 Solution

Write ${\ell }_g$ for the dimensional offset of the centre of mass, reserving $h$ below for the dimensionless swimming parameter. The weight $-mg\mathbf{k}$ acts at $-{\ell }_g\mathbf{p}$, so its torque about the geometric centre is $mg{\ell }_g\mathbf{p}\times \mathbf{k}$. Buoyancy acts through the geometric centre and has no such torque. For a spherical cell, the ambient angular velocity is $\mathbf{\omega }/2$ and the rotational resistance is ${\zeta }_r=\rho \nu {\alpha }_{\perp }v$. Neglecting angular inertia, torque balance givesSince $\dot{\mathbf{p}}={\mathbf{\Omega }}_{\mathrm{cell}}\times \mathbf{p}$, the bottom-heavy spherical-cell orientation dynamics isThus a steady orientation satisfiesThe printed algebraic equation has the opposite sign on its right side, if vorticity means the usual $\nabla \times \mathbf{u}$. Its later concentration equation is consistent with the corrected sign. In particular, linearizing the corrected equation about $\mathbf{p}=\mathbf{k}$ gives ${\mathbf{p}}_{\perp }=(B/2)(\mathbf{\omega }\times \mathbf{k}{)}_{\perp }$, whereas the printed sign would reverse that response. Merely changing the definition of vorticity would also require changing the ensuing fluid and orientation calculations.

The gyrotactic reorientation time $B$ measures gravitational restoration against rotational viscous resistance. In still fluid a tilt angle obeys $\dot{\theta }=-\sin \theta /B$, soFor small tilt it is precisely the exponential relaxation time. Applying the algebraic orientation balance to a time-dependent disturbance additionally assumes that this relaxation is fast enough to neglect orientation lag. Keeping the full dynamic orientation equation would introduce an additional relaxation factor into the perturbation problem.