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Extraembryonic Development in Insects and the Acrobatics of Blastokinesis ⁎ Kristen A

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Extraembryonic Development in Insects and the Acrobatics of Blastokinesis ⁎ Kristen A View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Elsevier - Publisher Connector Available online at www.sciencedirect.com Developmental Biology 313 (2008) 471–491 www.elsevier.com/developmentalbiology Review Extraembryonic development in insects and the acrobatics of blastokinesis ⁎ Kristen A. Panfilio University Museum of Zoology, Department of Zoology, Downing Street, Cambridge CB2 3EJ, UK Received for publication 10 July 2007; revised 11 October 2007; accepted 2 November 2007 Available online 17 November 2007 Abstract Extraembryonic development is familiar to mouse researchers, but the term is largely unknown among insect developmental geneticists. This is not surprising, as the model system Drosophila melanogaster has an extremely reduced extraembryonic component, the amnioserosa. In contrast, most insects retain the ancestral complement of two distinct extraembryonic membranes, amnion and serosa. These membranes are involved in several key morphogenetic events at specific developmental stages. The events of anatrepsis and katatrepsis–collectively referred to as blastokinesis–are specific to hemimetabolous insects. Corresponding events in holometabolous insects are simplified and lack formal names. All insects retain dorsal closure, which has been well studied in Drosophila. This review aims to resurrect both the terminology and awareness of insect extraembryonic development–which were last common currency in the late nineteenth and early twentieth centuries–as a number of recent studies have identified essential components of these events, through RNA interference of developmental genes and ectopic hormonal treatments. As much remains unknown, this topic offers opportunities for research on tissue specification, the regulation of cell shape changes and tissue interactions during morphogenesis, tracing the origins and final fates of cell and tissue lineages, and ascertaining the membranes' functions between morphogenetic events. © 2007 Elsevier Inc. All rights reserved. Keywords: Insect; Extraembryonic membranes; Morphogenesis; Tissue interactions; Epithelial reorganization; Amnion; Serosa; Blastokinesis; Anatrepsis; Katatrepsis Introduction “additional complication” (p. 130) to embryogenesis in some insect orders. Yet extraembryonic development is relevant to Insect extraembryonic development encompasses all aspects the development of all insects. Researchers who are currently of embryogenesis involving the amniotic and serosal mem- familiar with this fact include embryological endocrinologists branes. In this review I will focus primarily on the role of the (e.g., Truman and Riddiford, 1999) and classical morpholo- membranes in morphogenetic events, specifically the move- gists (extraembryonic membranes even feature in the logo of ments associated with membrane formation–anatrepsis–and the Arthropodan Embryological Society of Japan). Meanwhile, their last acts–katatrepsis and dorsal closure–prior to their Anderson's view is still prevalent among developmental demise. These movements are particularly well developed in geneticists. Blastokinesis does not occur in the model hemimetabolous insects (those with incomplete metamorpho- organism Drosophila melanogaster, the embryos of which sis) and are collectively known as blastokinesis. Extraembryo- have very reduced extraembryonic tissue (although this tissue is nic membrane ontogeny in the holometabolous insects (those still essential for development). As a consequence, many with complete metamorphosis: embryonic, larval, pupal, and researchers are either unaware of the phenomena of extraem- adult stages) includes a subset of these events. bryonic development or have glossed over them in their own The occurrence of blastokinesis was last reviewed in detail research species. This is reflected in searching the PubMed 35 years ago by Anderson (1972a), who regarded it as an database of biomedical and life science articles (http://www. ncbi.nlm.nih.gov), where there are thousands of articles on “insect embryo” generally but only a handful specifically for ⁎ Fax: +44 1223 336 679. “blastokinesis” or “insect extraembryonic development.” As I E-mail address: [email protected]. hope to show here, extraembryonic development is fascinating 0012-1606/$ - see front matter © 2007 Elsevier Inc. All rights reserved. doi:10.1016/j.ydbio.2007.11.004 472 K.A. Panfilio / Developmental Biology 313 (2008) 471–491 and worthy of researchers' attention in its own right for study of progressively. Among non-insect hexapods (see phylogeny in a multitude of developmental processes. Fig. 2), Collembola and Protura (springtails and allies) possess a This review is two-pronged: it surveys manifestations of serosa and a structure known as the primary dorsal organ (see blastokinesis, and it summarizes relevant functional data on Box 1), but they lack two distinct membranes (Anderson, 1973, these movements. To provide adequate context, I begin with a p. 205; Jura, 1972; Machida, 2006; Uemiya and Ando, 1991). background account of the evolution and possible functional Similarly the Diplura (two-pronged bristletails) have a single, roles of the membranes. To introduce the nature and complexity serosal covering over the yolk, though there is also later of the morphogenetic events, blastokinesis is first described in production of additional extraembryonic tissue, which has been detail in a representative species. This is followed by the termed “amnion” (Ikeda and Machida, 2001). Within the true comparative survey of extraembryonic morphogenesis across insects, the amnion is prefigured by the pro-amniotic/pro- the insects. I then assess the relationship of the membranes to serosal distinction in archaeognathan (bristletail) embryos the execution of these movements in different species. Lastly, I (Heymons and Heymons, 1905; Machida et al., 1994; see present recent data from perturbation studies that elucidate some Box 1). An amnion proper is first seen in the Thysanura of the molecular components underlying extraembryonic (firebrats; Anderson, 1972a, pp. 204–205; Jura, 1972, pp. 80– morphogenesis, and identify possible avenues for further 82). However the thysanuran amnion does not form a complete research. membrane but leaves a persistently open amniotic cavity (Heymons and Heymons, 1905; Woodland, 1957). Complete Phylogeny of extraembryonic membrane elaboration and amnions seem restricted to the pterygote (winged) insects, reduction although the apterygote species survey supporting this claim is limited. Extraembryonic cells or tissues are common features of As Dorn (1976) also indicates, there are deviations from embryogenesis in animals. This material has become elaborated possession of two extraembryonic membranes in some in the insects to comprise two distinct membranes, with each holometabolous insect species (see phylogeny in Fig. 2). possessing a distinct topography and role. The inner membrane, Here, the lack of a full membrane complement is a derived, the amnion, envelops the ventral side of the developing embryo, secondary condition. Wholesale changes in embryogenesis– creating the fluid-filled amniotic cavity. The outer membrane, such as rapid development from a large embryonic rudiment the serosa, lies just under the chorion (eggshell), and surrounds (relative to egg size)–correlate with a reduction in extraem- embryo, amnion, and yolk (Fig. 1). Presence of these two bryonic tissue (Anderson, 1972a; Roth, 2004). In some lower membranes is regarded as a synapomorphy (shared, derived Hymenoptera (sawflies), the amnion forms but degenerates very character) of the insect egg (Larink, 1997): early in development and may be fragmentary (Ivanova-Kasas, 1959; Shafiq, 1954). Apocritan (higher) Hymenoptera (bees, Serosa and amnion can be considered standard inventory of ants, and wasps) usually lack an amnion, or have only a the insect embryo. The absence of one or the other in temporary amniotic vestige that covers the yolk rather than the apterygote insects is probably indicative of their phyloge- embryo (Anderson, 1972b; Bull, 1982; Fleig and Sander, 1988). netic development, whereas deviations, as they occur in Cyclorrhaphous (higher) Diptera (true flies), including the fruit holometabolous insects, represent presumably environmen- fly Drosophila melanogaster, lack distinct amniotic and serosal tal adjustments. (Dorn, 1976) membranes, retaining a single membrane, the amnioserosa, As implied by the quotation, the membranes arose within the which also covers only the yolk (Anderson, 1972b; Schmidt- lineage of apterygote (primitively wingless) insects, perhaps Ott, 2000). Proposed functions of the extraembryonic membranes Given the conserved presence of the extraembryonic mem- branes in most insects, why do they exist? Their designation as extraembryonic signifies that the amnion and the serosa do not directly contribute to the final form of the body. What function do they fulfill to necessitate their temporary existence? Such questions have produced several suggestions. Because Fig. 1. Schematic cartoons of a germband stage embryo, illustrating the the membranes surround the embryo, a general protective positions of the serosa (blue) and amnion (orange) with respect to the embryo (grey) and yolk (yellow). The amniotic cavity (white) is the region between the function has been ascribed (Dorn, 1976; Oseto and Helms, amnion and the ventral surface of the embryo. (a) Mid-sagittal section view. (b) 1972; Zeh et al., 1989). Specifically, the fluid-filled
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