DOCSLIB.ORG

Evidence for Gonadotropin-Releasing Hormone-Like Peptides in a Cnidarian Nervous System

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
Evidence for Gonadotropin-Releasing Hormone-Like Peptides in a Cnidarian Nervous System General and Comparative Endocrinology 119, 317–328 (2000) doi:10.1006/gcen.2000.7524, available online at http://www.idealibrary.com on Evidence for Gonadotropin-Releasing Hormone-like Peptides in a Cnidarian Nervous System Michel Anctil De´partement de Sciences Biologiques and Centre de Recherche en Sciences Neurologiques, Universite´ de Montre´al, Montre´al, Que´bec, Canada H3C 3J7; and Whitney Laboratory, University of Florida, St. Augustine, Florida 32086 Accepted May 25, 2000 There is increasing evidence that peptides of the gonado- resented in invertebrates by native forms (De Loof et tropin-releasing hormone (GnRH) family, long consid- al., 1989), suggesting that peptide families are con- ered a vertebrate preserve, are also present in inverte- served through evolutionary lineages from inverte- brate (molluscan) nervous systems. The possibility was brates to vertebrates. This conservation may also exist examined that GnRHs are present and bioactive in cni- in the lowest phylum (Cnidaria) in which neurons are darians, considered to be representatives of the most identifiable. For example, neurons of several cnidarian primitive animals possessing a nervous system. Immuno- species contain peptides belonging to the FMRFamide reactive GnRH was detected in endodermal neurons of family (Grimmelikhuijzen et al., 1992; Grimmelikhui- two anthozoans, the sea pansy Renilla koellikeri and the jzen and Westfall, 1995), all but one (Yasuda et al., sea anemone Nematostella vectensis. In the sea pansy, 1993) unique to this phylum. immunoreactivity was detected throughout the auto- The gonadotropin-releasing hormones (GnRHs) zooid polyps, including gamete-producing endoderm. constitute a peptide family whose members are re- High-performance liquid chromatography and radioim- markably conserved in length and composition of munoassays of extracts from whole sea pansy colonies amino acid sequences and appear to have a distribu- yielded two elution peaks exhibiting GnRH immunoreac- tion restricted to vertebrates (Sherwood, 1987). The tivity with antisera raised against shark or mammalian possibility that GnRHs are also represented in inver- GnRH. Vertebrate GnRHs as well as the two sea pansy tebrates is supported by evidence showing that GnRH-like factors inhibited the amplitude and frequency GnRH-like immunoreactive material is present in neu- of peristaltic contractions in the sea pansy, and these rons of the gastropod mollusc Helisoma trivolvis and 1 actions were blocked by the LHRH analog [D-pGlu ,D- that extracts from its nervous system induced electro- 2 3,6 Phe ,D-Trp ]-LHRH. These results suggest that the physiological responses in Helisoma neurons and go- GnRH family of neuropeptides is more widespread in nadotropin release from dispersed goldfish pituitary metazoans than previously thought. Although our physi- cells (Goldberg et al., 1993). In addition, GnRH-like ological data are preliminary, they point to a role for immunoreactive material was detected in several GnRHs as inhibitory modulators of neuromuscular trans- protochordates (Georges and Dubois, 1980; Dufour mission in the sea pansy. © 2000 Academic Press et al., 1988; Kelsall et al., 1990; Mackie, 1995; Cameron Key Words: Cnidaria; GnRH; neuropeptide; sea pansy; et al., 1999) and two new peptides of the GnRH molecular evolution. family were identified in the tunicate Chelyosoma pro- ductum (Powell et al., 1996), thus demonstrating that Several of the families of structurally related neu- vertebrate GnRHs had ancestors in the protochordate ropeptides first identified in vertebrates are also rep- lineages. 0016-6480/00 $35.00 317 Copyright © 2000 by Academic Press All rights of reproduction in any form reserved. 318 Michel Anctil This paper addresses whether GnRH-like peptides diluted (30%) local sea water at the Whitney Laboratory. occur in invertebrates other than molluscs and tuni- They were fed three times weekly with brine shrimps, at cates and, focusing on cnidarians, whether the evolu- which times their culture medium was changed. tionary history of the GnRH family is traceable to the Immunohistochemistry. Ten sea pansy colonies earliest invertebrates known to possess a nervous sys- were prepared for whole-mount immunohistochemis- tem. The primary cnidarian used to investigate these try. Sea pansies were anesthetized in equal parts of questions was the sea pansy, Renilla koellikeri, an octo- 0.37 M MgCl2 and ASW. Autozooid polyps and their corallian belonging to the sea pen group (Pennatula- recessed reproductive tissue inside the colonial mass cea). The organization of the pennatulacean nervous (rachis) were excised and immersed in Zamboni’s fix- system is reasonably well known (Buisson, 1970; Sat- ative (4% paraformaldehyde and 7.5% saturated picric terlie et al., 1980), and the sea pansy was the focus of acid) freshly prepared in 0.1 M phosphate buffer with extensive mapping of neurotransmitter-specific neu- 2.4% NaCl (PBS) at pH 7.4. Fixation proceeded at rons (Umbriaco et al., 1990; Pani et al., 1995; Anctil and room temperature for 1 h and at 4° for the following Ngo Minh, 1997; Mechawar and Anctil, 1997). In ad- 24 h. After washing in PBS, tissues were incubated for dition to classical neurotransmitters, the sea pansy 48 h at 4° in either of three different primary antisera: contains large amounts of the peptide Antho-RFamide an antiserum raised in rabbit against chicken GnRH-II which is widely distributed in its nervous system and graciously provided by Dr. J. P. Chang, University (Grimmelikhuijzen and Groeger, 1987). of Alberta (Goldberg et al., 1993) and two other anti- A combination of immunohistochemical and chro- sera raised in rabbit against LHRH, one from a com- matographic techniques was used to gain evidence for mercial source (Sigma Chemicals) and the other ob- the presence of GnRH-like peptides in the sea pansy. tained as a courtesy from Dr. Robert Benoıˆt (LR-1: In addition, the physiological function of GnRHs was Benoıˆt et al., 1987). All three antisera were diluted investigated by comparing the activity of synthetic 1:1000 in PBS with 1% normal goat serum or 1% GnRHs with chromatographically purified sea pansy bovine serum albumin and 0.3% Triton X-100. After GnRH-like fractions on sea pansy muscle activity. Fi- further washing in PBS, tissues were treated for 1 h nally, confirmatory immunohistochemical data were with Cy3-conjugated goat anti-rabbit IgG (Jackson Im- obtained in another anthozoan species, the starlet sea munochemicals) diluted 1:100 in PBS containing 0.3% anemone Nematostella vectensis. Together, the results Triton X-100. After a final wash, tissues were mounted from these approaches provide strong evidence for the in a 1:3 mixture of glycerol and PBS and examined presence and physiological relevance of GnRHs in with a Wild-Leitz Dialux 20 fluorescence microscope. Cnidaria. Preliminary results of this study were pre- Controls were performed by replacing the primary sented to the XIIIth International Congress of Com- antisera with nonimmunized serum and by preab- parative Endocrinology (Anctil, 1997). sorption of the primary antisera with 0.1 mM LHRH or chicken GnRH-II. The double-labeling immunohistochemistry proce- MATERIALS AND METHODS dure of Wu¨ rden and Homberg (1993) was used to determine whether FMRFamide-like immunoreactiv- ity (Fair), reflecting the presence of native Antho-RF- Colonies of sea pansy (Renilla koellikeri Pfeffer) were amide, colocalizes with GnRH-like immunoreactivity purchased from Marinus Inc. (Long Beach, CA) and (GnRHir) in the same cells. Tissues were treated for maintained in aquaria with recirculating, filtered arti- 48 h in the LR-1 LHRH antiserum diluted 1:1000 fol- ficial sea water (ASW, Instant Ocean) at 12–16° and pH lowed by the Cy3-conjugated secondary antiserum as 8.0. The unfed colonies were exposed to a light–dark previously described. After washing in three 1-h cycle according to the photoperiod length experienced changes of rabbit IgG (Jackson Immunochemicals) di- in their environment before shipment and were pro- luted 1:25 in PBS, the tissues were treated for 24 h at 4° cessed within 2 weeks of shipment. in a biotinylated FMRFamide antiserum (also raised in Starlet sea anemones (Nematostella vectensis) were rabbit, Incstar), diluted 1:800 in PBS. The FMRFamide maintained in culture dishes at room temperature in antiserum was biotinylated by the succinimide ester Copyright © 2000 by Academic Press All rights of reproduction in any form reserved. GnRH-like Hormones in a Cnidarian 319 method of Harlow and Lane (1986). Tissues were then noassay (RIA) in some of the fractions, more extract incubated for3hinstreptavidin-conjugated fluores- was subsequently processed by HPLC to select those cein isothiocyanate (FITC, Jackson Immunochemicals) specific fractions and pool them for further purifica- diluted 1:20. tion steps. Starlet sea anemones were processed similarly to sea The selected fractions were run as above but on a pansies except that anesthesia was performed in equal smaller column (Brownlee RP-300, 2.1 ϫ 220 mm) at mixtures of 30% sea water and 0.12 M MgCl2, and PBS 0.5 ml/min, and a stronger ion-pairing agent, hepta- was prepared with 0.8% NaCl to reflect the salinity con- fluoro-butyric acid (HFBA) was substituted for TFA at ditions under which the animals were cultured. A dilu- the same dilution. Three successive purification steps tion of 1:500 of the primary LHRH antiserum (Sigma) were performed with this procedure. was used, and either Cy3 (1:100) or FITC (1:80) was used Radioimmunoassay. Iodination and RIA protocols as fluorophore conjugated to secondary antibodies. followed Price et al. (1987). Two iodinated GnRH pep- Peptide extraction. Whole sea pansies were ex- tides, mammalian GnRH and salmon GnRH, were used tracted during the reproductive season (May–August) in the RIA methods tested. Iodination was performed by by the method of Goldberg et al. (1993) after modifi- mixing 2–3 ␮lof125I (1–1.5 mCi) with 1 ␮l of peptide cations. Sea pansies were deflated to remove residual solution (1 mM in 0.5 M phosphate buffer, pH 7.0), sea water from their gastrovascular channels, followed by addition of 5 ␮l of 0.2% chloramine T and weighed, and immediately frozen in liquid nitrogen.
Load more

Recommended publications
  • The Green Fluorescent Protein
    P1: rpk/plb P2: rpk April 30, 1998 11:6 Annual Reviews AR057-17 Annu. Rev. Biochem. 1998. 67:509–44 Copyright c 1998 by Annual Reviews. All rights reserved THE GREEN FLUORESCENT PROTEIN Roger Y. Tsien Howard Hughes Medical Institute; University of California, San Diego; La Jolla, CA 92093-0647 KEY WORDS: Aequorea, mutants, chromophore, bioluminescence, GFP ABSTRACT In just three years, the green fluorescent protein (GFP) from the jellyfish Aequorea victoria has vaulted from obscurity to become one of the most widely studied and exploited proteins in biochemistry and cell biology. Its amazing ability to generate a highly visible, efficiently emitting internal fluorophore is both intrin- sically fascinating and tremendously valuable. High-resolution crystal structures of GFP offer unprecedented opportunities to understand and manipulate the rela- tion between protein structure and spectroscopic function. GFP has become well established as a marker of gene expression and protein targeting in intact cells and organisms. Mutagenesis and engineering of GFP into chimeric proteins are opening new vistas in physiological indicators, biosensors, and photochemical memories. CONTENTS NATURAL AND SCIENTIFIC HISTORY OF GFP .................................510 Discovery and Major Milestones .............................................510 Occurrence, Relation to Bioluminescence, and Comparison with Other Fluorescent Proteins .....................................511 PRIMARY, SECONDARY, TERTIARY, AND QUATERNARY STRUCTURE ...........512 Primary Sequence from
    [Show full text]
  • Coastal and Marine Ecological Classification Standard (2012)
    FGDC-STD-018-2012 Coastal and Marine Ecological Classification Standard Marine and Coastal Spatial Data Subcommittee Federal Geographic Data Committee June, 2012 Federal Geographic Data Committee FGDC-STD-018-2012 Coastal and Marine Ecological Classification Standard, June 2012 ______________________________________________________________________________________ CONTENTS PAGE 1. Introduction ..................................................................................................................... 1 1.1 Objectives ................................................................................................................ 1 1.2 Need ......................................................................................................................... 2 1.3 Scope ........................................................................................................................ 2 1.4 Application ............................................................................................................... 3 1.5 Relationship to Previous FGDC Standards .............................................................. 4 1.6 Development Procedures ......................................................................................... 5 1.7 Guiding Principles ................................................................................................... 7 1.7.1 Build a Scientifically Sound Ecological Classification .................................... 7 1.7.2 Meet the Needs of a Wide Range of Users ......................................................
    [Show full text]
  • Bioluminescence and Fluorescence of Three Sea Pens in the North-West
    bioRxiv preprint doi: https://doi.org/10.1101/2020.12.08.416396; this version posted December 9, 2020. 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-ND 4.0 International license. Bioluminescence and fluorescence of three sea pens in the north-west Mediterranean sea Warren R Francis* 1, Ana¨ısSire de Vilar 1 1: Dept of Biology, University of Southern Denmark, Odense, Denmark Corresponding author: [email protected] Abstract Bioluminescence of Mediterranean sea pens has been known for a long time, but basic parameters such as the emission spectra are unknown. Here we examined bioluminescence in three species of Pennatulacea, Pennatula rubra, Pteroeides griseum, and Veretillum cynomorium. Following dark adaptation, all three species could easily be stimulated to produce green light. All species were also fluorescent, with bioluminescence being produced at the same sites as the fluorescence. The shape of the fluorescence spectra indicates the presence of a GFP closely associated with light production, as seen in Renilla. Our videos show that light proceeds as waves along the colony from the point of stimulation for all three species, as observed in many other octocorals. Features of their bioluminescence are strongly suggestive of a \burglar alarm" function. Introduction Bioluminescence is the production of light by living organisms, and is extremely common in the marine environment [Haddock et al., 2010, Martini et al., 2019]. Within the phylum Cnidaria, biolumiescence is widely observed among the Medusazoa (true jellyfish and kin), but also among the Octocorallia, and especially the Pennatulacea (sea pens).
    [Show full text]
  • Appendix C - Invertebrate Population Attributes
    APPENDIX C - INVERTEBRATE POPULATION ATTRIBUTES C1. Taxonomic list of megabenthic invertebrate species collected C2. Percent area of megabenthic invertebrate species by subpopulation C3. Abundance of megabenthic invertebrate species by subpopulation C4. Biomass of megabenthic invertebrate species by subpopulation C- 1 C1. Taxonomic list of megabenthic invertebrate species collected on the southern California shelf and upper slope at depths of 2-476m, July-October 2003. Taxon/Species Author Common Name PORIFERA CALCEREA --SCYCETTIDA Amphoriscidae Leucilla nuttingi (Urban 1902) urn sponge HEXACTINELLIDA --HEXACTINOSA Aphrocallistidae Aphrocallistes vastus Schulze 1887 cloud sponge DEMOSPONGIAE Porifera sp SD2 "sponge" Porifera sp SD4 "sponge" Porifera sp SD5 "sponge" Porifera sp SD15 "sponge" Porifera sp SD16 "sponge" --SPIROPHORIDA Tetillidae Tetilla arb de Laubenfels 1930 gray puffball sponge --HADROMERIDA Suberitidae Suberites suberea (Johnson 1842) hermitcrab sponge Tethyidae Tethya californiana (= aurantium ) de Laubenfels 1932 orange ball sponge CNIDARIA HYDROZOA --ATHECATAE Tubulariidae Tubularia crocea (L. Agassiz 1862) pink-mouth hydroid --THECATAE Aglaopheniidae Aglaophenia sp "hydroid" Plumulariidae Plumularia sp "seabristle" Sertulariidae Abietinaria sp "hydroid" --SIPHONOPHORA Rhodaliidae Dromalia alexandri Bigelow 1911 sea dandelion ANTHOZOA --ALCYONACEA Clavulariidae Telesto californica Kükenthal 1913 "soft coral" Telesto nuttingi Kükenthal 1913 "anemone" Gorgoniidae Adelogorgia phyllosclera Bayer 1958 orange gorgonian Eugorgia
    [Show full text]
  • The Diet and Predator-Prey Relationships of the Sea Star Pycnopodia Helianthoides (Brandt) from a Central California Kelp Forest
    THE DIET AND PREDATOR-PREY RELATIONSHIPS OF THE SEA STAR PYCNOPODIA HELIANTHOIDES (BRANDT) FROM A CENTRAL CALIFORNIA KELP FOREST A Thesis Presented to The Faculty of Moss Landing Marine Laboratories San Jose State University In Partial Fulfillment of the Requirements for the Degree Master of Arts by Timothy John Herrlinger December 1983 TABLE OF CONTENTS Acknowledgments iv Abstract vi List of Tables viii List of Figures ix INTRODUCTION 1 MATERIALS AND METHODS Site Description 4 Diet 5 Prey Densities and Defensive Responses 8 Prey-Size Selection 9 Prey Handling Times 9 Prey Adhesion 9 Tethering of Calliostoma ligatum 10 Microhabitat Distribution of Prey 12 OBSERVATIONS AND RESULTS Diet 14 Prey Densities 16 Prey Defensive Responses 17 Prey-Size Selection 18 Prey Handling Times 18 Prey Adhesion 19 Tethering of Calliostoma ligatum 19 Microhabitat Distribution of Prey 20 DISCUSSION Diet 21 Prey Densities 24 Prey Defensive Responses 25 Prey-Size Selection 27 Prey Handling Times 27 Prey Adhesion 28 Tethering of Calliostoma ligatum and Prey Refugia 29 Microhabitat Distribution of Prey 32 Chemoreception vs. a Chemotactile Response 36 Foraging Strategy 38 LITERATURE CITED 41 TABLES 48 FIGURES 56 iii ACKNOWLEDGMENTS My span at Moss Landing Marine Laboratories has been a wonderful experience. So many people have contributed in one way or another to the outcome. My diving buddies perse- vered through a lot and I cherish our camaraderie: Todd Anderson, Joel Thompson, Allan Fukuyama, Val Breda, John Heine, Mike Denega, Bruce Welden, Becky Herrlinger, Al Solonsky, Ellen Faurot, Gilbert Van Dykhuizen, Ralph Larson, Guy Hoelzer, Mickey Singer, and Jerry Kashiwada. Kevin Lohman and Richard Reaves spent many hours repairing com­ puter programs for me.
    [Show full text]
  • The Sandy Beach Environment
    THE SANDY BEACH ENVIRONMENT The sandy beach is a region of shifting sands and crashing waves. Each time a wave pummels the shore, sediment is suspended and scattered in every direction. Together the wind and waves constantly rework and reshape the face of the beach. Throughout the year, the beach slope is continually changing. During the spring and summer, gentle constructive waves deposit sediment on the beach platform, building up the slope. The large destructive waves of fall and winter strip the sand from the beach, often leaving only cobblestones and the rocky beach platform. This unstable environment makes it difficult for plants and animals to settle. In addition to waves, the organisms must contend with the problems imposed by tidal fluctuations. The daily ebb and flow of the sea exposes the shore to all the elements. The animals living within the range of the high and low tides (intertidal) are subjected to extreme temperature changes, drying out, and other problems resulting from exposure to the air. Very few animals can survive these rigorous conditions, and no large plants can anchor in the shifting sands. To the casual observer, the sandy beach appears desolate and barren. However, within the sand grains live large numbers of diverse animals. To survive in this environment, these animals have evolved some interesting adaptations. Successful species either ride the waves, live just above the tideline, or burrow beneath the sand to protect themselves from the hammering waves. The upper beach -is full of small scavengers and beachhoppers. These animals live just above the breaking waves.
    [Show full text]
  • An Invitation to Monitor Georgia's Coastal Wetlands
    An Invitation to Monitor Georgia’s Coastal Wetlands www.shellfish.uga.edu By Mary Sweeney-Reeves, Dr. Alan Power, & Ellie Covington First Printing 2003, Second Printing 2006, Copyright University of Georgia “This book was prepared by Mary Sweeney-Reeves, Dr. Alan Power, and Ellie Covington under an award from the Office of Ocean and Coastal Resource Management, National Oceanic and Atmospheric Administration. The statements, findings, conclusions, and recommendations are those of the authors and do not necessarily reflect the views of OCRM and NOAA.” 2 Acknowledgements Funding for the development of the Coastal Georgia Adopt-A-Wetland Program was provided by a NOAA Coastal Incentive Grant, awarded under the Georgia Department of Natural Resources Coastal Zone Management Program (UGA Grant # 27 31 RE 337130). The Coastal Georgia Adopt-A-Wetland Program owes much of its success to the support, experience, and contributions of the following individuals: Dr. Randal Walker, Marie Scoggins, Dodie Thompson, Edith Schmidt, John Crawford, Dr. Mare Timmons, Marcy Mitchell, Pete Schlein, Sue Finkle, Jenny Makosky, Natasha Wampler, Molly Russell, Rebecca Green, and Jeanette Henderson (University of Georgia Marine Extension Service); Courtney Power (Chatham County Savannah Metropolitan Planning Commission); Dr. Joe Richardson (Savannah State University); Dr. Chandra Franklin (Savannah State University); Dr. Dionne Hoskins (NOAA); Dr. Charles Belin (Armstrong Atlantic University); Dr. Merryl Alber (University of Georgia); (Dr. Mac Rawson (Georgia Sea Grant College Program); Harold Harbert, Kim Morris-Zarneke, and Michele Droszcz (Georgia Adopt-A-Stream); Dorset Hurley and Aimee Gaddis (Sapelo Island National Estuarine Research Reserve); Dr. Charra Sweeney-Reeves (All About Pets); Captain Judy Helmey (Miss Judy Charters); Jan Mackinnon and Jill Huntington (Georgia Department of Natural Resources).
    [Show full text]
  • Retinoic Acid and Nitric Oxide Promote Cell Proliferation and Differentially Induce Neuronal Differentiation in Vitro in the Cnidarian Renilla Koellikeri
    Retinoic Acid and Nitric Oxide Promote Cell Proliferation and Differentially Induce Neuronal Differentiation In Vitro in the Cnidarian Renilla koellikeri Djoyce Estephane, Michel Anctil De´ partement de sciences biologiques and Centre de recherche en sciences neurologiques, Universite´ de Montre´ al, Case postale 6128, Succursale Centre-ville, Montre´ al, Que´ bec, Canada, H3C 3J7 Received 26 January 2010; revised 8 July 2010; accepted 9 July 2010 ABSTRACT: Retinoic acid (RA) and nitric oxide cell density. NO donors also induce cell proliferation in (NO) are known to promote neuronal development in polylysine-coated dishes, but induce neuronal differen- both vertebrates and invertebrates. Retinoic acid tiation and neurite outgrowth in uncoated dishes. No receptors appear to be present in cnidarians and NO other cell type undergoes differentiation in the pres- plays various physiological roles in several cnidarians, ence of NO. These observations suggest that in the sea but there is as yet no evidence that these agents have a pansy (1) cell adhesion promotes proliferation without role in neural development in this basal metazoan phy- morphogenesis and this proliferation is modulated pos- lum. We used primary cultures of cells from the sea itively by 9-cis RA and NO, (2) 9-cis RA and NO differ- pansy Renilla koellikeri to investigate the involvement entially induce neuronal differentiation in nonadherent of these signaling molecules in cnidarian cell differen- cells while repressing proliferation, and (3) the involve- tiation. We found that 9-cis RA induce cell prolifera- ment of RA and NO in neuronal differentiation tion in dose- and time-dependent manners in dishes appeared early during the evolutionary emergence of coated with polylysine from the onset of culture.
    [Show full text]
  • CEAR Doc#961)
    Vancouver Fraser Port Auihority PORT of i00 The Fcinie, 999 Canada Place ('Ð Varrcouyer. B.C. Canacia V6C 3T4 vancouver portvancouver.corn February 9,2018 Jocelyne Beaudet Panel Chair, Roberts Bank Terminal 2 Project C/O Debra Myles Panel Manager, Roberts Bank Terminal 2 Project Canadian Environmental Assessment Agency 22nd Floor, Place Bell 160 Elgin Street Ottawa, ON K1A 0H3 Dear Mme. Beaudet, From the Vancouver Fraser Port Authority re: Information Requests from the Review Panel for the Roberts Bank Terminal 2 Project Environmental Assessment: Responses (Select Responses from Packages 2 and 5 - February 9, 2018 Submission) The Vancouver Fraser Port Authority is pleased to submit to the Review Panel selected responses to Information Request Packages 2 and 5 related to the Roberts Bank Terminal 2 Project Environmental Impact Statement. We are making available the document Information Requests from the Review Panel for the Roberts Bank TermÌnal 2 Project Environmental Assessment: Responses (Se/ect Responses from Packages 2 and 5 - February 9,2078 Submission) which addresses the information requests IR2-19a, IR5-05, -06, -07, -1O, -45, and -47. Compilation of Panel Information Requests and Vancouver Fraser Port Authority Responses, which combines all available responses to the Review Panel's Information Requests, will be updated shortly While we will continue to regularly send responses as they are completed, we anticipate to complete Information Request Package 5 by March, Package 6 by April, and PackageT by June, and the Revised Project Description by May 2018. Submission of responses to Packages 8, 9 and 10 are still being planned and will be the subject of a further update to the Panel in the future.
    [Show full text]
  • Cladistic Analysis of the Pennatulacean Genus <I>Renilla</I
    View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by UNL | Libraries University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Papers in Entomology Museum, University of Nebraska State January 2001 Cladistic analysis of the pennatulacean genus Renilla Lamarck, 1816 (Coelenterata, Octocorallia) Carlos D. Pérez Laboratorio de Biología de Cnidarios (LABIC), Departamento de Ciencias Marinas, Facultad de Ciencias Exactas y Naturales, UNMdP, Funes 3250, 7600, Mar del Plata, Argentina Federico C. Ocampo University of Nebraska - Lincoln, [email protected] Follow this and additional works at: https://digitalcommons.unl.edu/entomologypapers Part of the Entomology Commons Pérez, Carlos D. and Ocampo, Federico C., "Cladistic analysis of the pennatulacean genus Renilla Lamarck, 1816 (Coelenterata, Octocorallia)" (2001). Papers in Entomology. 126. https://digitalcommons.unl.edu/entomologypapers/126 This Article is brought to you for free and open access by the Museum, University of Nebraska State at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Papers in Entomology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Published in Journal of Natural History 35:2 (January 2001), pp. 169–173; doi 10.1080/00222930150215305 Copyright © 2001 Taylor & Francis Ltd. Used by permission. http://www.tandf.co.uk/journals http://dx.doi.org/10.1080/00222930150215305 Accepted December 7, 1999. Cladistic analysis of the pennatulacean genus
    [Show full text]
  • The Global Diversity of Sea Pens (Cnidaria: Octocorallia: Pennatulacea)
    Review The Global Diversity of Sea Pens (Cnidaria: Octocorallia: Pennatulacea) Gary C. Williams* California Academy of Sciences, San Francisco, California, United States of America most brilliantly phosphorescent …. In some place nearly Abstract: Recent advances in deep-sea exploration everything brought up seemed full of luminous sparks. The technology coupled with an increase in worldwide biotic alcyonarians, the brittle-stars, and some annelids were the most surveys, biological research, and underwater photography brilliant. The Pennatulae, the Virgulariae, and the Gorgoniae shone in shallow water marine regions such as coral reefs, has with a lambient white light, so bright that it showed quite distinctly allowed for a relatively rapid expansion of our knowledge the hour on a watch …. We had another gorgeous display of in the global diversity of many groups of marine luminosity during this cruise …. The dredge came up tangle with organisms. This paper is part of the PLoS ONE review the long pink stems of the singular sea-pen Pavonaria quadrangularis. collection of WoRMS (the Worldwide Register of Marine Species), on the global diversity of marine species, and The Pavonariae were resplendent with a pale lilac phosphorescence treats the pennatulacean octocorals, a group of cnidarians like the flame of cyanogen gas: not scintillating …., almost commonly referred to as sea pens or sea feathers. This constant, sometimes flashing out at one point more brightly and also includes sea pansies, some sea whips, and various then dying gradually into comparative dimness, but always vermiform taxa. Pennatulaceans are a morphologically sufficiently bright to make every portion of a stem caught in the diverse group with an estimated 200 or more valid tangles or sticking to the ropes distinctly visible.
    [Show full text]
  • Hermit Crabs - Paguridae and Diogenidae
    Identification Guide to Marine Invertebrates of Texas by Brenda Bowling Texas Parks and Wildlife Department April 12, 2019 Version 4 Page 1 Marine Crabs of Texas Mole crab Yellow box crab Giant hermit Surf hermit Lepidopa benedicti Calappa sulcata Petrochirus diogenes Isocheles wurdemanni Family Albuneidae Family Calappidae Family Diogenidae Family Diogenidae Blue-spot hermit Thinstripe hermit Blue land crab Flecked box crab Paguristes hummi Clibanarius vittatus Cardisoma guanhumi Hepatus pudibundus Family Diogenidae Family Diogenidae Family Gecarcinidae Family Hepatidae Calico box crab Puerto Rican sand crab False arrow crab Pink purse crab Hepatus epheliticus Emerita portoricensis Metoporhaphis calcarata Persephona crinita Family Hepatidae Family Hippidae Family Inachidae Family Leucosiidae Mottled purse crab Stone crab Red-jointed fiddler crab Atlantic ghost crab Persephona mediterranea Menippe adina Uca minax Ocypode quadrata Family Leucosiidae Family Menippidae Family Ocypodidae Family Ocypodidae Mudflat fiddler crab Spined fiddler crab Longwrist hermit Flatclaw hermit Uca rapax Uca spinicarpa Pagurus longicarpus Pagurus pollicaris Family Ocypodidae Family Ocypodidae Family Paguridae Family Paguridae Dimpled hermit Brown banded hermit Flatback mud crab Estuarine mud crab Pagurus impressus Pagurus annulipes Eurypanopeus depressus Rithropanopeus harrisii Family Paguridae Family Paguridae Family Panopeidae Family Panopeidae Page 2 Smooth mud crab Gulf grassflat crab Oystershell mud crab Saltmarsh mud crab Hexapanopeus angustifrons Dyspanopeus
    [Show full text]