Mad11/Mad12 transheterozygotes exhibit abnormal L4/L5 wing veins, with no L2 or crossveins and a large margin notch.
Hemizygous viable. Transheterozygotes with Mad5 and Mad6 are viable and female sterile. Transheterozygotes with a deficiency or null Mad allele exhibit numerous defects in imaginal disc-derived structures.
Mad mutations can be placed in an allelic series based on relative severity of the maternal effect enhancement of weak dpp alleles: Mad1 < Mad7 < Mad12 < Mad10 < Mad5 < Mad3 < Mad2 < Mad11 < Mad6 < Mad4 < Mad8 < Mad9.