  Understanding and manipulating the interactions between foreign bodies and
cell membranes during endo- and phagocytosis is of paramount importance, not
only for the fate of living cells but also for numerous biomedical
applications. This study aims to elucidate the role of variables such as
anisotropic particle shape, curvature, orientation, membrane tension, and
adhesive strength in this essential process, using a minimal experimental
biomimetic system comprising giant unilamellar vesicles and rod-like particles
with different curvatures and aspect ratios. We find that the particle wrapping
process is dictated by the balance between the elastic energy penalty and
adhesion energy gain, leading to two distinct engulfment pathways, tip-first
and side-first, emphasizing the significance of the particle orientation in
determining the pathway. Moreover, our experimental results are consistent with
theoretical predictions in a state diagram, showcasing how to control the
wrapping pathway from surfing to partial to complete wrapping by the interplay
between membrane tension and adhesive strength. At moderate particle
concentrations, we observed the formation of rod clusters, which exhibited
cooperative and sequential wrapping. Our study not only contributes to a
comprehensive understanding of the mechanistic intricacies of endocytosis by
highlighting how the interplay between the anisotropic particle shape,
curvature, orientation, membrane tension, and adhesive strength can influence
the engulfment pathway but also provides a foundational base for future
research in the field.
