Pollen-tube growth and fertilization mode in Gymnostoma (Casuarinaceae): their characteristics and evolution

A unique mode of fertilization called "chalazogamy", whereby the pollen tube passes through the chalaza instead of the micropyle, is known in several species of derived genera in Casuarinaceae. In this paper we report the occurrence of chalazogamy in Gymnostoma ( G. poissonianum), the most primitive genus in the family. We also show that the pollen tube grows discontinuously from the stigma to ovules in about 3 months. At the time of pollination, the ovules have not yet formed in the ovary, and require a long time to develop. The pollen tube(s) lie in a zigzag line and are branched in the upper region of the style, and their growth is arrested there until the ovary develops further. Studies of the relevant literature further revealed discontinuous pollen-tube growth in relation to a prolonged period between pollination and fertilization, as well as chalazogamy, in Betulaceae, Juglandaceae and/or Fagaceae that are closely related to Casuarinaceae. This feature may have derived early in the evolution of Fagales.