Development of the unfertilized embryo sac and pollen tubes in the dwarf mistletoe Arceuthobium americanum (Viscaceae)

Abstract

Megasporogenesis, embryo sac development, and pollen tubes in Arceuthobium americanum Nutt. ex Engelm. were investigated with light, fluorescence, and electron microscopy. The orthotropous ovular structure of A. americanum lacked integuments and possessed a nucellus that was largely continuous with and indistinguishable from the placenta; we coined this structure the placental–nucellar complex (PNC). Two megasporocytes became evident in the tenuinucellate PNC by mid-April, and had undergone bisporic megasporogenesis by mid-May. The upper cell from each dyad (distal to the base of the PNC) became a functional megaspore, although only one would develop into a seven-celled embryo sac. Like typical angiosperm embryo sacs, that of A. americanum possessed an egg cell having the ultrastructure reflective of a quiescent cell, and lacked cellulosic and (or) hemicellulosic wall material between the egg apparatus and central cell. However, the egg apparatus arose at the lower embryo sac pole, not at the upper as expected for an orthotropous ovule. A hypothetical model for the development of Arceuthobium ovules is the ancestral fusion and subsequent reduction of two anatropous ovules to form two embryo sacs within the PNC, of which only one completes development. The synergids have no role in pollen tube guidance, as tubes could be seen below each functional megaspore prior to megagametogenesis.Key words: Arceuthobium, embryo sac, megasporogenesis, mistletoe, pollen tubes, ultrastructure.