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Efficient plant male fertility depends on vegetative nuclear movement mediated by two families of plant outer nuclear membrane proteins
Edited by Ueli Grossniklaus, Institute of Plant Biology and Zürich-Basel Plant Science Center, University of Zurich, Zurich, Switzerland and accepted by the Editorial Board July 8, 2014 (received for review December 16, 2013)

Significance
Double fertilization of the female gametophyte is vital for angiosperm reproduction and requires delivery of sperm cells through a pollen tube. The sperm cells and pollen vegetative nucleus together are called the male germ unit. The pollen tube grows a long distance from the stigma to the ovary, and the mode of male germ unit movement and sperm cell delivery is largely unknown. This study reveals that WPP domain-interacting protein, a plant analog of animal proteins known to control nuclear movement, and WPP domain-interacting tail-anchored protein are responsible for movement of the vegetative nucleus within pollen tubes and contribute to sperm cell-to-ovule migration. This finding marks a leap forward in our understanding of double fertilization and the mechanism of nuclear movement in plants as a whole.
Abstract
Increasing evidence suggests that nuclear migration is important for eukaryotic development. Although nuclear migration is conserved in plants, its importance for plant development has not yet been established. The most extraordinary plant nuclear migration events involve plant fertilization, which is starkly different from that of animals. Instead of evolving self-propelled sperm cells (SCs), plants use pollen tubes to deliver SCs, in which the pollen vegetative nucleus (VN) and the SCs migrate as a unit toward the ovules, a fundamental but barely understood process. Here, we report that WPP domain-interacting proteins (WIPs) and their binding partners the WPP domain-interacting tail-anchored proteins (WITs) are essential for pollen nuclear migration. Loss-of-function mutations in WIT and/or WIP gene families resulted in impaired VN movement, inefficient SC delivery, and defects in pollen tube reception. WIPs are Klarsicht/ANC-1/Syne-1 Homology (KASH) analogs in plants. KASH proteins are key players in animal nuclear migration. Thus, this study not only reveals an important nuclear migration mechanism in plant fertilization but also, suggests that similar nuclear migration machinery is conserved between plants and animals.
Footnotes
- ↵1To whom correspondence should be addressed. Email: meier.56{at}osu.edu.
Author contributions: X.Z. and I.M. designed research; X.Z. performed research; X.Z. and I.M. analyzed data; and X.Z. and I.M. wrote the paper.
The authors declare no conflict of interest.
This article is a PNAS Direct Submission. U.G. is a guest editor invited by the Editorial Board.
This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323104111/-/DCSupplemental.