 Solution

The quantities $n_i(\tau ,t)$ are cell-cycle age densities: $n_i(\tau ,t)d\tau$ counts biological cells whose time since entering phase $i$ lies in $[\tau ,\tau +d\tau ]$. Here $\tau$ is time spent in the current phase, rather than time since birth. In this two-stage approximation to the cell cycle, a biological cell passes from G1 phase to G2 phase, then completes cell division and produces two daughters entering G1 phase.

The time derivative measures change at fixed phase age. The age derivative transports biological cells towards larger $\tau$ at unit speed, because $d\tau /dt=1$. Thus each equation is an age-structured population equation. The nonnegative hazard function ${\Gamma }_i(\tau )$ means that, conditional on still being in phase $i$ at age $\tau$, the probability of leaving during the next $dt$ is ${\Gamma }_i(\tau )dt+o(dt)$. Its product with $n_i$ is the loss from that phase-age class; it is not an additional death term.

The boundary condition at $\tau =0$ specifies the incoming flux. Every G1 phase exit contributes one entrant to G2 phase, whereas every G2 phase exit contributes two entrants to G1 phase. The initial conditions specify the phase-age composition at $t=0$. Consequently integration over phase age gives the useful population balanceEach completed cell division increases the total number of biological cells by one.