title: Stem cells in Nanomia bijuga (Siphonophora), a colonial animal with localized growth zones.
abstract: BACKGROUND
  Siphonophores (Hydrozoa) have unparalleled colony-level complexity, precision of colony organization, and functional specialization between zooids (i.e., the units that make up colonies). Previous work has shown that, unlike other colonial animals, most growth in siphonophores is restricted to one or two well-defined growth zones that are the sites of both elongation and zooid budding. It remained unknown, however, how this unique colony growth and development is realized at the cellular level.
  
  
  RESULTS
  To understand the colony-level growth and development of siphonophores at the cellular level, we characterize the distribution of proliferating cells and interstitial stem cells (i-cells) in the siphonophore Nanomia bijuga. Within the colony, we find evidence that i-cells are present at the tip of the horn, the structure within the growth zone that gives rise to new zooids. Co-localized gene expression of vasa-1, pl10, piwi, nanos-1, and nanos-2 suggests that i-cells persist in the youngest zooid buds and that i-cells become progressively restricted to specific regions within the zooids until they are mostly absent from the oldest zooids. The examined genes remain expressed in gametogenic regions. No evidence for i-cells is found in the stem between maturing zooids. Domains of high cell proliferation include regions where the examined genes are expressed, but also include some areas in which the examined genes were not expressed such as the stem within the growth zones. Cell proliferation in regions devoid of vasa-1, pl10, piwi, nanos-1, and nanos-2 expression indicates the presence of mitotically active epithelial cell lineages and, potentially, progenitor cell populations.
  
  
  CONCLUSIONS
  We provide the first evidence for i-cells in a siphonophore. Our findings suggest maintenance of i-cell populations at the sites of growth zones and that these sites are the main source of i-cells. This restriction of stem cells to particular regions in the colony, in combination with localized budding and spatial patterning during pro-bud subdivision, may play a major role in facilitating the precision of siphonophore growth. Spatially restricted maintenance of i-cells in mature zooids and absence of i-cells along the stem may explain the reduced developmental plasticity in older parts of the colony.
authors.affiliation: Department of Ecology and Evolutionary Biology, Brown University, 80 Waterman St. Box GW, Providence, RI 02912 USA.
authors.forename: Stefan
authors.initials: S
authors.lastname: Siebert
authors.affiliation: Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, District of Columbia, 20004 Washington USA.
authors.forename: Freya E
authors.initials: FE
authors.lastname: Goetz
authors.affiliation: Department of Ecology and Evolutionary Biology, Brown University, 80 Waterman St. Box GW, Providence, RI 02912 USA.
authors.forename: Samuel H
authors.initials: SH
authors.lastname: Church
authors.affiliation: Department of Ecology and Evolutionary Biology, Brown University, 80 Waterman St. Box GW, Providence, RI 02912 USA.
authors.forename: Pathikrit
authors.initials: P
authors.lastname: Bhattacharyya
authors.affiliation: Department of Ecology and Evolutionary Biology, Brown University, 80 Waterman St. Box GW, Providence, RI 02912 USA.
authors.forename: Felipe
authors.initials: F
authors.lastname: Zapata
authors.affiliation: Monterey Bay Aquarium Research Institute, Moss Landing, CA 95039 USA.
authors.forename: Steven H D
authors.initials: SH
authors.lastname: Haddock
authors.affiliation: Department of Ecology and Evolutionary Biology, Brown University, 80 Waterman St. Box GW, Providence, RI 02912 USA.
authors.forename: Casey W
authors.initials: CW
authors.lastname: Dunn
keywords: Growth zone; Interstitial stem cell; Nanomia bijuga; Siphonophora; i-cell