Oxyanion-hole stabilization is rarely a single trick: the sources converge on directional hydrogen-bond donors, geometric preorganization, and coupled proton or charge networks, so disrupting any one can weaken stabilization even when the others remain.
Backbone or side-chain donors can aim a hydrogen bond at the developing oxyanion and lower its energy, but effect depends on donor placement and angle, not donor count alone.
An active site can be shaped so donors are already aligned before binding, and loss of that shape can weaken stabilization even if the donor groups stay in place.
Nearby proton relays and charge-balancing residues can reinforce the oxyanion hole indirectly, so changes in the wider network can collapse stabilization without directly removing a key hydrogen bond.
The evidence points to a triad rather than a single mechanism, and the balance among donors, shape, and charge control can differ across enzymes.
The strongest stabilization comes from donor placement, a pre-shaped pocket, and a supporting electrostatic network working together.
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