It stands to reason that the feminin came to be at the boundary between these.
[1] https://en.wiktionary.org/wiki/Reconstruction:Proto-Indo-Eur...
PS: This is fun ... Der Mensch -> Die Familie -> Das Dorf. Der Krieger, die Krieger, das Kriegen. Der Aktionär -> die Aktie. Die Aktionärinnen ... das Aktienpaket. Der Karren (car), Die Karre (load), das Ankarren, das Herangekarrte, das Karree (See [2] for more on four wheels, strange animism, etc. Ultimately inconclusive).
Note that the pluralizing morpheme -en was written -in in old german. Also note that many case inflections come out as -en, so I suppose that female forms were in the objectified part of speech more often, where the gender is not reflected in the article (den Kindern, Männern, Frauen).