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Blastodermal Cuticle Formation Contributes to Desiccation Resistance in Springtail Eggs: Eco-Evolutionary Implications for Insect Terrestrialization

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Blastodermal Cuticle Formation Contributes to Desiccation Resistance in Springtail Eggs: Eco-Evolutionary Implications for Insect Terrestrialization bioRxiv preprint doi: https://doi.org/10.1101/767947; this version posted September 12, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC 4.0 International license. Vargas et. al., Springtail eggs. This preprint has not been submitted to any journal. September 12th, 2019. Blastodermal cuticle formation contributes to desiccation resistance in springtail eggs: eco-evolutionary implications for insect terrestrialization Helena Carolina Martins Vargas1,2; Kristen A. Panfilio3; Dick Roelofs2; Gustavo Lazzaro Rezende1,3 ¹Laboratório de Química e Função de Proteínas e Peptídeos, Centro de Biociências e Biotecnologia, Universidade Estadual do Norte Fluminense Darcy Ribeiro, Campos dos Goytacazes, Brazil. ²Department of Ecological Science, Faculty of Science, Vrije Universiteit, De Boelelaan 1085, 1081, HV Amsterdam, The Netherlands. 3School of Life Sciences, University of Warwick, Gibbet Hill Campus, Coventry, CV4 7AL, UK. HCM: ORCID 0000-0001-8290-8423, [email protected] / KAP: ORCID 0000-0002-6417-251X, [email protected] DR: ORCID 0000-0003-3954-3590, [email protected] / GLR: ORCID 0000-0002-8904-7598, [email protected] /[email protected] Abstract Land colonization was a major event in the history of life. Among animals, insects had a staggering terrestrialization success, due to traits usually associated with post-embryonic life stages, while the egg stage has been largely overlooked in comparative studies. In many insects, after blastoderm differentiation, the extraembryonic serosal tissue wraps the embryo and synthesizes the serosal cuticle, an extracellular matrix that lies beneath the eggshell and protects the egg against water loss. In contrast, in non-insect hexapods such as springtails (Collembola) the early blastodermal cells synthesize a blastodermal cuticle. Here, we investigate the relationship between blastodermal cuticle formation and egg resistance to desiccation in the springtails Orchesella cincta and Folsomia candida, two species with different oviposition tenvironments and developmental rates. The blastodermal cuticle becomes externally visible in O. cincta and F. candida at 22 and 29% of embryogenesis, respectively. To contextualize, we describe the stages of springtail embryogenesis, exemplified by F. candida. Physiological assays showed that blastodermal cuticle formation supports egg viability in a dry environment, significantly contributing to hatching success. However, protection differs between species: while O. cincta eggs survive at least 2 hours outside a humid environment, the survival period recorded for F. candida eggs is only 15 minutes, which correlates with this species’ requirement for humid microhabitats. We suggest that the formation of this protective cuticle is an ancestral trait among hexapods, predating and facilitating the process of terrestrialization that occurred during insect evolution. Keywords: blastodermal cuticle, egg resistance to desiccation, embryogenesis, land colonization, springtails, water loss. Introduction insects accounts for 79% and 66% of the total, Life originated in a marine environment and respectively (Suppl. Figure 1). In other words, subsequent colonization of land was a significant event considering all living animal species, 2/3 are insects for the evolution of many organisms on Earth, such as (Zhang, 2011). arthropods (Little, 1983; Selden, 2016). The arthropod In order to survive on land, insects evolved phylum is comprised of the paraphyletic "Crustacea" traits that allow them to overcome low or irregular and the monophyletic Chelicerata, Myriapoda and water availability, such as an internalized respiratory Hexapoda (constituted of Collembola, Protura, Diplura system and an exoskeleton that minimizes water loss and Insecta) (Kukalová-Peck, 1987; Misof et al., 2014) (Dunlop et al., 2013; Hadley, 1994; Little, 1983). (Suppl. Figure 1). It is estimated that during arthropod Moreover, it was suggested that traits related to the evolution independent events of terrestrialization egg stage contributed to their successful occurred, standing out the ones of hexapods, terrestrialization, in addition to juvenile and adult myriapods and chelicerates (Dunlop et al., 2013; characteristics (Zeh et al., 1989). Freshly laid insect Grimaldi and Engel, 2005; Rota-Stabelli et al., 2013). eggs are covered by eggshell layers that are produced Specifically in the case of insect evolution, during oogenesis within the maternal body. These colonization of land led to an unparalleled maternal eggshell layers will be collectively termed diversification among animals. There are 1,527,660 here as 'chorion'. During the early stages of insect described species of extant animals; arthropods and embryogenesis, the blastoderm differentiates into three 1 bioRxiv preprint doi: https://doi.org/10.1101/767947; this version posted September 12, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC 4.0 International license. Vargas et. al., Springtail eggs. This preprint has not been submitted to any journal. September 12th, 2019. types of cells: the two extraembryonic tissues, serosa Orchesella cincta (Entomobryidae) is a litter- and amnion, and the embryonic ones. After dwelling (epidaphic) species. Adults are pigmented differentiation the serosal cells envelop the embryo and measure about 6 mm in length. Its embryogenesis and secrete the serosal cuticle, an extracellular matrix lasts for about 6 days at 19-24 ºC, while egg size at the (Handel et al., 2000; Panfilio, 2008; Rezende et al., chorion rupture stage is about 200 - 250 μm in 2016, 2008). Present in most insects, this post-zygotic diameter, when two blastodermal cuticles are cuticle constitutes the innermost eggshell layer, located produced. This cuticle is covered with thorns (Bretfeld, below the chorion. Moreover, it is fundamental to 1963). Its postembryonic development takes at least 44 protect the insect egg against water loss (Jacobs et al., days before reaching the adult stage (Hopkin, 1997; 2013; Rezende et al., 2016, 2008). The serosa and its Joosse et al., 1972). The species reproduces sexually cuticle are considered as crucial traits that fostered the through indirect sperm transfer via spermatophores. evolution of insects on land (Jacobs et al., 2013). The individuals molt during adult life, and sperm cells However, nothing is known about the role of post- cannot be stored by females. After each adult molt a zygotic cuticle in other lineages, given that female enters a new reproductive cycle and take up a comparative studies with other arthropods are still new spermatophore (Hopkin, 1997). Orchesella cincta lacking. is employed in ecotoxicology tests (Hopkin, 1997) and Indeed, some other arthropods produce post- its genome was recently sequenced (Faddeeva- zygotic eggshell layers named blastodermal cuticle Vakhrusheva et al., 2016). Also, several molecular (also termed blastodermic cuticle or blastoderm techniques were recently developed for this species, cuticle) (Machida, 2006; Rezende et al., 2016). For such as RNAi-driven gene silencing and in situ this trait, it is feasible to study Collembola, a hexapod hybridization in eggs (Konopova and Akam, 2014). group closely related to the insects, but lacking such a Folsomia candida (Isotomidae) is a tremendous diversification: collembolans, colloquially parthenogenetic, soil-dwelling (eudaphic) species. called 'springtails', account for only 0.5% of the total Adults are unpigmented, blind and measure between 1 species number of living animals (Suppl. Figure 1). to 3 mm in length. Its embryogenesis lasts about 10 Collembolan embryogenesis and blastodermal cuticle days and hatchlings take 24 days to reach the adult formation have been described in many species stage at 20 ºC. Its eggs are smaller than those of O. (Bretfeld, 1963; Jura, 1972; Jura et al., 1987; Marshall cincta: they measure about 80 μm when laid and and Kevan, 1962; Prowazek, 1900; Tamarelle, 1981; increase in size up to 165 μm at the chorion rupture Tomizuka and Machida, 2015; Uemiya and Ando, stage (Marshall and Kevan, 1962). Adults can survive 1987a, 1987b) (Figure 1). Briefly, during early under relatively dry conditions by increasing embryogenesis the springtail blastoderm differentiates osmolality through production of sugars and polyols, into two types of cells: an extraembryonic tissue and facilitating absorbance of water vapor (Fountain and the embryonic rudiment (Figure 1A, B). Depending on Hopkin, 2005; Hopkin, 1997; Roelofs et al., 2013). For the species the blastodermal cuticle is secreted by cells more than 40 years, F. candida has been used as a of either the uniform undifferentiated blastoderm or model organism for the study of soil contamination the differentiated blastoderm (Figure 1C). In parallel, and is used in an internationally standardized test (ISO water uptake increases egg volume, which leads to 11267) to assess the level of toxicity of contaminated chorion rupture (Figure 1D), thus forming the polar soils and to monitor their remediation (Fountain and caps (i.e. chorion remnants, which lie at both poles of Hopkin, 2005). The F. candida genome was recently the egg) and the blastodermal cuticle becomes the sole sequenced (Faddeeva-Vakhrusheva
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