The embryology of Arceuthobium pusillum
Megasporogenesis in Arceuthobium pusillum begins in March–April, 19 months after emergence of the pistillate aerial shoots. The embryo sacs are fully developed within 1 month. The first division results in a functional dyad and a nonfunctional dyad. The nucleus in the nonfunctional dyad divides once and the resulting nuclei disintegrate. Meiosis in the functional dyad follows immediately. Development of the embryo sac is of the bisporic Allium type. Staminate and pistillate aerial shoots flower in April–May. Fertilization occurs within several days after pollen has adhered to and germinated on the stigmas of pistillate flowers. One sperm nucleus fuses with the egg nucleus, the other with the polar nuclei. A haustorial extension then elongates downward into the ovarian papilla and becomes filled with a file of endosperm cells. Concurrently a uniseriate layer of endosperm has developed around the zygote. The endosperm then grows to a rather massive size before the zygote begins development. A transverse wall forms after the first division of the zygote. The terminal cell is bisected first, followed by bisection of the basal cell. Succeeding anticlinal and periclinal divisions result in growth of the proembryo. Embryonic protoderm is visible as a distinct tissue by the time the embryo has assumed an ovoid shape. When the seeds are forcibly expelled from their fruits in mid-September, the embryo has four distinct histological zones: protoderm, promeristem, ground meristem, and procambium.