## Problem 159 — rank:276

**Title:** Standing variation versus new mutation in adaptation

**Added binding assumptions (LaTeX):**

```latex
Under epistasis there is no canonical nonnegative additive ``fraction of adaptive effect.'' Fix an attribution rule assigning nonnegative weights to contributing allele copies or predefined additive trait components; require positive total weight and define $Z$ as the pre-change-weight sum divided by the total. State that this is rule-dependent. Evaluate at the first threshold-hitting time on $A$, and propagate uncertainty in mutation origins, linkage and unsampled standing variants.
```

**Direct edits:** 0. **Fresh external checks:** None; supplied sources retained as provenance.

