Full title: Collective cell migration due to guidance-by-followers is robust to multiple stimuli Authors: R. Müller, D. Jahn, J. Starruß, A. Boutillon, N. B. David, L. Brusch Contributors: D. Jahn Date: 10.02.2023 Software: Morpheus (open-source). Download from https://morpheus.gitlab.io Model ID: https://identifiers.org/morpheus/M0008 Units: [time] = min, [space] = μm Reference: This model is the original used in the publication, up to technical updates: R. Müller, A. Boutillon, D. Jahn, J. Starruß, N. B. David, L. Brusch: Collective cell migration due to guidance-by-followers is robust to multiple stimuli. Front. Appl. Math. Stat. 9: 2297-4687, 2023. https://doi.org/10.3389/fams.2023.1163583 Comment: Extension of the guidance-by-followers mechanism (https://identifiers.org/morpheus/M0006) comparing robustness of various integrations of multiple simultaneous cell-cell contact stimuli.

1 Strength of the DirectedMotion plugin Strength of the DirectedMotion plugin Angle between cell movement and direction to the neighbor. Below max_angle, the cell is considered as moving towards the neighbouring cell and will influence its direction Minimal velocity for a cell to influence its neighbors Needed for NeighborhoodReporters neighbor_velocity_x and neighbor_velocity_y time == init_time axial_cell_front_y polster_cell_front_y (axial_cell_front_y - axial_cell_front_y_init) / (time - init_time) (polster_cell_front_y - polster_cell_front_y_init) / (time - init_time) Total number of polster cells orientated ±45° polster_cells_orientated_total/polster_cells_total 0 = none, 1 = contact-weighted mean, 2 = largest contact takes all, 3 = min angle takes all celltype.cell.size Cell colors: - white: uninitialized cells (time >= init_time) - yellow: axial cells (RandT_or_Mech_motion_strength == 0) - red: RaT polster cells not orientated ±45° (mech_induced_dir.abs == 0 and !polster_cell_orientated) - orange: guided polster cells not orientated ±45°(mech_induced_dir.abs > 0 and !polster_cell_orientated) - bright green: guided polster cells orientated ±45°(mech_induced_dir.abs > 0 and polster_cell_orientated) - dark green: RaT polster cells randomly orientated ±45° (mech_induced_dir.abs == 0 and polster_cell_orientated) if(time >= init_time, if(RandT_or_Mech_motion_strength > 0, if(mech_induced_dir.abs > 0, if(polster_cell_orientated, 5, 3), if(polster_cell_orientated, 6, 2)), 4), 1) After init_time, set the properties of axial cells time == init_time and cell.center.y <= 360 directed_motion_strength_global 0 Set to strength = 0 for polster cells After init_time, set the properties of polster cells time == init_time and cell.center.y > 360 RandT_or_Mech_motion_strength_global 0 (time >= tumble.last + tumble.run_duration) time run_duration_adjustment * rand_gamma(0.5, 5) angle, 0 , 1

Needs time-step = 1

Needs time-step = 1

py_induced_dir_x, py_induced_dir_y, 0.0 if(mech_induced_dir.abs > 0, mech_induced_dir, RaT_dir) RandT or mechanically induced movement. Set to strength = 0 for axial cells cell.center.y * (RandT_or_Mech_motion_strength == 0) cell.center.y * (RandT_or_Mech_motion_strength > 0) The axial cell furthest ahead determines the axial cell front

The polster cell furthest ahead determines the polster cell front

Returns: - '1' if cell is a polster cell - '0' if cell is a polster cell RandT_or_Mech_motion_strength > 0 and time >= init_time

Returns: - '1' for polster cells moving in y direction ±45° - '0' for polster cells moving in other directions and all axial cells RandT_or_Mech_motion_strength > 0 and velocity.phi >= 1/4 * pi and velocity.phi <= 3/4 * pi Sums up polster cells oriented ±45°

Transform the angle of the cell from Morpheus' 0° in x-axis direction to 0° when the cell moves in y-axis direction. The direction of movement of the cell is then given as an absolute difference in degrees from the ideal movement in the y-axis direction. atan2(-cos(velocity.phi), sin(velocity.phi)) / pi * 180 1 3 1 For plot 'Velocity Angle ±45°'

For plot 'Velocity Angle Distribution'

Do not turn off. Helper logger ensuring that the VectorEquations for 'mech_induced_dir' and 'dir' are periodically evaluated by the TimeScheduler.