Sutton and Boveri conducted experiments to understand the pattern of inheritance of traits in organisms.

Which among the following statements was /were understood as the result of experiments by Sutton and Boveri?

1: Genes are located on homologous chromosomes.

2: All genes on the same chromosome are linked.

3: Genes on homologous chromosomes get separated during anaphase of meiotic division.

4: The frequency of gene pairs on the same chromosome is a measure of the distance between the genes.