not the proliferating-cell death hazard. Any characteristic root with positive real part makes the equilibrium linearly unstable. A nonzero imaginary crossing satisfies and ; these are its Hopf conditions, subject to nondegeneracy.
On slow time , the unit delay becomes . Taylor expansion gives . Dividing the equation by therefore yields
This approximation requires slow smooth variation and a denominator bounded away from zero; a vanishing denominator marks failure of this slow reduction. On the fast time , omit only the explicit order- terms, retaining the finite delay:
Replacing the delayed argument by the present value on that fast scale would incorrectly erase the dynamics. These are the slow and fast limits of a delayed stem-cell model.