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Vitellogenesis and Post-Vitellogenic Maturation of the Insect Ovarian Follicle L Entomology Publications Entomology 2005 Vitellogenesis and Post-Vitellogenic Maturation of the Insect Ovarian Follicle L. Swevers National Centre for Scientific Research ‘‘Demokritos,’’ Athens, Greece A. S. Raikhel University of California, Riverside T. W. Sappington Iowa State University, [email protected] P. Shirk U.S. Department of Agriculture K. Latrou National Centre for Scientific Research ‘‘Demokritos,’’ Athens, Greece Follow this and additional works at: https://lib.dr.iastate.edu/ent_pubs Part of the Entomology Commons, Molecular Genetics Commons, and the Population Biology Commons The ompc lete bibliographic information for this item can be found at https://lib.dr.iastate.edu/ ent_pubs/482. For information on how to cite this item, please visit http://lib.dr.iastate.edu/ howtocite.html. This Book Chapter is brought to you for free and open access by the Entomology at Iowa State University Digital Repository. It has been accepted for inclusion in Entomology Publications by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. Vitellogenesis and Post-Vitellogenic Maturation of the Insect Ovarian Follicle Abstract Female insects typically produce prodigious numbers of eggs to assure the propagation of their genes, and invest considerable resources towards this end. Ulti- mately, the egg of an insect must contain a haploid set of chromosomes, sufficient nutrients to supply the growing embryo with resources to last until the larva or nymph ecloses and begins feeding, and a set of determinants to direct the organization and progres- sion of embryogenesis, including the differentiation of a new cluster of germ cells. As with all organs, the morphology of the ovary reflects the physical and genetic requirements of its physiological role, which in this case is the functional assembly of the various components of the oocyte. Visual inspection shows that the polytrophic ovary of holometabolus insects, which represent a major focus of this chapter, is comprised of a series of ovarioles that contain linear arrays of progres- sively developing follicles starting with dividing germ stem cells at one end and ending with mature oocytes ready for fertilization at the other ( Figure 1). Essentially, the ovariole can be considered an assem- bly line leading to the production of the egg. How this assembly line operates within different species to produce similar end products, i.e., the mature oocytes, depends on the insect, its life style, and its evolutionary history. Disciplines Ecology and Evolutionary Biology | Entomology | Genetics and Genomics | Molecular Genetics | Population Biology Comments This is a chapter from Swevers, L., A. S. Raikhel, T. W. Sappington, P. Shirk, and K. Iatrou. 2005. Vitellogenesis and post-vitellogenic maturation of the insect ovarian follicle. In: L. I. Gilbert, S. Gill, and K. Iatrou (eds.), . Vol. 1, Reproduction and Development. Elsevier, NY, pp. 87-155. doi: 10.1016/B0-44-451924-6/00093-4. Posted with permission. Rights Works produced by employees of the U.S. Government as part of their official duties are not copyrighted within the U.S. The onc tent of this document is not copyrighted. This book chapter is available at Iowa State University Digital Repository: https://lib.dr.iastate.edu/ent_pubs/482 1.3 Vitellogenesis and Post-Vitellogenic Maturation of the Insect Ovarian Follicle L Swevers, National Centre for Scientific Research ‘‘Demokritos,’’ Athens, Greece A S Raikhel, University of California, Riverside, CA, USA T W Sappington, Iowa State University, Ames, IA, USA PShirk, USDA ARS CMAVE, Gainesville, FL, USA K Iatrou, National Centre for Scientific Research ‘‘Demokritos,’’ Athens, Greece ß 2005, Elsevier BV. All Rights Reserved. 1.3.1. Introduction 87 1.3.2. Previtellogenesis: The Development of the Ovarian Structure 87 1.3.2.1. Drosophila melanogaster 87 1.3.2.2. Lepidoptera 91 1.3.3. Vitellogenesis 93 1.3.3.1. The Transition from Previtellogenesis to Vitellogenesis 93 1.3.3.2. Ovarian Yolk Proteins 101 1.3.3.3. The Transition from Vitellogenesis to Choriogenesis 113 1.3.3.4. Ecdysteroidogenesis and Resumption of Meiosis 120 1.3.3.5. Osmotic Swelling and Loss of Patency 121 1.3.3.6. Patterning and Movements of the Follicular Epithelium 121 1.3.3.7. Nurse Cell Dumping and Apoptosis 122 1.3.3.8. Vitelline Membrane Synthesis 123 1.3.4. Choriogenesis 125 1.3.4.1. Chorion Genes and Regulation of Chorion Gene Expression 127 1.3.5. Ovulation and Egg Activation 136 1.3.6. Conclusions and Perspectives 136 1.3.1. Introduction Essentially, the ovariole can be considered an assem- bly line leading to the production of the egg. How Female insects typically produce prodigious numbers this assembly line operates within different species of eggs to assure the propagation of their genes, and to produce similar end products, i.e., the mature invest considerable resources towards this end. Ulti- oocytes, depends on the insect, its life style, and its mately, the egg of an insect must contain a haploid evolutionary history. set of chromosomes, sufficient nutrients to supply the growing embryo with resources to last until the larva or nymph ecloses and begins feeding, and a set of 1.3.2. Previtellogenesis: The determinants to direct the organization and progres- Development of the Ovarian Structure sion of embryogenesis, including the differentiation of a new cluster of germ cells. As with all organs, the In this section, the development of the ovary and morphology of the ovary reflects the physical and oocyte in the fruit fly, Drosophila melanogaster, genetic requirements of its physiological role, which is examined briefly and then extend the discussion in this case is the functional assembly of the various to other insects, mainly Lepidoptera and non- components of the oocyte. Drosophila Diptera, where most of the additional Visual inspection shows that the polytrophic relevant information is available. ovary of holometabolus insects, which represent a 1.3.2.1. Drosophila melanogaster major focus of this chapter, is comprised of a series of ovarioles that contain linear arrays of progres- The organogenesis of the ovary and the production sively developing follicles starting with dividing of eggs in Drosophila is one of the most completely germ stem cells at one end and ending with mature examined developmental systems. From the forma- oocytes ready for fertilization at the other (Figure 1). tion of germ cells in the embryo to the localization 88 Vitellogenesis and Post-Vitellogenic Maturation of the Insect Ovarian Follicle Figure 1 Ovaries of holometaboulous insects. The upper panel shows ovaries dissected from previtellogenic (newly eclosed) and vitellogenic (four day old) Drosophila melanogaster adult females. The increase in size of the ovaries is due to the uptake of yolk proteins and other nutrients by the maturing follicles. The lower panel shows ovaries dissected from early vitellogenic (112 h) and prefollicular (white or 0 h) Plodia interpunctella female pupae. The increase in size is due to the growth of the germ and somatic tissues of the ovary. The linear array of developing follicles can be seen within each ovariole extending from the germarium to the lateral oviduct. The size of the ovary will increase again as the follicles complete vitellogenesis. of germ cell determinants in the oocytes within Raftery, 2000; Houston and King, 2000) (see the ovary of the adult female, the growth and activ- Chapter 1.2). ities of the ovary and follicles have been examined In Drosophila, the female molts into the adult extensively to determine the fundamental genetic stage with an immature ovary but, under the influ- and developmental processes that are involved in ence of juvenile hormone (JH), it initiates yolk the production of an egg. As a result, the structural protein production (vitellogenesis) and follicle organization of the ovary and the developmental maturation within the first day of adulthood processes controlling its formation have been stud- (reviews: Postlethwait and Shirk, 1981; Bownes, ied in-depth for over three-quarters of a century 1986; Lasko, 1994). The female continues producing (Dobzhansky, 1930; Kerkis, 1930; Demerec, 1950; eggs throughout its life, provided a sufficient supply King et al., 1968; King, 1970; Mahowald and of nutrients is available (Bownes, 1986). Because the Kambysellis, 1980; Spradling, 1993; Lasko, 1994; process of egg production is continuous, plentiful Mahajan-Niklos and Cooley, 1994; Dobens and experimental material is always at hand, providing Vitellogenesis and Post-Vitellogenic Maturation of the Insect Ovarian Follicle 89 a rich resource for the manipulation and analysis of germ cells (Hay et al., 1988; Lasko and Ashburner, oocyte production and maturation. 1988). Nearly half of the originally formed pole cells Under the influence of determinants localized are lost during embryogenesis, as they migrate from within the posterior pole plasm of the embryo, the the posterior of the germ band to their position germ cells are the first to cellularize in the syncytial within the embryonic gonads, and only 10–15 embryos (reviews: Spradling, 1993; Houston and germ cells are present within the late embryonic King, 2000) (see Chapter 1.2). The formation of gonad (Lasko and Ashburner, 1990; Mueller, the pole plasm within the oocyte is dependent 2002) (see Chapter 1.2). upon the maternal contribution and posterior locali- Throughout most of the larval stages, the germ zation of transcripts for oskar (osk; Lehman and cells divide mitotically to produce an unorganized Nu¨ sslein-Volhard, 1986), nanos (nos;Nu¨ sslein- mass of primordial germ cells mixed with somatic Volhard et al., 1987), and germ cell-less (gcl; cells (Kerkis, 1930). Midway through the third in- Jongens et al., 1992, 1994), as well as the proteins star, however, the metamorphosis of the ovary is Staufen (Stau; Schu¨ pbach and Wieschaus, 1986), initiated, leading to the organization of the somatic Valois (Vls; Schu¨ pbach and Wieschaus, 1986), Vasa and germ cells into the adult organ (King et al., 1968) (Vas; Schu¨ pbach and Wieschaus, 1986), Tudor (Tud; (see Chapter 1.2).
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