DOCSLIB.ORG

Mathilde Klein Thesis (PDF 4MB)

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
Mathilde Klein Thesis (PDF 4MB) CHARACTERISATION OF DUPLICATED HAEMOGLOBIN GENES IN BIVALVES Mathilde Klein Bachelor of Medical Laboratory Science, QUT Submitted in fulfilment of the requirements for the degree of Master of Applied Science (Research) School of Biomedical Sciences, IHBI Faculty of Health Queensland University of Technology 2017 Keywords Arcoida, Bivalves, Gene duplication, Genomics, Globins, Haemoglobin, Limoida, Molluscs, Transcriptome Keywords i Abstract Haemoglobins (Hbs) are found in virtually all phyla and are some of the most investigated proteins in biomedical sciences. These proteins exhibit an extraordinary diversity of form and function in invertebrate lineages. This provides a unique opportunity to explore the origin and evolution of Hbs yet little is known about their distribution, function and evolution in invertebrate lineages. To explore further the functions and evolution of those Hbs, recent transcriptome data for the Arcid bivalve Anadara trapezia is investigated here. This species shows the presence of duplicated Hb encoding genes suggesting that gene duplication may have been more extensive than previously thought in bivalves. This study tests the hypothesis that these duplicated genes show patterns of tissue specific expression and evidence of neofunctionalisation. This is shown here for at least three Hb encoding genes present in A. trapezia with strong tissue specific expression in haemolymph compared to other tissues. Furthermore, the expression of these genes remains unaffected by prolonged air exposure suggesting that neofunctionalisation may confer an evolutionary advantage to this bivalve. As well as the unique Hbs found in the bivalve order Arcoida, Hbs are also found in three other bivalve orders: Carditoida, Solemyoida and Veneroida. These four orders that possess Hbs provide compelling evidence for the independent evolution of these proteins in multiple bivalve lineages. To expand data on the distribution of Hbs in bivalves, a transcriptome sequence for Ctenoides ales in the order Limoida was generated in this project. Interrogation of the transcriptome shows the presence of at least three globin-like encoding genes including two Hb-like encoding genes providing preliminary evidence for another independent origin of Hb in a bivalve lineage. Overall, this study provides novel insights into the function, evolution and distribution of Hbs in bivalves by investigating two distantly related species. Results of this study are consistent with current theories that Hb diversity in bivalves is a result of repeated rounds of gene duplication providing the raw material for evolution. Investigation of hypoxic resistance also reinforces that greater expression of Hbs in haemolymph confers a physiological advantage suggesting that Hb would evolve more often in some lineages during adaptation to unfavourable environment conditions, particularly Abstract ii hypoxia and prolonged air exposure. The finding of Hb-like encoding genes in another bivalve lineage also supports the evolution of this gene family through independent evolution and gene duplication, and gives insight into the distribution of globin genes in bivalves which is still poorly understood. The investigation of Hb genes in these bivalves also contributes to further understand the role of Hbs and provides potential novel insights for resistance in hypoxic environments, disease control and resistance to pollution in aquaculture. Abstract iii Table of Contents Keywords .................................................................................................................................. i Abstract .................................................................................................................................... ii List of Figures ......................................................................................................................... vi List of Tables .......................................................................................................................... xi List of Abbreviations ............................................................................................................. xii Statement of Original Authorship ......................................................................................... xiii Acknowledgements ............................................................................................................... xiv Chapter 1: Introduction ............................................................................................ 1 1.1 Functional and structural diversity within the globin superfamily .................................1 1.1.1 Recently discovered globin genes ....................................................................................... 2 1.1.2 Myoglobins and haemoglobins ........................................................................................... 2 1.2 Evolution of haemoglobins .............................................................................................4 1.2.1 Gene duplication ............................................................................................................... 5 1.2.2 Divergent evolution of duplicated genes .......................................................................... 8 1.2.3 Convergent evolution ........................................................................................................ 9 1.2.4 Invertebrate haemoglobins .............................................................................................. 10 1.3 Bivalve haemoglobins .......................................................................................................12 1.3.1 Haemoglobins in the family Arcidae, Pteriomorphia subclass.......................................... 16 1.4 Aims ..................................................................................................................................19 1.5 Thesis Outline ...............................................................................................................20 Chapter 2: Tissue specificity and neofunctionalisation of haemoglobin genes in Anadara trapezia ....................................................................................................... 22 2.1 Background .......................................................................................................................22 2.2 Material and Methods .......................................................................................................23 2.2.1 Sample acquisition and tissue dissection .......................................................................... 23 2.2.2 RNA extraction and cDNA synthesis ............................................................................... 24 2.2.3 RT–PCR (Real Time PCR) ............................................................................................... 25 2.2.4 Candidate gene validation ................................................................................................. 26 2.2.5 RT-qPCR (Real Time quantitative PCR) for quantification of gene expression .............. 26 2.2.6 RT-qPCR data analysis ..................................................................................................... 27 2.3 Results ...............................................................................................................................28 2.3.1 Haemolymph analysis ....................................................................................................... 28 2.3.2 RT-PCR ............................................................................................................................ 30 2.3.3 Candidate gene validation ................................................................................................. 31 2.3.4 RT-qPCR for quantification of gene expression ............................................................... 31 2.4 Discussion .........................................................................................................................35 2.4.1 Haemolymph characteristics ............................................................................................. 35 2.4.2 Tissue specific expression and neofunctionalisation ......................................................... 36 2.5 Conclusion ........................................................................................................................37 Chapter 3: Functional annotation of the Ctenoides ales transcriptome .............. 39 3.1 Background .......................................................................................................................39 Table of contents iv 3.2 Materials and Methods ......................................................................................................41 3.2.1 Sample collection .............................................................................................................. 41 3.2.2 RNA extraction, library preparation and sequencing ........................................................ 41 3.2.3 Transcriptome assembly and validation ............................................................................ 44 3.2.4 Functional annotation of transcripts and mapping ............................................................ 44 3.2.5 Comparative transcriptomics ............................................................................................ 46 3.2.6 Candidate genes identification .........................................................................................
Load more

Recommended publications
  • Teresa Haigh Thesis Submitted for the Degree of Doctor of Philosophy
    r DIELECTROPHORETIC INVESTIGATIONS OF HAEMATOLOGICAL CELLS PROCEDURES AND APPLICATIONS Teresa Haigh Thesis submitted for the degree of Doctor of Philosophy University of York Department of Biology March 1995 DEDICATION This thesis is dedicated to my family and friends for all their support. 11 ABSTRACT The dielectrophoretic phenomenondescribes the translationalmotion of a particle such as a biological cell, in responseto a non-uniform electric field. Both the magnitude and the direction of the induced movementare dependentupon the electrical properties of the cell with respectto its surroundingsand thus are a function of cell composition, and vary according to the alternation frequency of the electric field. Quantification of the responsein terms of the number of cells which exhibit positive dielectrophoretic behaviour, i. e. towards greatestfield intensity, as a function of field frequency,enables characteristicspectra to be compiled. Exploitation of a dielectrophoretic technique for biological analysis offers several advantages; notably that measurements are non-invasive and require no pre- modification of the cell, thus potentially permitting the separation of populations of viable cells. The clinical applications of this phenomena have to date been restricted, since methods for investigating dielectrophoretic response have relied upon manual quantification and been limited by cell sample size. Such difficulties have been minimised by the development of automated detection and analysis systems, enabling a typical collection spectra to be generated within an hour. The development of a dielectrophoretic technique for rapid analysis of haematological cells has been described. A computer-based system was used to control voltage application to a micro-electrode chamber through which a cell suspension was circulated.
    [Show full text]
  • Taxonomy of Tropical West African Bivalves V. Noetiidae
    Bull. Mus. nati. Hist, nat., Paris, 4' sér., 14, 1992, section A, nos 3-4 : 655-691. Taxonomy of Tropical West African Bivalves V. Noetiidae by P. Graham OLIVER and Rudo VON COSEL Abstract. — Five species of Noetiidae are described from tropical West Africa, defined here as between 23° N and 17°S. The Noetiidae are represented by five genera, and four new taxa are introduced : Stenocista n. gen., erected for Area gambiensis Reeve; Sheldonella minutalis n. sp., Striarca lactea scoliosa n. subsp. and Striarca lactea epetrima n. subsp. Striarca lactea shows considerable variation within species. Ecological factors and geographical clines are invoked to explain some of this variation but local genetic isolation could not be excluded. The relationships of the shallow water West African noetiid species are analysed and compared to the faunas of the Mediterranean, Caribbean, Panamic and Indo- Pacific regions. Stenocista is the only genus endemic to West Africa. A general discussion on the relationships of all the shallow water West African Arcoidea is presented. The level of generic endemism is low and there is clear evidence of circumtropical patterns of similarity between species. The greatest affinity is with the Indo-Pacific but this pattern is not consistent between subfamilies. Notably the Anadarinae have greatest similarity to the Panamic faunal province. Résumé. — Description de cinq espèces de Noetiidae d'Afrique occidentale tropicale, ici définie entre 23° N et 17° S. Les Noetiidae sont représentés par cinq genres. Quatre taxa nouveaux sont décrits : Stenocista n. gen. (espèce-type Area gambiensis Reeve) ; Sheldonella minutalis n. sp., Striarca lactea scoliosa n.
    [Show full text]
  • Questioning the Functional Relevance of Mitochondrial Supercomplexes by Time-Resolved Analysis of the Respiratory Chain
    Questioning the functional relevance of mitochondrial PNAS PLUS supercomplexes by time-resolved analysis of the respiratory chain Martin Trouillarda, Brigitte Meunierb, and Fabrice Rappaporta,1 aInstitut de Biologie Physico-Chimique, Unité Mixte de Recherche 7141 Centre National de la Recherche Scientifique-Univ P et M Curie, 13 rue P et M Curie, 75005 Paris, France; and bCentre de Génétique Moléculaire, Unité Propre de Recherche 3404 Centre National de la Recherche Scientifique, Avenue de la Terrasse, 91198 Gif-sur-Yvette, France Edited by Marten Wikstrom, University of Helsinki, Helsinki, Finland, and accepted by the Editorial Board September 19, 2011 (received for review June 13, 2011) Mitochondria are the powerhouses of eukaryotic cells as they feed zation. Since then, many supercomplex assemblies from a wide metabolism with its major substrate. Oxidative-phosphorylation variety of organisms or organs have been biochemically charac- relies on the generation, by an electron/proton transfer chain, of terized, with a large diversity of stoichiometries and complex an electrochemical transmembrane potential utilized to synthesize compositions [see e.g., the recent exhaustive review by Lenaz and ATP. Although these fundamental principles are not a matter of Genova (11)]. This culminated in the isolation by Acin-Pérez, debate, the emerging picture of the respiratory chain diverges et al. of a functional “respirasome” from mammal cells, transfer- from the linear and fluid scheme. Indeed, a growing number of ring electrons all the way from NADH or succinate to molecular pieces of evidence point to membrane compartments that possibly oxygen (12). In addition, the composition and abundance of the restrict the diffusion of electron carriers, and to supramolecular biochemically characterized supercomplexes have been shown to assembly of various complexes within various kinds of super- vary with growth or physiological conditions (13, 14), supporting complexes that modulate the thermodynamic and kinetic proper- their physiological significance.
    [Show full text]
  • Journal Ofbiotechnology
    Journal of Biotechnology 187 (2014) 1–9 Contents lists available at ScienceDirect Journal of Biotechnology journal homepage: www.elsevier.com/locate/jbiotec In vivo biodistribution and oxygenation potential of a new generation of oxygen carrier Tony Le Gall a,b, Valérie Polard c, Morgane Rousselot c, Auréline Lotte c, Mouna Raouane d, Pierre Lehn a,b, Paule Opolon e, Elisabeth Leize f, Eric Deutsch d, Franck Zal c, Tristan Montier a,b,g,∗ a Unité INSERM 1078, SFR 148 ScInBioS, Université de Bretagne Occidentale, Université Européenne de Bretagne, 46 rue Félix Le Dantec, CS51819, 29218 Brest Cedex 02, France b Plateforme SynNanoVect, SFR 148 ScInBioS, Université de Bretagne Occidentale, Faculté de Médecine, 22 rue Camille Desmoulins, 29200 Brest, France c HEMARINA SA, Aéropôle centre, Biotechnopôle, 29600 Morlaix, France d Unité INSERM 1030, Radiothérapie Moléculaire, Université Paris XI, Institut Gustave Roussy, 114 rue Edouard Vaillant, 94805 Villejuif, France e Unité de Pathologie Expérimentale, Institut Gustave Roussy, 114 rue Edouard Vaillant, 94805 Villejuif, France f CHRU de Brest, Département de Prothèses, UFR Odontologie, Brest F29238, France g DUMG, Université de Bretagne Occidentale, CHRU de Brest, service de biochimie et de pharmaco-toxicologie, 5 avenue du Maréchal Foch, 29200 Brest, France article info a b s t r a c t Article history: Natural giant extracellular hemoglobins (Hbs) from polychaete annelids are currently actively inves- Received 3 October 2013 tigated as promising oxygen carriers. Their powerful oxygenating ability and their safety have been Received in revised form 4 July 2014 demonstrated in preclinical studies, motivating their development for therapeutic and industrial applica- Accepted 7 July 2014 tions.
    [Show full text]
  • Biochemical Responses in Penaeids Caused by Contaminants
    Aquaculture 191Ž. 2000 163±168 www.elsevier.nlrlocateraqua-online Biochemical responses in penaeids caused by contaminants Afonso Celso Dias Bainy ) Departamento de Bioquõmica,ÂàCentro de Ciencias Biologicas, UniÕersidade Federal de Santa Catarina, Florianopolis, SC, 88040-900 Brazil Received 1 March 2000; accepted 9 May 2000 Abstract The shrimp aquaculture industry has become increasingly aware of the need for developing sensitive and precise diagnostic toolsŽ. Biomarkers with predictive capability for assessing the toxic effect of commonly encountered chemicals on shrimp culture. Potentially damaging com- pounds used in shrimp culture include disinfectants, therapeutics, feed additives, algicidals, pesticides, and fertilizers. These chemicals may cause biological damage at all life stages during shrimp production. Since many chemicals may be stressors to the organisms, lower production rates may occur, compromising the sustainability of the shrimp production. This manuscript gives a brief overview about toxic effects associated with the chemicals used directly or indirectly during shrimp production. q 2000 Elsevier Science B.V. All rights reserved. Keywords: Penaeid; Biomarker; Contamination 1. Introduction In many countries, the consumption of shrimps has grown yearly but the natural stocks of these decapod crustaceans have been intensively depleted. Alternatively, shrimp culture has been adopted and consolidated as one of the largest profitable aquaculture activities all over the world. Methods of shrimp cultivation adopted by the shrimp aquaculture companies vary from intensive farming to extensive rearing systems. The former requires auxiliary inputs and capital, while the second is to a great extent dependent on natural processes ) Tel.: q55-48-3316561; fax: q55-48-3319672. E-mail address: [email protected]Ž.
    [Show full text]
  • Part 4 Functional Biochemistry
    МІНІСТЕРСТВО ОХОРОНИ ЗДОРОВ'Я УКРАЇНИ Харківський національний медичний університет PART 4 FUNCTIONAL BIOCHEMISTRY Self-Study Guide for Students of General Medicine Faculty in Biochemistry ЧАСТИНА 4 ФУНКЦІОНАЛЬНА БІОХІМІЯ Методичні вказівки ДЛЯ ПІДГОТОВКИ ДО ПРАКТИЧНИХ ЗАНЯТЬ З БІОЛОГІЧНОЇ ХІМІЇ (Для студентів медичних факультетів) Затверджено вченою радою ХНМУ. Протокол № 4 від 27.04.2017 р. Approved by the Scientific Council of KhNMU. Protocol 4 (April 27, 2017) Харків ХНМУ 2017 Self-study guide for students of general medicine faculty in biochemistry. Part 4. Functional biochemistry / comp. O. Nakonechna, S. Stetsenko, L. Popova, A. Tkachenko. – Kharkiv : KhNMU, 2017. – 80 p. Методичні вказівки для підготовки до практичних занять з біологічної хі- мії (для студентів медичних факультетів). Частина 4. Функціональна біохімія / упоряд. О.А. Наконечна, С.О. Стеценко, Л.Д. Попова, А.С. Ткаченко. – Харків : ХНМУ, 2017. – 80 с. Authors Nakonechna O. Stetsenko S. Popova L. Tkachenko A. Автори: О.А. Наконечна С.О. Стеценко Л.Д. Попова А.С. Ткаченко - 2 - SOURCES For preparing to practical classes in "Biological Chemistry" Basic Sources 1. Біологічна і біоорганічна хімія: у 2 кн.: підручник. Кн. 2. Біологічна хі- мія / Ю.І. Губський, І.В. Ніженковська, М.М. Корда, В.І. Жуков та ін.; за ред. Ю.І. Губського, І.В. Ніженковської. – К.: ВСВ «Медицина», 2016. – 544 с. 2. Губський Ю.І. Біологічна хімія. Підручник / Губський Ю.І. – Київ- Вінниця: Нова книга, 2007. – 656 с. 3. Губський Ю.І. Біологічна хімія / Губський Ю.І. – Київ–Тернопіль: Укрмедкнига, 2000. – 508 с. 4. Гонський Я.І. Біохімія людини. Підручник / Гонський Я.І., Максимчук Т.П., Калинський М.І. – Тернопіль: Укрмедкнига, 2002.
    [Show full text]
  • The 1940 Ricketts-Steinbeck Sea of Cortez Expedition: an 80-Year Retrospective Guest Edited by Richard C
    JOURNAL OF THE SOUTHWEST Volume 62, Number 2 Summer 2020 Edited by Jeffrey M. Banister THE SOUTHWEST CENTER UNIVERSITY OF ARIZONA TUCSON Associate Editors EMMA PÉREZ Production MANUSCRIPT EDITING: DEBRA MAKAY DESIGN & TYPOGRAPHY: ALENE RANDKLEV West Press, Tucson, AZ COVER DESIGN: CHRISTINE HUBBARD Editorial Advisors LARRY EVERS ERIC PERRAMOND University of Arizona Colorado College MICHAEL BRESCIA LUCERO RADONIC University of Arizona Michigan State University JACQUES GALINIER SYLVIA RODRIGUEZ CNRS, Université de Paris X University of New Mexico CURTIS M. HINSLEY THOMAS E. SHERIDAN Northern Arizona University University of Arizona MARIO MATERASSI CHARLES TATUM Università degli Studi di Firenze University of Arizona CAROLYN O’MEARA FRANCISCO MANZO TAYLOR Universidad Nacional Autónoma Hermosillo, Sonora de México RAYMOND H. THOMPSON MARTIN PADGET University of Arizona University of Wales, Aberystwyth Journal of the Southwest is published in association with the Consortium for Southwest Studies: Austin College, Colorado College, Fort Lewis College, Southern Methodist University, Texas State University, University of Arizona, University of New Mexico, and University of Texas at Arlington. Contents VOLUME 62, NUMBER 2, SUmmer 2020 THE 1940 RICKETTS-STEINBECK SEA OF CORTEZ EXPEDITION: AN 80-YEAR RETROSPECTIVE GUesT EDITed BY RIchard C. BRUsca DedIcaTed TO The WesTerN FLYer FOUNdaTION Publishing the Southwest RIchard C. BRUsca 215 The 1940 Ricketts-Steinbeck Sea of Cortez Expedition, with Annotated Lists of Species and Collection Sites RIchard C. BRUsca 218 The Making of a Marine Biologist: Ed Ricketts RIchard C. BRUsca AND T. LINdseY HasKIN 335 Ed Ricketts: From Pacific Tides to the Sea of Cortez DONald G. Kohrs 373 The Tangled Journey of the Western Flyer: The Boat and Its Fisheries KEVIN M.
    [Show full text]
  • Biodiversità Ed Evoluzione
    Allma Mater Studiiorum – Uniiversiità dii Bollogna DOTTORATO DI RICERCA IN BIODIVERSITÀ ED EVOLUZIONE Ciclo XXIII Settore/i scientifico-disciplinare/i di afferenza: BIO - 05 A MOLECULAR PHYLOGENY OF BIVALVE MOLLUSKS: ANCIENT RADIATIONS AND DIVERGENCES AS REVEALED BY MITOCHONDRIAL GENES Presentata da: Dr Federico Plazzi Coordinatore Dottorato Relatore Prof. Barbara Mantovani Dr Marco Passamonti Esame finale anno 2011 of all marine animals, the bivalve molluscs are the most perfectly adapted for life within soft substrata of sand and mud. Sir Charles Maurice Yonge INDEX p. 1..... FOREWORD p. 2..... Plan of the Thesis p. 3..... CHAPTER 1 – INTRODUCTION p. 3..... 1.1. BIVALVE MOLLUSKS: ZOOLOGY, PHYLOGENY, AND BEYOND p. 3..... The phylum Mollusca p. 4..... A survey of class Bivalvia p. 7..... The Opponobranchia: true ctenidia for a truly vexed issue p. 9..... The Autobranchia: between tenets and question marks p. 13..... Doubly Uniparental Inheritance p. 13..... The choice of the “right” molecular marker in bivalve phylogenetics p. 17..... 1.2. MOLECULAR EVOLUTION MODELS, MULTIGENE BAYESIAN ANALYSIS, AND PARTITION CHOICE p. 23..... CHAPTER 2 – TOWARDS A MOLECULAR PHYLOGENY OF MOLLUSKS: BIVALVES’ EARLY EVOLUTION AS REVEALED BY MITOCHONDRIAL GENES. p. 23..... 2.1. INTRODUCTION p. 28..... 2.2. MATERIALS AND METHODS p. 28..... Specimens’ collection and DNA extraction p. 30..... PCR amplification, cloning, and sequencing p. 30..... Sequence alignment p. 32..... Phylogenetic analyses p. 37..... Taxon sampling p. 39..... Dating p. 43..... 2.3. RESULTS p. 43..... Obtained sequences i p. 44..... Sequence analyses p. 45..... Taxon sampling p. 45..... Maximum Likelihood p. 47..... Bayesian Analyses p. 50..... Dating the tree p.
    [Show full text]
  • Structural and Functional Diversity of Microbial Globins
    European Journal of Molecular & Clinical Medicine ISSN 2515-8260 Volume 07, Issue 07, 2020 Structural And Functional Diversity Of Microbial Globins 1Shiv Kumar 1Department of Medical Laboratory Sciences (School of Physiotherapy and Paramedical Sciences), Lovely Professional University, Phagwara, Punjab (India) – 1444 Email ID: [email protected]; ABSTRACT Hemoglobin and myoglobin are heme proteins that can bind oxygen reversibly and were first reported in vertebrates. Later on, occurrence of hemoglobin was found in all forms of life ranging from prokaryotes to eukaryotes. Myxobacteria are Gram-negative, rod-shaped eubacteria and that belong to delta proteobacteria. They predominantly thrive in soil and have very large genome size with high G+C content in comparison to other eubacteria. It is evident that the globins have been characterized predominantly from bacteria involved in photosynthesis, nitrogen fixation andpathogenicity, where these are implicated in and the protection of photosynthetic machinery from the oxidative damage,protection of nitrogen fixing enzyme from nitrosative and oxidative stress and the virulence of the host, respectively. Keywords: Myxobacteria, hemoglobin, myoglobin, phytoglobin, androglobin. 1. INTRODUCTION Globins are heme containing proteins which comprise five to eight α-helices that are connected by short intervening loops and have a common characteristic of binding oxygen reversibly [1-4]. These are widely reported in all the three domains of life forms such as Archaebacteria, eubacteria and eukaryotes [1]. In addition to their well-known function of oxygenstorage and transport, they are also involved in nitric oxide detoxification, oxygensensing or act as oxidation-reduction sensors [2]. Globins containhighly conserved structural folds, which build the protein crevice to hold the heme group [5, 6].
    [Show full text]
  • Circulatory Systems Vertebrate Hearts Respiratory Pigments
    3/25/2015 Circulatory Systems Vertebrate hearts • Chambers isolate pulmonary and • Open vs. closed systemic circuits • Components of • Blood pressure and regulation of hemolymph or blood flow rates • Hearts • Vasoconstriction and dilation – Simplest forms are thickened smooth • Flow velocity minimized, area muscle in arteries maximized in capillaries • Flow pressure minimized in largest veins Respiratory Pigments • Molecules that bind oxygen, facilitate transfer from respiratory surface to tissues that need it. Tissues Lungs Bind Release High PO2 Low PO2 • Oxygen affinity – [ ] • % = x 100 [] 1 3/25/2015 Structure of Hemoglobin Respiratory Pigments Oxygen Molecular • Oxyhemoglobin – bound with O2, reversible Color Cells or Pigment Structure capacity Weight Animal Groups (change) Solution • Deoxyhemoglobin – not bound with oxygen, Fe reduced (ml g-1) (kDa) Mollusks, • Carbaminohemoglobin – bound with CO , reversible Hemocyanin Blue 2 Protein+Cu2+ 0.3-0.5 25-7000 Solution cephalopods, (colorless) • Carbon monoxide hemoglobin – combined with CO, not arthropods Nematodes, reversible Hemoglobin Protein+heme Red 1.2-1.4 16-2000 Either annelids, +Fe2+ (purple/blue) vertebrates Protein+heme Annelids, marine Chlorocruorin Green 0.6-0.9 3000 Solution +Fe2+ polychaetes Protein Violet Brachiopods, some Hemerythrin 1.6-1.8 16-125 Either +Fe2+ (colorless) marine annelids • Intracellular vs. solution • Various other forms, recall gene families Before/after methemoglobinemia P50 Oxygen carrying capacity • Total oxygen capacity of blood depends on
    [Show full text]
  • Patterns of Morphological Diversity Among and Within Arcid Bivalve Species Pairs Separated by the Isthmus of Panama
    J. PaleonL, 75(3), 2001, pp. 590-606 Copyright © 2001, The Paleontological Society 0022-3360/01/0075-590$03.00 PATTERNS OF MORPHOLOGICAL DIVERSITY AMONG AND WITHIN ARCID BIVALVE SPECIES PAIRS SEPARATED BY THE ISTHMUS OF PANAMA PETER B. MARKO'-3 AND JEREMY B. C. JACKSON'.^ 'Smithsonian Tropical Researcii Institute, Box 2072, Balboa, Republic of Panama, and ^Scripps Institute of Oceanography, Geosciences Research Division, University of California San Diego, La JoUa 92093-0224 U.S.A. ABSTRACT•Geminate species are morphologically similar sister-species found on either side of the Isthmus of Panama. The existence of all geminates in the tropical Eastern Pacific ocean and the Caribbean Sea is most often explained by vicariance: closure of the Central American Seaway 3.1 to 3.5 Ma simultaneously isolated populations of species with amphi-American distributions. In this paper, we test the potential of morphological measurements for discriminating between Recent geminate species pairs from three genera (Area, Arcopsis, and Barbatia) in the bivalve family Arcidae and examine the prospects for distinguishing nominal species in the fossil record. Fourteen morphological variables were used to characterize shell shape and multivariate methods were used to discriminate between five Recent species pairs. Collection sites were also used as a priori groups for discrimination to describe patterns of intraspecific morphological variation and to evaluate differences among samples from different geographic regions. On average, 84 percent of specimens within geminate pairs are classified correctly following five separate discriminant analyses with nominal species as the grouping variable. Although all but one arcid species pair are discriminated with high statistical significance, some collection sites within species are highly morphologically distinct.
    [Show full text]
  • Documenting Benthic Invertebrate from Rocky Environments in the REMAPE
    MANUAL ON DOCUMENTING BENTHIC INVERTEBRATE COMMUNITIES FROM ROCKY ENVIRONMENTS IN THE MARINE RESERVE EL PELADO, SANTA ELENA, ECUADOR ISBN: 978-9942-36-314-5 Sascha Steiner, Antonella Lavorato, Jenny Rodríguez A MANUAL ON DOCUMENTING BENTHIC INVERTEBRATE COMMUNITIES FROM ROCKY ENVIRONMENTS IN THE MARINE RESERVE EL PELADO, SANTA ELENA, ECUADOR 1, 2 2 2 Steiner SCC , Lavorato AL , Rodríguez J 1 Secretaría Nacional de Educación Superior, Ciencia, Tecnología Innovación (SENESCYT), Guayaquil, Ecuador 2 Escuela Superior Politécnica del Litoral (ESPOL), Centro Nacional de Acuicultura e Investigaciones Marinas (CENAIM), Guayaquil, Ecuador Proyecto Prometeo: Creación e implementación de un sistema de información marino y atlas dinámico georeferenciado de biodiversidad del ecosistema del área marina protegida El Pelado, Santa Elena, Ecuador Proyecto: Caracterización de la biodiversidad microbiológica y de invertebrados de la reserva marina El Pelado a escalas taxonómica, metabolómica y metagenómica, para uso en salud humana y animal This handbook will be cited as follows Steiner, S. Lavorato, A. Rodríguez, J. A MANUAL ON DOCUMENTING BENTHIC INVERTEBRATE COMMUNITIES FROM ROCKY ENVIRONMENTS IN THE MARINE RESERVE EL PELADO, SANTA ELENA, ECUADOR. Escuela Superior Politécnica del Litoral. 88 pp. Guayaquil, Ecuador. Technical Review: María José Brito Cover and text design: María José Brito and Juan Carlos Castillo Cover Pictures: Karla Jaramillo Supervision and Approval: Jenny Rodríguez and Gabriela Agurto Printer and binder: Gabriel Zurita Correspondence and Exchange: Cecilia Campoverde ISBN: 978-9942-36-314-5 Short Title Documenting benthic invertebrate from rocky environments in the REMAPE. Autorización de Investigación Científica: N°005-17 IC-FAU- DPSE/MA Contrato Marco de Acceso a Recurso Genético: MAE- DNB-CM-2015-0021 PROLOGUE Marine biodiversity offers a wide range of opportunities for biotechnological development.
    [Show full text]