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. 2021 Jul 26;10:e71585. doi: 10.7554/eLife.71585

Figure 1. Structure and activity of cohesin.

Figure 1.

(A) Structural overview of the cohesin complex subunits. The two main subunits Smc1/Smc3 (gray and yellow) associate at a hinge on one end and at their ATPase heads on the other. These subunits form a ring-like shape when they are bound by a long flexible kleisin protein (magenta). Kleisin also interacts with additional subunits: the cohesin loader complex (green and blue) and an Scc3 protein (orange). The ATPase heads form a gate that DNA can go through called the head gate, while the interaction between kleisin and the ATPase head of Smc3 forms another gate called the N-gate. (B) Topological entrapping: the kleisin N-gate opens, allowing a free strand of DNA to enter it. The N-gate then closes, trapping the DNA strand between the N-gate and the head gate. Once in the DNA gripping position, the head gate is opened by ATP hydrolysis to complete DNA entry. The Smc hinge moves away from the ATPase heads straightening up the cohesin, and leading to topological DNA entrapping. (C) Loop extrusion: in this case, a free DNA strand becomes bound by the hinge module and the ATPase heads forming a loop without crossing either gate. Since the kleisin N-gate does not open, even if the head gate is opened by ATP hydrolysis, DNA cannot cross the cohesin ring. Once in this alternative DNA gripping position, Smc hinge, with Scc3 and DNA bound, moves away from the ATPase heads, pulling the DNA with it, and lengthening the loop. (D) Schematic for the loop extrusion process. Step 1: in the DNA gripping state, prior to ATP hydrolysis, the two DNA binding modules (hinge and heads) are in close proximity. Step 2: following ATP hydrolysis, the hinge, along with Scc3 bound to DNA, detaches from the head and moves away like a swing. This swinging motion of the hinge module is considered to be driven by diffusion (biased Brownian motion), and pulls the DNA with it. Steps 3 and 4: the next loop extrusion cycle proceeds to promote loop growth when the hinge module returns to form a new gripping state.