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Inheritance studies of testa color in the cultivated peanut ( Arachis hypogaea L.) began in the Netherlands East Indies (Java), now Indonesia, by J. E. van der Stok in 1910. He was the first to report the results from crossing peanut plants with red‐brown seed coat (red testa) × plants with light red (pink testa). The first (F 1 ) generation was red, and the second (F 2 ) generation segregated in three times as many plants with red as with pink (3:1 ratio). Currently, there are five basic testa colors known in the peanut (white, tan, pink, red, and purple) with variegated testa having red and white, purple and white, purple stripes on tan, and white spots on red. Genetic inheritance studies over the past 100 yr have established at least 12 genes controlling basic testa color ( F 1 , F 2 , D 1 , D 2 , R 1 , R 2 , R 3 , P , W 1 , W 2 , Wh 1 , and Wh 2 ). These genes interact with at least five variegated genes ( V , Vsp 1 , Vsp 2 , Wsp 1 , and Wsp 2 ) to result in the phenotypic testa color expression found within the present‐day peanut germplasm collections.
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W. D. Branch (2010) studied this question.
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