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. 2021 Mar 22;19:1806–1828. doi: 10.1016/j.csbj.2021.03.019

Table 1.

Overview of protein secretion systems and the substrate features in Gram-negative bacteria.

Secretion system Secretion step(s) Membrane spanning Secretion signal Substrate state
Sec 1 Inner N-terminus Unfolded
Tat 1 Inner N-terminus Folded
T1SS 1 Inner + Outer C-terminus Unfolded
T2SS 2 (Sec/Tat) Inner + Outer N-terminus Folded
T3SS1 1 or 2 (Sec) Inner + Outer (+Host) N-terminus Unfolded
T4SS2 1 Inner + Outer (+Host) C-terminus Unfolded
T6SS 1 Inner + Outer + Host N-terminus? Folded
T5SS 2 (Sec) Outer N-terminus Unfolded
Pili/ T7SS 2 (Sec) Outer N-terminus Folded
Curli/ T8SS 2 (Sec) Outer N-terminus Unfolded
T9SS 2 (Sec) Inner + Outer C-terminus Folded

Notes: 1 T3SSs include non-flagella T3SSs and flagella T3SSs. Non-flagella T3SSs are translocation systems delivering substrates into host cells in one step, while flagella T3SSs involve two steps to secrete substrates extracellularly. 2 T4SSs translocate substrate proteins into host cells like T3SSs, or transport the proteins into extracellular milieu.