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Chromosome Visualization in a Temperate Anemone, Anthopleura Elegantissima

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Chromosome Visualization in a Temperate Anemone, Anthopleura Elegantissima Chromosome visualization in a temperate anemone, Anthopleura elegantissima by Lucy Shaffer A THESIS submitted to Oregon State University Honors College in partial fulfillment of the requirements for the degree of Honors Baccalaureate of Science in BioResource Research (Honors Scholar) Presented May 25, 2018 Commencement June 2018 AN ABSTRACT OF THE THESIS OF Lucy Shaffer for the degree of Honors Baccalaureate of Science in BioResource Research presented on May 25, 2018. Title: Chromosome visualization in a temperate anemone Anthopleura elegantissima. Abstract approved: _____________________________________________________ Eli Meyer Anthopleura elegantissima (Cnidaria: Anthozoa) is an emerging model organism for studying the endosymbiotic relationship between dinoflagellate (genus Symbiodinium) and the cnidarian host. To be developed as a model organism, more genetic resources, like integrated genome and linkage map, are needed. An important validation for developing genomic resources is chromosome number. To count chromosomes, we need to be able to visualize them. In this study, I developed protocols for single cell dissociation, tissue imprinting, fixation, and DAPI staining for Anthopleura elegantissima. I visualized chromosomes for this anemone and propose that these methods can be used to karyotype Anthopleura elegantissima, to further develop this model. Key Words: cell biology, cell visualization, chromosome visualization, anemone, Anthopleura elegantissima Corresponding e-mail address: [email protected] ©Copyright by Lucy Shaffer May 25, 2018 All Rights Reserved Chromosome visualization in a temperate anemone, Anthopleura elegantissima by Lucy Shaffer A THESIS submitted to Oregon State University Honors College in partial fulfillment of the requirements for the degree of Honors Baccalaureate of Science in BioResource Research (Honors Scholar) Presented May 25, 2018 Commencement June 2018 Honors Baccalaureate of Science in BioResource Research project of Lucy Shaffer presented on May 25, 2018. APPROVED: _____________________________________________________________________ Eli Meyer, Mentor, representing Department of Integrative Biology _____________________________________________________________________ Stephen Atkinson, Secondary Mentor, representing Department of Microbiology _____________________________________________________________________ Holland Elder, Committee Member, representing Department of Integrative Biology _____________________________________________________________________ Katherine Field, Committee Member, Director, BioResource Research _____________________________________________________________________ Toni Doolen, Dean, Oregon State University Honors College I understand that my project will become part of the permanent collection of Oregon State University, Honors College. My signature below authorizes release of my project to any reader upon request. _____________________________________________________________________ Lucy Shaffer, Author INTRODUCTION Coral reefs are estimated to harbor around one third of all described marine species (Wilkinson 2004). However, the rapid decline in coral abundance and loss of reef habitats are one of the most pressing environmental issues of our time. The coral reef depends upon a functional symbiosis between cnidarian hosts and unicellular, photosynthetic, dinoflagellate algae of the genus Symbiodinium (Baumgarten et al. 2015). The endosymbiotic relationship between the coral host and dinoflagellates is necessary for the survival of most reef building corals in the nutrient-poor seas of the tropics (Muscatine and Porter 1977). The endosymbiont provides 95% of the coral’s energy budget in the form of photosynthetically-fixed carbon - 3- (Trench 1993; Yellowlees et al. 2008). The host supplies nitrates (NO3 ) and phosphates (PO4 ) to sustain the endosymbiont’s productivity (Rädecker et al. 2018). The host and the endosymbiont live interdependently as they exchange nutrients. However, this relationship is disrupted by heat stress. Sea temperatures above the average maximum for a region cause corals to eject their symbionts in a process called bleaching (Hughes et al. 2017). Without their symbionts, the corals risk starvation and death. If temperatures do not return to the mean temperature for the region, there is often coral mortality (Hoegh-Guldberg et al. 2017). Despite the importance of coral dinoflagellate symbiosis, the specific molecular and cellular biological mechanisms of onset and the breakdown of this symbiosis are not well known. This is in part because of the difficulties of maintaining corals in the laboratory and experimental set-ups. Nearly half of all cnidarians contain symbiotic algae, including sea anemones, therefore other classes of Cnidaria can be used as an algal-symbiosis model (Douglas et al. 2009). Aiptasia pallida, a tropical sea anemone species, has been used as a model system to make advances in the field of cnidarian dinoflagellate symbiosis (Baumgarten et al. 2015). However, A. pallida. associates with multiple Symbiodinium species (LaJeunesse et al. 2004). This introduces additional complications to studying symbiosis in this system. Anthopleura elegantissima, commonly known as the aggregating anemone or clonal anemone, is a good candidate for a model of cnidarian dinoflagellates symbiosis, because it is symbiotic with a species of the dinoflagellate, Symbiodinium muscatinei, and with a single-celled alga, Elliptochloris marina (Lajeunesse and Trench 2000). The anemone is easy to maintain in a laboratory setting, and is the most abundant intertidal sea anemone on the Pacific Coast of North America (Quesada et al. 2016). Anthopleura elegantissima also exists naturally in three distinctly colored symbiotic states: zooxanthellate (brown,hosting the dinoflagellate S. muscatinei), zoochlorellate (green, hosting the chlorophyte Elliptochloris marina), and asymbiotic (white, lacking symbionts) (Lajeunesse and Trench 2000; Lewis and Muller-Parker 2004; Letsch et al. 2009). Having symbiotic relationships with fewer species and distinct symbiotic states will simplify experimental designs which can determine the mechanisms for of onset and breakdown of symbiosis. 1 Despite its potential as a model for symbiosis, there is a need for genomic resources for A. elegantissima, which are required to design complicated cell biology experiments targeting specific genes that play a role in mechanisms of symbiosis. A transcriptome assembly of functional genes has been published (Kitchen et al. 2015). An integrated genome and linkage map is in development by the Meyer lab at Oregon State University. The draft genome for the A. elegantissima is 310 Mb, with a linkage map that indicates that there are 16 to 18 linkage groups, which suggests that there are 32 to 36 chromosomes in the diploid organism (i.e 2n=32-36). We expected that A. elegantissima would have 32 to 36 chromosomes, both from the number of linkage groups, and from the karyotypes of other anemones and corals (Order: Anthozoa) which have been karyotyped have similar numbers of chromosomes. Karyotyped anemones include Haliplanella luciae with 2n=36 (Fukui, 1993) and Nematostella vectensis with 2n=30 (Guo et al. 2018). Corals that have been karyotyped include Goniopora lobata with 2n=14 (Heyward 1985), Favia pallida (2n=28), Galaxea fascicularis (2n=26), Acropora millepora (2n=28), Acropora spathulata (2n=28), Acropora papillare (2n=26) and Acropora nasuta (2n=40) (Flot et al. 2006). While a great diversity of chromosome numbers is represented here, two anemones and three coral species have similar numbers of chromosomes to the number of chromosomes for A. elegantissima indicated by the linkage map. My project was to support the conclusions of the linkage map, with a karyotype of the anemone to confirm the number of chromosomes present. Previous anemone and coral karyotyping studies used fertilized eggs or embryos because they are naturally in high mitotic states (Fukui 1993; Choe et al. 2000). Gamete collection is time consuming and limited due to A. elegantissima displaying annual cycle of sexual reproduction, only spawning in the fall. (Ford 1964). Mitotic karyotype from live polyp of hydra has been done before, which suggests that chromosome visualization can be done in adult sea anemone because both are in the classes of Cnidaria, and are similar in body structure including tissue layers (Anokhin and Nokkala 2004). However, karyotyping from adult anemone tissue have not been done before, therefore specific conditions under which the chromosomes can best be visualized had to be developed. This study followed the general karyotyping procedures for specimen preparation, chromosome preparation and chromosome staining (Anokhin et al. 2010). Specimen preparation included a single-cell dissociation protocol, tissue imprinting and DNA staining. Through these techniques, we visualized cell nuclei and chromosomes in A. elegantissima. These techniques could be applied to karyotype not only A. elegantissima but other species of sea anemones. MATERIALS AND METHODS Field sampling Anthopleura elegantissima were collected from Seal Rock Oregon from the rocky intertidal coast under Oregon Department of Fish and Wildlife (ODFW) scientific permit #18876. They were transported to Oregon State University in Corvallis, OR, USA and maintained in recirculating aquaria at 12˚C. 2 Single cell dissociation protocol To obtain single cells from live Anthopleura elegantissima specimens, a modified coral
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