Mechanisms of Endocytosis: Membrane Curvature
and Vesicular Scission
February 19, 2026
1 Introduction
Endocytosis is the fundamental process by which cells internalise extracellular fluid, so-
lutes, and macromolecules. This involves the de novo formation of vesicles from the
plasma membrane. This article focuses on the energetics of membrane deformation and
the molecular machinery, such as clathrin and dynamin, required to overcome the physical
barriers to scission.
2 The Energetics of Membrane Bending
The formation of an endocytic vesicle requires significant mechanical work to transform
a planar membrane into a highly curved spherical structure. According to the Helfrich
model, the bending energy per unit area, w, is:
1
w = κ(c1 + c2 − c0 )2 + κ̄c1 c2
2
Where:
• κ is the bending modulus.
• c1 and c2 are the principal curvatures.
• c0 is the spontaneous curvature.
• κ̄ is the Gaussian curvature modulus.
For a spherical vesicle of radius r, the total bending energy Ebend = 8πκ. Given that
κ ≈ 20 kB T , the energy required is roughly 500 kB T , necessitating the recruitment of
specialised proteins.
3 Clathrin-Mediated Endocytosis (CME)
CME is the most well-characterised pathway. Clathrin triskelions assemble into a poly-
hedral cage that provides a scaffold for the budding vesicle.
1
3.1 Cargo Recruitment
Adaptor proteins, such as AP2, bind to cargo-receptor complexes and the plasma mem-
brane. The density of cargo, σc , affects the rate of pit formation. The chemical potential
of a cargo molecule in the pit relative to the bulk membrane is:
ϕ
µ = µ0 + kB T ln − ϵbind
1−ϕ
Where ϕ is the area fraction of cargo and ϵbind is the binding energy to the adaptor.
4 Membrane Scission and Dynamin
The final stage of endocytosis is the ”pinching off” of the vesicle. This is an energetically
unfavourable topological change. The large GTPase dynamin polymerises around the
neck of the budding vesicle.
4.1 The Helical Constriction Model
Dynamin undergoes a conformational change upon GTP hydrolysis, resulting in a reduc-
tion of the helical pitch and radius. The torque, Γ, generated by dynamin must exceed
the resistance of the membrane:
dEmem
Γ≥
dθ
The work done during scission is supplied by the free energy of GTP hydrolysis,
∆GGTP ≈ 10–15 kB T per molecule.
5 Fluid-Phase Pinocytosis
Unlike receptor-mediated endocytosis, pinocytosis involves the non-specific uptake of ex-
tracellular fluid. The volume of fluid internalised, Vi , per unit time is:
dVi 4
= N · πr3
dt 3
Where N is the number of vesicles formed per second. This process is essential for
membrane recycling and controlling the surface-to-volume ratio of the cell.
6 Conclusion
Endocytosis is a triumph of biological engineering, converting chemical energy into the
mechanical force needed to remodel the lipid bilayer. By coordinating protein assembly
with membrane tension, the cell maintains its composition and internalises the vital
nutrients required for survival.