title: Posterior localization of ApVas1 positions the preformed germ plasm in the sexual oviparous pea aphid Acyrthosiphon pisum.
abstract: BACKGROUND
  Germline specification in some animals is driven by the maternally inherited germ plasm during early embryogenesis (inheritance mode), whereas in others it is induced by signals from neighboring cells in mid or late development (induction mode). In the Metazoa, the induction mode appears as a more prevalent and ancestral condition; the inheritance mode is therefore derived. However, regarding germline specification in organisms with asexual and sexual reproduction it has not been clear whether both strategies are used, one for each reproductive phase, or if just one strategy is used for both phases. Previously we have demonstrated that specification of germ cells in the asexual viviparous pea aphid depends on a preformed germ plasm. In this study, we extended this work to investigate how germ cells were specified in the sexual oviparous embryos, aiming to understand whether or not developmental plasticity of germline specification exists in the pea aphid.
  
  
  RESULTS
  We employed Apvas1, a Drosophila vasa ortholog in the pea aphid, as a germline marker to examine whether germ plasm is preformed during oviparous development, as has already been seen in the viviparous embryos. During oogenesis, Apvas1 mRNA and ApVas1 protein were both evenly distributed. After fertilization, uniform expression of Apvas1 remained in the egg but posterior localization of ApVas1 occurred from the fifth nuclear cycle onward. Posterior co-localization of Apvas1/ApVas1 was first identified in the syncytial blastoderm undergoing cellularization, and later we could detect specific expression of Apvas1/ApVas1 in the morphologically identifiable germ cells of mature embryos. This suggests that Apvas1/ApVas1-positive cells are primordial germ cells and posterior localization of ApVas1 prior to cellularization positions the preformed germ plasm.
  
  
  CONCLUSIONS
  We conclude that both asexual and sexual pea aphids rely on the preformed germ plasm to specify germ cells and that developmental plasticity of germline specification, unlike axis patterning, occurs in neither of the two aphid reproductive phases. Consequently, the maternal inheritance mode implicated by a preformed germ plasm in the oviparous pea aphid becomes a non-canonical case in the Hemimetabola, where so far the zygotic induction mode prevails in most other studied insects.
authors.affiliation: Laboratory for Genetics and Development, Department of Entomology/Institute of Biotechnology, College of Bioresources and Agriculture, National Taiwan University, No. 27, Lane 113, Roosevelt Road, Sec. 4, Taipei 106, Taiwan ; Laboratory of Ecological Genetics, Graduate School of Environmental Science, Hokkaido University, N10 W5, Kita-ku, Sapporo, Hokkaido 060-0810, Japan.
authors.forename: Gee-Way
authors.initials: GW
authors.lastname: Lin
authors.affiliation: EMBL-European Bioinformatics Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge CB10 1SD, UK.
authors.forename: Charles E
authors.initials: CE
authors.lastname: Cook
authors.affiliation: Laboratory of Ecological Genetics, Graduate School of Environmental Science, Hokkaido University, N10 W5, Kita-ku, Sapporo, Hokkaido 060-0810, Japan.
authors.forename: Toru
authors.initials: T
authors.lastname: Miura
authors.affiliation: Laboratory for Genetics and Development, Department of Entomology/Institute of Biotechnology, College of Bioresources and Agriculture, National Taiwan University, No. 27, Lane 113, Roosevelt Road, Sec. 4, Taipei 106, Taiwan ; Research Center for Developmental Biology and Regenerative Medicine, National Taiwan University, Taipei 100, Taiwan ; Genome and Systems Biology Degree Program, National Taiwan University, Taipei 106, Taiwan.
authors.forename: Chun-Che
authors.initials: CC
authors.lastname: Chang
keywords: Aphid; Asymmetric localization; Developmental plasticity; Germ cells; Germline specification; Vasa