Synthesis of the amino acid histidine is a multistep anabolic pathway that uses the products of 13 genes (hisA to hisM) in E. coli. Two independently isolated  E. coli mutants, designated his1⁻ and his2⁻ are studied in a conjugation experiment. A his⁺ F' donor strain that carries a copy of the hisJ gene on the plasmid is mated with a his1⁻ recipient strain in Experiment 1 and with a his2⁻ recipient in Experiment 2. The exconjugants are grown on plates lacking histidine. Growth is observed among the exconjugants of Experiment 2 but not among those of Experiment 1.

What is the genotype of exconjugants in Experiment 2?