After rediscovering Mendel's laws of heredity, which apply to
chromosomal inheritance, he undertook experiments with the
four o'clock (
Mirabilis jalapa) to investigate apparent counterexamples to Mendel's laws in the heredity of variegated (green and white mottled) leaf color. Correns found that, while Mendelian traits behave independently of the sex of the source parent, leaf color depended greatly on which parent had which trait. For instance, pollinating an
ovule from a white branch with pollen from another white area resulted in white progeny, the predicted result for a
recessive gene. Green pollen used on a green stigma resulted in all green progeny, the expected result for a
dominant gene. However, if green pollen fertilized a white stigma, the progeny were white, but if the sexes of the donors were reversed (white pollen on a green stigma), the progeny were green.
This
non-mendelian inheritance pattern was later traced to a
gene named
iojap which codes for a small protein required for proper assembly of the
chloroplast ribosome. Even though
iojap assorts according to Mendel's rules, if the mother is
homozygous recessive, then the protein is not produced, the chloroplast ribosomes fail to form, and the plasmid becomes non-functional because the ribosomes cannot be imported into the
organelle. The progeny could have functional copies of
iojap, but since the chloroplasts come exclusively from the mother in most
angiosperms, they would have been inactivated in the previous generation, and so will give white plants. Conversely, if a white father is paired with a green mother with functional chloroplasts, the progeny will only inherit functional chloroplasts, and will thus be green. In his 1909 paper, he established variegated leaf color as the first conclusive example of
cytoplasmic inheritance.