On the Origin of Mitosing Cells by Lynn Sagan appeared in the March 1967 edition of the Journal of Theoretical Biology. At the time the article was published, Lynn Sagan had divorced astronomer Carl Sagan, but kept his last name. Later, she remarried and changed her name to Lynn Margulis, and will be referred to as such throughout this article. In her 1967 article, Margulis develops a theory for the origin of complex cells that have enclosed nuclei, called eukaryotic cells. She proposes that three organelles: mitochondria, plastids, and basal bodies, which are all parts of eukaryotic cells, were once free-living cells that took residence inside primitive eukaryotic cells. This process Margulis called endosymbiosis. Margulis' theory explained the origin of eukaryote cells, which are the fundamental cell type of most multicellular organisms and form the basis of embryogenesis. After fertilization, embryos develop from a single eukaryotic cell that divides by mitosis.

All cells that have a nucleus, including plant, animal, fungal cells, and most single-celled protists, also have mitochondria. Mitochondria are particles called organelles found outside the nucleus in a cell's cytoplasm. The main function of mitochondria is to supply energy to the cell, and therefore to the organism. The theory for how mitochondria evolved, proposed by Lynn Margulis in the twentieth century, is that they were once free-living organisms. Around two billion years ago, mitochondria took up residence inside larger cells, in a process called endosymbiosis, becoming functional parts of those cells. Within each mitochondrion is the mitochondrial DNA (mtDNA), which is different from the DNA in the cell's nucleus (nDNA). Organisms inherit their mitochondria only from their mothers via egg cells (oocytes). Mitochondria contribute to the development of oocytes, the release of the oocyte from the ovary (ovulation), the union of oocyte and sperm (fertilization), all stages of embryo formation (embryogenesis), and growth of the embryo after fertilization.