ALGAE AND MAN ALGAE AND MAN Based on lectures presented at the NATO Advanced Study Institute July 22 -August 11, 1962 Louisville, Kentucky Edited by Daniel F. Jackson Department of Civil Engineering Syracuse University Syracuse, New York PLENUM PRESS NEW YORK 1964 First Printing - November 1964 Second Printing - April 1966 Library of Congress Catalog Card Number 63-21218 ISBN-13: 978-1-4684-1721-0 e-ISBN-13: 978-1-4684-1719-7 DOl: 10.1007/978-1-4684-1719-7 ©1964 Plenum Press Softcover reprint of the hardcover 1st edition 1964 A Division of Consultants Bureau Enterprises, Inc. 227 West 17th Street New York, N. Y. 10011 No part of this publication may be reproduced in any form without written permission f1·om the publisher This book is dedicated to the Science Committee of the North Atlantic Treaty Organization in tribute to its activities in fostering international scientific thinking Preface With the continuous increase in human population and its constant demands on the aquatic environment, there has been a compounding of the interrelationships between algae and man. These relatively simple green plants not too long ago were often considered as merely biological curiosities. Within the past twenty-five years, with advances in technology and the increased eutrophication of lakes and streams, the interplay between algae and man has become more complex and more im portant. Problems of taste, odor, toxicity, or obnoxious growth caused by algae are unfortunately quite familiar to the water supplier and to the public health worker. Algae have met their role in the space age as a possible source for food or as a gas ex changer. In order to explore any of these practical problems, it is essential to have adequate, basic knowledge of algal taxonomy, physiology, cytogenetics and ecology. This book is the outgrowth of a North Atlantic Treaty Organi zation Advanced Study Institute in which authorities in both the applied and basic fields of phycology, as well as in cognate disci plines, met and discussed various topics related to algae. It is of significance to note that this was the first NATO Advanced Study Institute to be held in the United States and that it had for its theme a subject which is of import for the welfare of all mankind. Special acknowledgment is due the Scientific Affairs Division of NATO for providing the funds with which to conduct the Advanced Study Institute. Dr. Hans J0rgen Helms and Dr. B. Coleby, both of the Scientific Affairs Division, were instrumental in sponsorship of the program. Recognition and thanks are ex tended to President Philip G. Davidson for his graciousness in permitting the Advanced Institute to be held on the University of Louisville campus. vii viii Algae and Man Among the many other individuals who have helped in either conducting the Advanced Institute or in preparing this book are the members of the editorial staff of Plenum Press, who have been most helpful in all respects, and, as always, in any of my endeavors, my wife, Bettina Jackson, who is my unfailing partner. A salute is due each of the contributors to this book for his cooperation as well as to his scientific achievements. Through the conscientious efforts of each, a more complete understanding of the relations between algae and man has been achieved. DANIEL F. JACKSON Syracuse, New York Contents Contributions of Current Research to Algal Systematics . 1 G. W. Prescott Criteria and Procedures in Present-Day Algal Taxonomy... 31 C. van den Hoek The Gross Classification of Algae . 59 Tyge Christensen The Cytology of the Phaeophyta - A Review of Recent Develop- ments, Current Problems, and Techniques . 65 Margaret Roberts Environmental Conditions and the Pattern of Metabolism in Algae ...................... 77 G. E. Fogg Micronutrient Requirements for Green Plants, Especially Algae 86 Clyde Eyster Some Problems Remaining in Algae Culturing. 120 A. G. Wurtz The Ecology of Benthic Algae. 138 F. E. Round A Discussion of Natural and Abnormal Diatom Communities 185 Ruth Patrick The Ecology of Plankton Algae . 205 James B. Lackey Principles of Primary Productivity: Photosynthesis Under Com- pletely Natural Conditions 221 Jacob Verduin Algae in Water Supplies of the United States. 239 C. Mervin Palmer Algal Problems Related to the Eutrophication of European Water Supplies, and a Bio-Assay Method to Assess Fertilizing Influences of Pollution on Inland Waters . 262 Olav Skulberg The Biotic Relations with Water Brooms. 300 George P. Fitzgerald Toxic Algae 307 Paul R. Gorham ix x Contents Extracellular Products of Algae .... . . . . . . . . .. . .. 337 Marcel Lefevre Algae and Medicine ........................................ . 368 David Schwimmer and Morton Schwimmer Mass Culture of Microalgae for Photosynthetic Gas Exchange 413 Richard 1. Benoit The Future of Phycology .............................................. . 426 F. Evens Contributions of Current Research to Algal Systematics G. W. Prescott Department of Botany Michigan State University East Lansing, Michigan It is fitting, and a compliment to phycologists, that this sym posium is not entitled "Man and Algae"-those organisms which today represent the most venerable of all chlorophyll-bearing life on earth. Algae are certainly here to stay and of course we are less sure about Man, so it is appropriate that emphasis be placed on algae and that we think of what these plants mean to Man, this marauding upstart on the biological scene. It is also appropriate that attention be directed toward system atics to throw up a backdrop for the consideration of the many aspects of phycology related to Man. In thinking about the contri butions of modern research to systematics it is impossible, conver sely, to divorce the role of systematics in research. Therefore, in following the subject at hand I may consciously or unconsciously take some license and reverse the emphasis. At the same time I shall take the opportunity to single out some phycological problems much in need of investigation. In so doing it may be that some fences will be broken down and I shall encroach, apologetically, upon the fields assigned to other participants in this conference. It will be impossible to avoid doing this, so intimately enmeshed are the facets of phycology. The current conservative concept of the groups of organisms which constitute the "algae" recognizes the following nine phyla: 1) Chlorophyta; 2) Cyanophyta; 3) Chrysophyta; 4) Euglenophyta; 5) Pyrrhophyta; 6) Rhodophyta; 7) Phaeophyta; 8) "Cryptophyta"; 9) "Chloromonadophyta". Such a division into nine phyla is based mostly upon the delineations proposed by Pascher (1931) who, 1 2 Prescott however, relegated the Cyanophyta to a position exterior to the algae "proper." (See also van Oye, 1961.) Whereas this scheme is subject to modification by some students of the algae it seems, in general, to be the most natural classification yet conceived. At the same time it must be recognized and kept in mind that research and the accumulation of knowledge require that the classification sys tem be fluid and that the composition of the phyla may be contin ually shifting. This is disconcerting of course to those who look for a definite and static system of classification such as is more nearly approached in the arrangement of land or "higher" plants. The familiarly recognized foundations for the definitions of algal phyla are: the chemistry of pigments and their relative amounts; the chemistry of food reserve; and cytology, including flagellation, flagella morphology and wall structure. Therefore, it is only to be expected that as our knowledge becomes more refined, the criteria for assignment of organisms to this or to that phylum, or the definitions of an algal group, should be revised. Especially is this true in respect to the varying degrees of importance attached to flagellation and to chemistry of food reserve. Recent and current information on such features no doubt will bring about some re alignments and the present Pascherian system eventually will give way to new, modern schemes that will see a reconstitution of the algal phyla. This is especially predictable for the Chrysophyta which, as we know, includes three subphyla that, at best, are somewhat incompatible. The majority of biologists accept the premise that systematics has the same fundamentally important relation to biology that math ematics have to the physical sciences. This is not to say, of course, that mathematical science is not fundamental as well to certain facets of biology. But the describing, pigeonholing and name-giving biologist, independent of mathematics, is something of a philosoph ical would-be artist who creates a fundamental backdrop for all other aspects of biology. His goal is to create order and to provide arrangement, a goal for which the giving of names to living and fossil organisms is only part of the task. It is also the systematist who collects facts, interrelates them, and accordingly attempts to derive concepts that may answer questions relative to phylogeny and evolution-questions which continually pique and tantalize the inquisitive mind. Because of the basically important tasks and duties of the
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