Novel features of Equisetum arvense spermatozoids:: insights into pteridophyte evolution

To characterize structural diversity within Equisetum and among pteridophytes, architectural features of the sperm cell are described here in a second subgenus of Equisetum, a divergent basal group in the fern clade. Transmission electron microscopy observations of prereleased spermatozoids of Equisetum arvense were correlated with three-dimensional scanning electron microscopy images of swimming cells. The mature spermatozoid completes a helix of approximately 2.5 revolutions. At the cell anterior is a complex multilayered locomotory apparatus with staggered flagella. Mitochondria (elongated-rounded) are aggregated near the locomotory apparatus and organelles extend along the cell length. The spline contains up to 300 microtubules and wraps in part around the long cylindrical nucleus. In swimming sperm cells, the anterior of the cell remains tightly coiled while the posterior relaxes and extends in a trailing fashion. Spermatozoids of Equisetum arvense are smaller than those of Equisetum hyemale but structurally similar, except for nuclear shape. Conservation of cellular features suggests recent radiation of the genus. Equisetum spermatozoids share several critical features with ferns, including Psilotum, and support monophyly of a fern-Equisetum assemblage. Entry of the male gametes of Equisetum in their entirety into the archegonial venters indicates possible biparental inheritance of chloroplast and mitochondrial genomes.