I am making my way through the linked “Nature” artricle.
I don’t understand all of the genetic technical terminology, but I can follow the conclusions. There are several relevant points.
The “slow graying” genetic variant, aka G2, is relatively rare. They only found two groups - a family of Connemaras in Sweden and a family of 5 TBs in Japan, with the G2 versions. MOST grays have the G3, or “fast graying” variant. (Note that they use the term G1 for the non-gray variant)
The “slow graying” horses maintain their “base coat” color, but getting lighter, until their teens. They only start to look dappled gray in their late teens, and never lose the dapples, even at age 30. " If a slow greying foal has face markings (like a star or blaze) the white parts are not well-defined and the white hairs will spread out in time." “The first signs of greying in a slow greying horse will normally not appear until 5–7 years of age, with grey hairs often expanding from white head markings.” “When a slow greying horse has reached an age of about 15 years the coat color may turn into a dappled steel or bluish grey shade.” “…they will never become a visually white horse.”
Within both families there were some descentants with the slow graying variant, and some with the fast graying variant. They examined 25 “late graying” horses between 15 and 35 and found no sign of melanomas on any of them.
In the general population, 50% to 80% of gray horses in this age group show signs of melanoma. Even though they only examined a small population of “slow graying” horses, this difference is statistically “highly signifcant” ( Χ 2 = 25.0, d.f. = 1; P = 6 × 10-7).
One of the conclusions is:
“The G1/G1 genotype shows no greying and low incidence of melanoma, G1/G2 shows slow greying and low incidence of melanoma, G1/G3 shows fast greying and high incidence of melanoma, and G3/G3 homozygotes grey very fast and show a very high incidence of melanoma as previously demonstrated.”