Osmolarity, tonicity and three distinct readings
Osmolarity counts dissolved particles; tonicity describes a solution's effect on the volume of a particular cell. They can differ when the membrane passes a solute: an isoosmotic urea bath can be hypotonic because urea equilibrates across the membrane.
Each compartment is therefore summarised by three sums. They are not redundant results: each answers a different question.
C is in mol/L; i and σ are dimensionless; T is in kelvin and R = 8.314462618 J·mol⁻¹·K⁻¹. The factor 1000 converts litres to cubic metres.
Reflection coefficient and initial flux
Staverman's reflection coefficient σ measures how strongly a solute gradient contributes to osmotic water flow: σ = 1 represents complete reflection and σ = 0 no effective contribution. It is neither a literal fraction of crossing molecules nor a replacement for permeability. In this model, a solute marked impermeant uses σ = 1.
Jv> 0 is defined as outward flow: a positive effective difference produces initial water efflux; a negative one produces influx. Lp is the hydraulic permeability of the membrane.
Equilibrium volume
Over time, permeant solutes equalise their concentrations and drop out of the balance. Intracellular impermeant osmoles remain trapped and conserved; at equilibrium their final concentration equals the impermeant concentration in the bath.
If the outside contains no impermeant osmoles, no finite equilibrium volume exists; if they are absent only from the inside, the model leads to collapse.
Transient reversal
Initial direction depends on effective osmolarity; final volume depends on impermeant osmolarity. Defining both differences as outside minus inside, the two point in opposite directions when:
This pattern can arise while loading permeant cryoprotectants: a cell first shrinks and then recovers volume, or vice versa. Only the initial instant and equilibrium are calculated here, not the time course between them.