The cell conservation equation equates the change of cell number density to minus the divergence of its total flux. The flux is advection by the water, is the mean swimming flux, and is the dispersal flux. The tensor describes effective translational diffusion, whereas in part (i) describes rotational diffusion; they have different dimensions. This conservation law neglects birth, death and sources at the walls.
The equation expresses incompressible flow. In the momentum equation, is the reference suspension density, is material acceleration times density, is the pressure force after subtracting the pure-water hydrostatic pressure, and is the viscous force for constant dynamic viscosity in a Newtonian fluid. Since counts cells per volume, in is excess downward weight per cell; it is not an acceleration. The small cell-fluid density difference permits the Boussinesq approximation: retain that difference in buoyancy while using constant elsewhere. Diluteness permits neglect of cell-cell interactions, changes of viscosity, and active cell stresses. Any separate settling speed is assumed negligible or already incorporated in the specified mean drift. The light strength and swimming speed are uniform.
Consider the intended long-channel bulk approximation. At the sidewalls impose zero normal cell flux and the no-slip boundary condition. The closed ends require zero total fluid flux through each horizontal section, so ; localized turning regions are omitted from the bulk equations. The mean concentration is . The final paragraph below distinguishes these bulk conditions from exact closed-end cell conservation.
For , the vorticity is . Hence , and the leading horizontal mean swimming direction is . With and , steady cell conservation equation says that is constant. Its wall value is zero, so with . The phototactic concentration in a vertical channel is therefore
This result neglects the correction to horizontal swimming. For positive phototaxis, and cells accumulate towards the illuminated wall .
The steady inertial terms vanish for , since . The horizontal momentum equation gives . The vertical momentum equation then requires a constant excess pressure gradient and gives . Twice integrating produces
Set . The two no-slip boundary conditions give and . The zero fluid-flux condition follows by integrating this expression:
Consequently the remaining constants are
The resulting velocity can also be written without separate polynomial constants as
Here is the zero-flux pressure gradient in a cell-driven channel. It is negative for : pressure decreases upwards, creating the upward pressure force that balances the excess cell weight and drives the return flow. Its magnitude is determined by zero net fluid transport, not by prescribing pressure independently. In the zero-light limit, the continuous result is , and .
For the requested large- sketch, cells occupy a boundary layer of thickness near . Put and . The exact dimensionless profile above is
Away from the illuminated-wall layer it tends to . There is broad upward return flow for and downward flow on the illuminated side. The outer maximum is at . Near , the exponential term is approximately and restores the no-slip boundary condition from the negative outer velocity to zero at the wall. The finite- profile shown below retains that layer. The small-vorticity assumption must still be enforced, for example by sufficiently weak excess weight or a sufficiently dilute suspension; alone does not ensure .
Figure 1.
Downward flow near the illuminated wall and upward return flow in a phototactic channel
. Details.
There is a genuine global qualification to the bulk construction. For an exactly steady channel with impermeable ends as well as impermeable sidewalls, the closed-channel cell-flux constraint also requires at every height. In this bulk ansatz the first-order orientation closure gives
since integration by parts, and imply . Moreover, multiply by and integrate. The wall values and zero fluid flux give
for a nontrivial profile. Thus zero axial cell flux generally fails unless the extra factor vanishes, or the flow is trivial. A complete steady closed-channel solution generically needs axial concentration variation. The explicit profile solves the stated bulk equations, sidewall conditions, concentration normalization and zero fluid-flux condition; it is not, for arbitrary parameters, an exact globally closed steady cell distribution. Ignoring end regions gives the intended bulk approximation, but does not by itself prove the stronger global existence claim.
Neglecting cell rotational inertia, the light and viscous torque sum to zero. Therefore the angular velocity is . A material orientation evolves by . The vector triple-product identity gives
The last term is the projection of onto the tangent plane to the unit sphere. In particular, , as required to preserve the unit orientation. It aligns the swimmer towards the light; the vorticity rotates that orientation.
For rotational diffusion coefficient , the phototactic orientation Fokker-Planck equation balances deterministic orientation drift with diffusion on the sphere. Its density is normalized with respect to solid angle, not . When , there is no azimuthal drift and the zero-current condition is . Thus , and normalization gives
The limiting density at is .
To justify the angular dependence of the first correction, define the operators
The steady Fokker-Planck equation is . Its regular expansion therefore gives
The coefficients of are independent of , so each azimuthal Fourier harmonic is preserved. The forcing has only the sine harmonic with azimuthal number one. There can be no extra unforced harmonic in a smooth normalized first correction: writing a homogeneous solution as gives . Multiplication by and integration by parts on the sphere gives , hence is constant. Normalization removes that multiple of . It follows that
No explicit calculation of is needed. Its dependence is on and the parameter , and its behavior at the poles must make the density smooth.
In the stated spherical coordinates, . Azimuthal integration makes the leading components and the first-order corrections vanish. The surviving coefficients are
Thus the small-vorticity phototactic orientation response is
Both coefficients depend only on . In particular, is the Langevin function of and is positive for positive light strength; its integral representation above suffices.
For isotropic rotational diffusion, orientation drift and probability density obey the displayed Fokker-Planck equation, normalized with respect to solid angle. Without flow, constant light gives the zero-current density , . Integration over the sphere gives unit mass; its mean projection is the Langevin function. The zero-light limit is the uniform density .