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The genetic basis for a striking aleurone color variegation of a maize population from Colombia is described. The variegation is the result of the interaction of a particular r allele, designated r-cu , and a specific factor, Fcu (Factor Cuna), that collectively compose a controlling element system. In the absence of Fcu, a variable dilute (nonvariegated) pigmentation is expressed in the aleurone. The degree of variegation is determined by differences in the dosage of the Fcu element; differences in variegation between kernels with one r-cu vs. two r-cu's cannot be detected. Because of the specificity of the interaction, Fcu is different from the three other well-known controlling elements, Dt, Ac and En(=Spm), and therefore represents a fourth controlling element system in maize. In addition to Fcu, the Cuna tribal maize contains two other controlling elements, Dt and a very weak En. The prevalence of controlling elements in natural maize populations is discussed.

Controlling-elements in a tribal maize from Colombia: FCU, a two-unit system