Fertilization in Plumbago zeylanica: entry and discharge of the pollen tube in the embryo sac

Abstract

The ultrastructural organization of the megagametophyte of Plumbago zeylanica, which lacks synergids, was examined in chemically and physically fixed ovules after entry of the pollen tube. Similar to angiosperms with conventionally organized megagametophytes, the pollen tube enters the ovule through a micropyle, formed by the inner integument, and approaches the female gametophyte by growing between nucellar cells. Unlike other described female gametophytes, however, continued pollen tube growth results in direct penetration of the base of the egg through cell wall projections forming a filiform apparatus and is completed between the egg and central cell without disrupting either of these cells' plasma membranes. A terminal pollen tube aperture forms when the pollen tube reaches an area of strong curvature near the summit of the egg; this results in the release of two sperm cells, the vegetative nucleus, and a limited amount of pollen cytoplasm. The formerly continuous chalazal egg cell wall is locally disrupted near the tip of the pollen tube and apparently is thus modified for reception of male gametes. Discharged pollen cytoplasm rapidly degenerates between the egg and central cell, but unlike pollen tube discharge in conventionally organized megagametophytes, it is unassociated with the degeneraton of any receptor cell within the female gametophyte. Sperm nuclei are transmitted, one to the egg and the other to the central cell, to effect double fertilization by nuclear fusion with their respective female reproductive nuclei. The vegetative nucleus and discharged pollen cytoplasm degenerate between the developing embryo and endosperm during early embryogenesis. The emerging concept that the egg of Plumbago possesses combined egg and synergid functions is supported by the present study and suggests that the megagametophyte of this plant displays a highly specialized egg apparatus composed exclusively of a single, modified egg cell.