88 THE ORIGIN OF GYNANDROMORPHS. 



or not at all, and no mating occurs. The copulatory organs show the 

 strangest combinations of the male and female type, but there are 

 still typical but rudimentary ovaries left. When the race, Jap. X 

 female is crossed to Eur. F male, a still higher degree of intersexuality 

 appears. Externally the daughters are "almost indistinguishable 

 from true males." The instincts are entirely male and the moths 

 try unsuccessfully to mate with females. The gonads look like testes, 

 but in sections show a mixture of ovarian and testicular tissue. A step 

 further and the daughters would be transferred into males. 



The next cross gives this final stage. When Jap. male is crossed 

 to any race of European female, only males are produced, i. e., all the 

 daughters become sons. 



The reverse picture is given by those combinations in which the 

 intersexes are sons partly changed over into daughters, a condition 

 that Goldschmidt terms male intersexuality. The wings are generally 

 streaked and in the extremest type only a few brown spots appear on 

 the wing-veins. The testis may contain some ovarial tissue, but the 

 changes in the gonads do not appear to run parallel to those seen on 

 the surface. 



The explanation that Goldschmidt offers for these intersexes is 

 entirely different from the explanation that is demonstrated for the 

 gynandromorphs of Drosophila. He accepts in part the chromosome 

 theory of sex determination and applies it to the present case on the 

 basis that the female is heterozygous for the sex chromosome Mm, 

 and the male homozygous MM. In addition, however, Goldschmidt 

 adds another set of sex-determining factors that he calls FF (inclosing 

 them in brackets) , which he locates in the cytoplasm, that is, outside 

 the chromosomal mechanism. These factors do not segregate (the 

 desirability of two F's is therefore not apparent) and are transmitted 

 from the female to her sons and daughters alike. The FF factors stand 

 for femaleness, which apparently includes the eggs, ovaries, secondary 

 sexual characters, and genitalia, in fact, all parts associated with the 

 female. The sex of a given individual is dependent on the balance 

 struck by the activity of the factors Mm and FF, one in the chromo- 

 somes and the other in the cytoplasm. 



The FF factors are supposed to be located in the cytoplasm because 

 if a certain numerical value is assigned to the egg, this value adheres 

 to the maternal line, no matter which sex chromosomes are introduced 

 from the male side in successive generations. If the factors for female- 

 ness were carried by the male and like other paternal characters in- 

 fluence the cytoplasm, their value would be affected by the kind of 

 males that were employed ; but Goldschmidt has sho-v^Ti that his results 

 work out on the assumption that no such effects need be postulated. 



There is, however, another way in which the inheritance of certain 

 factors along the maternal line may be treated. Goldschmidt has 

 himself admitted this as a possible interpretation, although he has 



