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THE ROLE OF
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ALGAE AND PLANKTON
IN MEDICINE
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Morton Schwimmer, M.D.,
Clinical Assistant in Medicine,
The New York Medical College,
Metropolitan Medical Center
and
David Schwimmer, M.D., F.A.C.P.,
Assistant Professor of Medicine,
The New York Medical College,
Metropolitan Medical Center
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10430 A003 YH10N 3H
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GRUNE & STRATTON
NEW YORK AND LONDON
1953

·Library of Congress Catalog Card No 53-11353
Copyright 1955
GRUNE & STRATTON, INC.
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·ACKNOWLEDGMENT WE GRATEFULLY ACKNOWLEDGE the extraordinary courtesies extended us by the library staffs of the following: The New York Academy of Medicine, New York Public Library, New York Botanical Gardens, Museum of Natural History, Explorers Club, Rockefeller Institute for Medical Research, Columbia University, The New York Times, Library of Con. gress, Army Medical Library, and the New York Medical College To Di Jorgen Jorgensen and Dr. Enrique Tejera, of the Venezuelan Ministry of Health, we express appreciation for patiently demonstrating to one of us (MS) the extensive algal culture techniques and feeding projects at the Cabo Blanco Leprosatium at Maiquetia, Venezuela. We are especially indebted to Miss Irma Frisch Farrington, of the New York Academy of Mediane library staff, for her invaluable aid in locating clusive material, arranging for translations, and helping prepare the bibliography Sincere thanks are also extended to the following for translations from the French, German, Italian, Spanish, Russian Japanese, and Afrikaans. Miss Freda Monkarz and her colleagues at the United Nations Headquarters in New York City; and Des Roger Duvoisin, Johan Senekal, Peter Masley, and Ulrich Batzdorf, of the Metropolitan Hospital The Auibors ·FOREWORD THIS MONOGRAPH on the role of algae and plankton in medicine is a timely one It 15, so far as the present writer knows, the only medical survey of the field, and it appears at a time when specialists in various non-medical branches of biology are exhibiting a keen interest in the subject The literature is expanding so rapidly that it may soon be difficult to keep abreast of the advances Although algae and plankton have been empirically utilized as food since ancient times, scientific observations were at first limited mainly to academic botanical aspects In recent years, world food shortages have focused attention on the nutritional utility of these substances for men and the animals upon which he is dependent An important corollary--the relationship of these forms of life to cliatcal medicine—appears to have been neglected until the present publication It should be stressed at the outset that the authors have been content merely to point out the great medical potential of algae and plankton, rather than to promote panaceas Nor have they sought, with their his. torical references, to justify vacious dietary regimens and practices of primitive medicine The fact that additional investigations might rationally explain ancient or aboriginal medical practices is presented merely as a matter of interest rather than importance However, in the opinion of this writer, no apology would be necessary even if the authors had utilized the historical approach and examined modern medicine in the light of primitive conceptions Even Hippocrates, in his "On Ancient Mediane," stated that the greatest medical discovery up to his time (400 BC) had involved countless experiments ranging from field to kitchen and finally resulted in the baking of bread Perhaps the discovery of sugar was almost as complex The historical approach would also have revealed that cinchona for malacia, ipecac for amoebic dysentery, chaulmoogra for leprosy, iron filiags for anemia, and digitalis for the heart all represent refinements of the dross of folk medicine and likewise require no apology. Pertinent examples can be ated from medi·vi FOREWORD cine of the present century About 25 years ago, Indian physicans brought reports about the action of a drug and specimens of the preparation to the writer for evaluation and criticism. Preoccupation with seemingly more pressing problems led to filing the reports and discarding the specimens Had time permitted an examination, Rauwolfa might have entered medicine at that time The authors are less concerned with past neglect than they are with emphasizing possible immediate medical applications in a promising field Much important data has been presented on nutritional, physiological, toxicological, and public health problems Also, some rather interesting clinical hypotheses have been propounded, particularly in the neurologic realm Since a vast majority of the references included in their excellent bibliography are derived from non-medical sources, the implication of neglect by physicians does not seem unwarranted Although countless other subjects occur to the writer (and, it is hoped, to the reader), an unwritten tule must be obeyed the introduction should not be longer than the actual text In summary, the authors have surveyed the world literature on algae and plankton in relation to medı. ane. They have appended an excellent bibliography of widely scattered, but generally available, references, most of which will not be found in the usual medical index They have accented the recent literature dealing with promising fields of investigation and possible application to current medicine There has been made available a concise introduction to a subject old as recorded medical history, yet so new that many medical investigators do not realize its existence. LINN | BOYD, M.D, F.A.C.P Professor of Medicine The New York Medical College, Metropolitan Medical Center ·CONTENTS FOREWORD, by Linn ) Boyd, MD, FACP, v 1. INTRODUCTION, 1 Increasing importance of algae and plankton Multiplicity of reports in nonmedical publications. Physicians' general ignorance of subject Objectives of book IT DEFINITIONS AND CLASSIFICATIONS, 2 A ALGAE, 2 Definition Distribution and habitat Size and structure Metabolism and composition Role of photosynthesis Confusion in classifications Current classzcatron into seven Divisions Most important groups B PLANKTON, 3 Interrelationship of algae and plankton Definitions Significance in aquatic nutritional cycle Zooplankton Phytoplankton Ultraplankton Naanoplankton Hekzstoplankton Littoral, benthal, and limnetic growths Variation of growth with environment III MACROSCOPIC ALGAE (SEAWEED), 5 A NUTRITIONAL ASPECTS, 5 Seaweeds as food in ancient times Popularity in Orient Diversity of names and uses Purple laver Green laver Dulse Agar and isenglass Irish moss Seaweed bread Sausage casings Custard powder Nutritional content Preponderance of carbohydrate Structure of carbohydrates Paucity of proteins Amino acids Lipid content Minerals and trace clements Iodine Seaweed as fertilizer and animal provender Feeding experiments Neurologic disorder in lambs. Relatron to "swapback" Effectiveness of supplementing diet with copper B MEDICINAL Uses, 13 Legendary and empirical origins Employment by Chinese Lavers for throat a:lments, gouter, and dropsy Seaweed derivatives for menstrual difficulties, abscesses, cancer, and inflammations. Birds' nest soup as sphrodisiac, Algal preparations in India for broachocoelc and goiter Syphilis Poultices of flamentous algae Hawasan extracts for digestive disturbances Scabies Constipation Earache Goiter Renal disorders. Infections and sprains. Roman use of algaz for sores, wounds, and gout Corraline as external absorbent for skin lesions, as internal absorbent for hepatic and digestive disorders "Sea lettuce" as source of salt for Iroquois. Todine. containing seaweeds in thyroid disease Agar as bulk, demulceat, and absorbent in gastro-intestinal disturbances Seaweeds as anthelmintics Irish moss as respiratory ·viii CONTENTS demulcent. Carragheen poultices. Scrofula. Dulse for pyrexia Bladderwrack and agar in obesity Stipes for fistulas, wounds, and labor Present status of agar, algio, and carragheenin Seaweed "gums" or phycocolloids, Trichomonal vaginitis Cellular plasma of manne algte Agar as culture medium and as vehicle for medications Algin combined with oral and parenteral medications. Possible allergenicity of alginates. Alginates in constipation and other digestive disorders Other uses of alginates in medicine and dentistry. Phycocolloids in dairy and baking industries Antibiotic properties of macroscopic algae. Anticoagulant activity of algal deriva- Laminarın as blood plasma substitute Radioactivity of algae exposed to Atomic explosions C MISCELLANEOUS USES, 21 Diverse and exotic applications of macroscopic algae tives IV PLANKTON, 22 A ZOOPLANKTON, 22 Definition and composition Distribution of various types Marine 200. plankton as food for sailors Earliest reports on eating of zoop Biochemical analyses War interest in zooplankton Tidal flow filtration Studies at Heligoland Plankton content of Russian seas. Batish and American observations Effect of zooplankton on water requirements of castaways Human feeding studies Rat feeding with marine and fresh water zooplankton Experience with plankton by passengers of the Kon-Tikr, the Heretique, and the Seven while Sisters B PHYTOPLANKTON (MICROSCOPIC ALGAE), 27 1 GENERAL NATURE, 27 Composition and chief types Basic role as ultimate nutrient of marine life Effect on color and phosphorescence of sea Early studies on microscopie algae More recent observations on culture, photosynthesis, and metabolism Review manuals on microalgae 2 METABOLISM, CULTURE, AND NUTRITIONAL VALUES, 29. Importance of phytoplankton as world food supplement Excellent digestibility. High content of protesn, amino acids, lipids, vitamins, and minerals Comparative analyses of algae and feld plants. Cellular structure Key pigments chlorophylls, carotenoids, phy. cobilins Distribution, nature, and functions of pigments Protoporphyrin precur. sors Chlorophyll and heme Vital role of pigments in photosynthesis Fluorescence Algae without pigment Algal reproduction Binary fisston. Sexuality and chemical factors Phases of algal growth in culture. Environmental and nutrient effects upon algal composition Enzyme systems Algal proteins and amino acids. Nitra gen fixation in algat culture Nitrogen fixation in agriculture, Algal carbohydrates Importance of lipids Variability of quantity and qualsty of lipids Chemical characteristics. Sterols Their use in synthesis of steroidal hormones High vitamin content of microalgae Variations with environmental and enzymatic dianges Vitamin 2551ys Major requited nutrients Trace elements or micronutrients Chelating agents Unidentified growth factors Mass culture of microscopic algae Basic requirements Types of apparatus Pilot plant studies in various countries Feeding experiments Palatability and acceptability.

·CONTENTS IX 3 MEDICAL ASPECTS, 45 a F'ater supply and Sewage Disposal, 45 Possible adverse effects of phytoplanktonic organisms "Water bloom" in fresh water bodies "Red tide," "yellow tide," "red snow." Unpleasant odors of water due to phytoplankton Origin of bad odors and tastes. Mechanism of production Chemical substances present Algal ouls Products of photosynthesis Products of algal decomposition Death of fish and other aquatic organisms Oxygenation Measures to control algal growth Algal oxidation ponds for sewage treatment Simplicity and efficiency of oxidation ponds Algal oxidation to convert nitrogenous wastes into food protein Radioactivity of phytoplankton in waters near atomic plants Algae and mosquito control b Animal Intoxication, 48 Many reports of animal intoxication and death due to toxic phytoplankton Table summary of 38 incidents of animal intoxication reported Confusion with effects of microorganisms, metallıc and chemical toxins, and poisonous land plants Symptomatology varied Predominance of neuromuscular and respiratory involvement Photosensitivity Jaundice Weakness and paralysis Convulsions Dyspnea Negativity of autopsy studies Experimental studies Effects of Microrystir extracts on nervous system, circulation, liver. Degrees of intoxication Parenchymatous, hydropic, fatty degeneration, necr Studies on toxios Their chemical and physical natures c Human Intoxications, 33 Phytoplanktonic role Clinical pictures dysenterral disorders, systemic allergies, local allergic eruptions Table suromary of reported human reactions to phytoplankton Epidemic intestinal disturbances in US National Parks and West Virginia Symptomatology Association with algal bloom in water supply Systemic allergy to Oscillatoracede in Wisconsin Lake Positive skin reactions to Microcysts Local allergic phenomena Urticarial "sexbathers' cruption" Acute urticaria from "red tide" in Gulf of Mexico Dermatitis from phycocyanın pigment in Anabaena in Pennsylvania lake Indirect role of phytoplankton in human disease states Prototheca and Blastocystis in tropical sprue Microscopic algae in mycotic skin lesions actinomycosis, paracoccidiosts Relation of antibiotic chlorellin" Antibiotic phyceine Antibacterial agents in Sh. chococcus and Protosyphor Algae in the diagnosis of death by drowning Intoxcations resulting from ingestion of certasa fish Summary of 58 documented episodes of fish poisoning Acute "paralytic" shellfish porsoning Predominantly neurologic manifestations Epidemiology Etiology Relationship to ingestion of toxic Gon ul.sx Nature of toxin in Gonyaular Negatwity of autopsy findings in acute paralytic shellfish poisoning Positive findings in chronically poisoned experimental animals Central nervous system changes Renal damage Other toxins from algae Inhecent toxicity of puffer fish and some barracuda Epidemics of fish poisoning during World War II Listing of poisonous tish Clinical classification of Ichthyosarcotoxicosis Teirsodon (puffer) poisoning Nature of retraodontoxía Clinical manifestations Gymnothorax (moray cel) poisoning Neurotoxie manifestations Cigualera and its clinical picture Scombroid poisoning Histamine-like manifestations "Haffkrankheit" Relation to contact with 6sh and plankton Symptomatology Pathological findings Methods of treating fish poisoning ·CONTENTS V, DISCUSSION, 62 Need for lateral diffusion of knowledge between sciences Meagerness of physicians' knowledge of algae sad plankton. Early emphasis on seaweeds Shift to zooplankton, Recent prominence of phytoplankton, Nutritional utility of seaweed as fertilizer and animal provender. Importance of seaweed decivatives: alginates, Ratibiotics, anticoagulants. Evaluation of zooplankton as emergency ration and food supplement. Greater potential of phytoplankton Increased food needs of expanding population. Culture methods. "Aquaculture " Hydroponics "Mariculture" "Manuring the sea. Phytoplanktonic feeding and acceptability Algal cultures ta atomic submarines and space ships Allergic dermatitides Respiratory irritations Chronic sinusitis. Nephropathies Gastroenteritides Acute neurotoxic and myotoxic disorders Multiple sclerosis Muscular dystrophy. Amyotrophic lateral sclerosis Inadence in Chamorros Poliomyelits "Swayback" Hepatitis and cirrhosis, BIBLIOGRAPHY, 66 INDEX, 81 ·I. INTRODUCTION DURING THE PAST SEVERAL DECADES, there have appeared increasingly insistent reports on ligge and plankton—reports concerned chiefly with developing a new source of food for a world population rapidly outgrowing its arable land areas These artides, appearing in technical journals:-1! and in the public press, 18-2% have been authored primarily by botanists, autritionists, economic philosophers, and Lay saentific writers. The concern of these individuals seems amply justified, not merely by future census potentialities, but also by actual present shortages These include the frequent caloric fastines affecting many nations, as well as the qualitative autritional deficiences. The latter are pointedly exemplified by the dietary fat shortages in Europe during World War II, the cotorious hypovitaminoses of many Asians, and the recently highlighted proteza deficiency, ku ashrorkor,27,29 of the Africans. With the growing recognition of the importance of autritive and brochemical substances in the prevention and therapeusis of disease, it is remarkable how siagularly little attention has been paid by the medical press to algae and plankton As Tildenhas so aptly put it. "Here la America only a small group of college professors of botany are faciliar with marine plants. Phycology has been a science far removed from all other subjects. It is too difficult for the biochemist, the zoologist, or the physicist to attempt to work with forms of which be knows nothing is for physicians, the depth of tbeir igno1.37in the subject is abysm.d." The present monograph attempts to assemble in tangible for sach knowledge of algae and plankton as may be pertinent to medianeknowledge which presently is diffused in a wide variety of non-medical publications. Such a compilation of badkground dut3, of artent concepts, zod of future potential might conceivably serve as a stimulus for wider activity in this beld.

·II. DEFINITIONS AND CLASSIFICATIONS A. ALGAE ALGAE ARE THE SIMPLEST FORM of plant life.S0 The term "alga, -ae" derives directly from the Latin word meaning "seaweed, "31 but as used today encompasses many additional forms While the vast majority of algae are characteristically aquatic, some few species have become adapted to growth in soil and even in such exotic habitats as floating icebergs Some are unicellular, others multicellular, with great variations in cellular structure They may vary in length from less than one micron for the slimy growths, to hundreds of feet for the giant kelps 32 Unlike the higher plants, algae possess no true roots, stems, leaves, or seeds The basic metabolic processes of nutrition and reproduction are all localized in the individual cell, without the transport system of land plants The one feature common to all algae is the property of photosynthesis the utilization of light energy to convert inorganic to organic substances By this method, algae can convert carbon dioxide, water, nitrogen, and mineral substances into diverse types of proteins, carbohydrates, lipids and sterols, vitamins, and other metabolic complexes 33,84 Algae have been extensively studied during the past sixty years, and over 17,000 species have been described 35,88 Their classification has been varjed, unstandardized, and in many instances rather badly confused For many years, they were simply lumped into four principal groups based on gross coloration green, brown, red, blue-green 37 With increased understanding of pigments, of metabolic reactions, and of reproductive mechanisms, algac are today classified into sevea main Divisions, each subdivided into Classes, Orders, families, Genera, and Species3R.41; I CHLOROPHYTA, grass greea, unicellular, multicellular, a few macroscopic, cell wall of cellulose and pectins, mostly freshwater, some marine 11 EUGLENOPHYTA, grass green, unscellular; motile; lacking a cell wall, largely freshwater ·DEFINITIONS AND CLASSIFICATIONS 3 III. CHRYSOPHYTA: yellow greca to golden brown (xanthophylls and carotenes may mask chlorophylls); microscopis, mostly unicellular, cell wall containing silica (eg diatoms); largely freshwater, but many marine forms IV PYRROPHYTA: yellow green to dark brown, xanthophylls predominant, unicellular, motile, cellulose cell wall; many marine forms V. PHAEOPHYTA, olive green to dark brown; mostly macroscopic, cellulose and pectin cell wall, chiefly marine VI CYANOPHYTA. blue green, phycocyanio; multicellular, but usually microscopic; usually with gelatinous sheath, fresh water, marine, some terrestrial VII RHODOPHYTA: red, phycoerythrın, usually macroscopic, pectin and cellulose cell wall, mostly marine These seven Divisions include over a dozen Classes, about sixty Orders, hundreds of Families, and thousands of Genera However, only a few score Genera are of sufficient historical or current medical importance to merit individual attention Among these are the following: CHLOROPHYTA Chlorella, Scenedesmus, Ulva, Nutella, Chlamydomonas, Polysomella, Prolosheca, Sticbococcus, Cladophora, Oedogonium EUCLENOPHYTA Euglena CHRYSOPHYTA N:lzschia (diatoms) PYRROPHYTA Gymnodinium, Prorocentrum, Gonyaulax PHAEOPHYTA: Macrocystis, Nereocystis, Laminaria, Fucus CYANOPHYTA. Cbroococcus, Microcystis RHODOPHYTA, Porphyra, Gelidium, Corralina, Chondrus, Rbody. menia, Gigartina B PLANKTON The interrelationships of plankton and algae can be simply illustrated by the following schematic diagram PLANKTON ALGAE Zooplankcon Phytoplankton = Microscopic Algze Macroscopic Algae (Seaweed) ·4 THE ROLE OF ALGAE AND PLANKTON IN MEDICNB "Plankton," directly from the Greek, is defined by Websterl as "the passively loating or weakly swimming animal and plant life of a body of waten" The term appears to have been first used by Hensen in Germany about 1889, although Johannes Mueller collected microscopic organisms as early as 1845 by dragging a very fine net through the sea, 80 As used today, the term "plankton" generally encompasses the countless billions of minute animals and plants (excepting bacteria) which reside in water. This includes the algal growths seen Boating in fresh water ponds or lakes, and also the minute organisms in the sea. The latter provide, directly or indirectly, the basic food for all forms of aquatic life, from the smallest polyps to the largest whales, with each devouring some form smaller or weaker than itself Whipplenz describes it thus "Fish feed upon crustacea and insect larvae, the crustacea prey upon the rotifera and the protozoa, the rotifers and protozoa consume algae and bacteria, and finally algae nourish themselves by the absorption of soluble substances and gases provided in part by the decomposition of animal and vegetable matter brought about by bacteria " This biological sequence is perhaps more graphically characterized in the old Chinese proverbis. Big fish eat little fish Latele fish eat shrimp Shrimp cat mud And the "mud" comprises the multitudinous microscopic organisms at the base of this amazing nutritional pyramid It is apparent, then, that plankton exists in two basıc forms, zoo. plankton (animal) and phytoplankton (plant) ** i Zooplankton, consisting of shrimplike crustaceans, floating fish eggs and larvae; and larvae of crabs, molluscs, and a gum. ber of other small animals. Most important, in volume and edibility, are the copepods and krills. 2 Phytoplankton, consisting almost exclusively of microscopic algae, Main types are the diatoms and unicellular flagellates The term "ultraplankton” refers generally to the most minute forms (naonaplankton) of marine plankton, as differentiated from the bulk of phytoplanktonic organisms too large to be eaten by young larvae Cole's suggests substituting the term "bekistoplankton" for greater accuracy; he ·MACROSCOPIC ALGAE (SEAWEEDS) 5 defines it as including all those elements in plankton ten micra or less in diameter of cell body. Other descriptive terms sometimes used are "littoral" and "benthal," indicating growths along the shore of in deep waters; and "lumnetic," designatiog growths occuring in moving water of great practical importance is the fact that the quantities and composition of plankton vary tremendously with geographical atez, sea. son, temperature, light, water depth and currents, and nutrient concen trations. IL MACROSCOPIC ALGAE (SEAWEEDS) A. NUTRITIONAL ASPECTS THE USE OF MACROSCOPIC ALGAE (scaweeds and Occasional fresh water forms) as food extends far back into antiquity, most notably in the Oriental community 8 One of the earliest records of algae is found in the Chinese Book of Poetry, circa 800 BC, wherein "pondweed" and "duckweed" appear as edible delicacies worthy even of sacrifice to revered ancestors. References to the emblematic beauty of algae are made in the Book of History and in the Analects of Confucius 15 In the Far East, seaweed generically goes various namesti: in China, it is called hai t' sar; in Japan, nori; in Guarn, lumut, in Tahiti, rimu; and in Hawaii, limu, Hawaii has the largest variety of edible scaweeds in the world75 types but these are not all of choice quality The considerable regard the Japanese manifest for algal foods has been related to the fact that, apart from nee, they have very few vegetables Yet, in spite of the sumerous types of seaweed avadable, st is striking ·THE ROLE OF ALGAE AND PLANKTON IN MEDICINE 10) that seineeds provide less than one per cent of the total vegetable food of the Nipponese. Although scores of different seaweeds have been eaten in various parts of the world, only a few have enjoyed truly widespread consumption 18,19 Probably the most cosmopolitan seaweed, in geographical distribution and usage, has been the membranaceous Porphyra, commonly known as purple laver The Chinese used it mainly as a base for soups Called Amanori in Japan, it was at one time considered one of the most important food plants in that land, and was consumed chiefly as a salad It grew both naturally and under cultivation in surrounding waters, in fact, the most famous grounds for cultivation of Porphyra lenera were in Tokyo Bay itself The purple laver was also well known in the British Isles and in France, it went by such diversified names as sloke, slouk, slack, sloucawn, and marine sauce Its preparation was also more varied here The Europeans not only used it as a salad, but they also fried fat, or boiled it into a dark brown semi-fluid breakfast dish, or even baked it into a laver bread The British sometimes sealed the fresh plant hermetically for use as a food by the crews of whaling vessels In Alaska, the use of Porphyra as "sea lettuce" was common with the Indians On the Pacific coast, from Canada to Mexico, Porphyra perforata was the only local seaweed in common use Green laver, or Ulta lactuca, was known to the Orientals as Wakone, but was chiefly eaten in Scotland, either as a salad or as a soup The kelp Lammaria japonica was utilized primarily as a soup ingredient by the Japanese (who called it Kombu) and by the Alaskan Indiaas. Dulse (Rhodymenia palmala), also variously designated as water leaf, sea kale, dilluk, sol, and crannogh. was one of the favorite seaweed nutrients in Scotland, Ireland, and the northeastern United States in the nineteenth century It was chewed fresh or after drying, or used as a favoring, salad, or relish Outstanding among seaweeds finding a place in the cuisine were those of the Gelidiaceae famıly Granilania lichenoides, Gelidium corneum, and other closely related species were the prime source of agat, familiarly referred to as seaweed isenglass and kan ten ("cold weather") The latter, a pearly white, shiny, transparent, tasteless and odorless extract, was in. troduced by the Chinese to Japan to 1662 A,D. It was consumed mainly as a soup stock, summer jelly, dessert, or candy. Chondrus crispus (Rhodophyta) became highly popular in the production of jellies This alga was the source of Irish moss, also designated as carragheen, Dorset weed, ·MACROSCOPIC ALGAE (SEAWEEDS) 7 menu pearl moss, jelly moss, rock moss, gristle moss, and Tsunomata (among the Japanese) This was the seaweed so popular in Ireland and in the United States in the making of blanc mange Chondrus crispus and Laminaria saccharina were at one time combined on the coast of Armorica to make a jelly-like "pain des algues" or seaweed bread The general utilization of seaweeds as food in Europe-particularly in Scotland, Ireland, and Scandinavia diminished notably in the latter half of the nineteenth century With the food shortages incident to World War II, there was some revival of interest in macroscopic algae It has been reported that German occupation troops in Norway built two bakeries to make bread from dried algae which had been desalınated and ground 17 German technicians are also credited with having fabricated from Norwegian seaweed an edible casing for sausages weighing one. twenty-ninth as much as cellophane ? Prior to the war, the English made a custard powder from brown seaweed, the New Zealanders were market. ing a similar product after the war In 1944, Yatham*7 told of a botanical banquet some years earlier in Wisconsin that featured a dehydrated algal fried seaweed, puree of seaweed, coast seaweed, and devilled Seaweed It is clear that the employment of macroscopic algae as food was on a purely empirical basis---some of it founded on local custom, some on superstition, some simply on the belly-filling qualities of seaweed and its freshwater relatives As with many other traditional practices, modern science has tried to explain, post facto, why the ancients were actually right That many macroscopic algae are edible and not poisonous is amply evidenced by the survival of both the custom and the customer That these algae are also endowed with some desirable nutritional qualities is being gradually demonstrated The exact composition has been determined for only a relatively few species Even in these, marked variations are found from crop to crop and season to season 50,51 The main components appear to be a variety of carbohydrates, some proteins and fats, mineral salts, vitamins, and water Carbohydrates are present in large amounts, chrefly as cell wall components and as intracellular storage matter. 40 52-54 Most of them are of a heterocyclic six-membered pyranose structure Glucose and other reducing sugars are either absent, or present in only trace quantities Free pentoses are likewise of no quantitative significance Mannstol, a hexahydrıc alcohol derived by reduction from mannose or fructose, is found in amounts up to 30 per cent of dry weight in species of Phaeophyceae Mannitan (as ·8 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE mannitol anhydride), dulcitol and sorbitol (mannitol isomers), and Bori. doside (a glycerol-D-galactoside) all occur in various species of Rhodophyceae. Trehalose, a glucopyranoside, is present only in fresh water species of Rhodophyceae. The chief intracellular reserve products in many species are glucose polymers. Thus floridean starch is characteristic of the Rhodophyceae, it is composed exclusively of glucose residues, but differs from ordinary starch in containing a large proportion of 1:3 linkages and in being able to resist the effect of B-amylase The Phaeophyceae lack starch and allied substances Laminasin, another glucose polymer found in some Phaeophyceae, comprises up to 25 per cent of the plant on a dry weight basis. Cellulose is the main cell wall component the Phaeophyceae and Rhodophyceae, and it makes up from 2 .0 15 per cent of their dry weight. By chemical and radiological examination, it appears to be similar to the cellulose of higher plants Other related carbohydrate substances in algae include chitin, pectin, and hemicellulose Algın, a characteristic intracellular substance of the Phaeophyceae, forms about 25 per cent of the seaweed on a dried basis. It is a calcium-magnesium salt of algınıc acid, a polymer of B-D-mannuronic acid Xylan, a polysaccharide from which xylose is derived by hydrolysis, has been found in Rhodymensa palmala Other intracellular polysaccharides, composed residues esterified by sulfuric aad, include agar-agar, carragheenin, and fucoidin. Relatively little is known about the protein components of macroscopic algae, except that they are present in only small quantity and that they are minimally assimilable by human beings According to Cameron,6% analyses indicated 685 per cent protein in Gelidium corneum, and 3.49 to 5 82 per cent in various species of Laminaria Analyses of various kelp meals and feeds for animals (who may utilize contained protends better than humans) indicated levels of 7 04, 13.12, and 5.60 per cent, respectively At the 1952 International Seaweed Symposium, 66 the Youngs indicated that the proteins and peptides were difficult to extract from algae, with only 50 per cent being obtained even by strong alkalı. The nitrogen of various com son brown seaweeds consisted of volatile base nitrogen, nitrate nitrogen, and residual Kjeldahl nitrogen E. T. Young also stated that the a-amino-nitrogen after hydrolysis was always lower than the residual Kjeldahi nitrogen because of amino-acid decomposition The amino-acids found on acid hydrolysis of Laminaria saccharina and Alatia nodosum included glycine, alanine, valine, leucine, aspartic acid, serine, glutamic acid, threonine, proline, phenylalaninc, iysine, and argin- ·MACROSCOPIC ALGAE (SEAWEEDS) 9 ine. Some algae have also been shown to contain nucleic aads, and some various water-soluble peptides 67-69 The lipid content of macroscopic algae may vary from zero to 3 or 4 per cent of the fatty aads identified, the commonest are the unsaturated ones with 16, 18, and 20 carbons These are present in most algae, with the exception of Alaria crassifolia, which has over half its fatty aads in saturated forms The predominant algal fatty acid is the 16-carbon palmitic aad Besides these fats, there are varying small quantities of such lipord substances as photosynthetic pigments, sterols, and hydrocarbons 61-63 The mineral ash content varied from 50 to 38 9 per cent in a number of reported studies The water-soluble ash exceeded the insoluble frac. tion by as much as four or five times One investigation of kelps 64 listed identified substances and their percentage of the total ash as shown in Table 1 TABLET Sulfates Calcium oxide Magaesium oxide Phosphates (as P,0) 614-11 24% 335-446 199— 434 232-560 Potassium Sodium Chlorine Iodine Bromine 940--10 43% 10 51--13 57 27 62—32 59 006–009 Trace The seaweeds containing the most 10dine are Lammarsa digitata, Laminaria sascharma, Fucus vesiculosus, and Saccharina bulbosa The richest in bromine is Fucus serratus Vitamins in good quantity have been found in some varieties of seaweed investigated 47 Vitamin A has been demonstrated in the sea lettuce Ulva lactuca, Laminaria digitata, and Codrum tomentosum, as well as in ground meal prepared from Pacific coast kelps This vitamin is even more abundant in phytoplankton, which will be discussed later Freudenthal65 in 1949 conducted biological investigations on three species of algae from the coast of Denmark-Furcellaria fastigiata, Fucus serratus, and Fucus vesiculosus He reported that they possessed a growthpromoting and anti-xerophthalmic vitamin activity (in Wistar rats) ex. ceeding that of the best Danish summer butter He also claimed that feeding these algae prevented the epithelial dysplasia and rickets that developed in animals maintained on vitamin-deficent diets Freudenthal, incideatally, takes issue with the common belief that nickets is related to Vitamın D deficiency, he feels that the disease is "the retarded, in- ·10 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE hibited growth dynamics of cartilage and bone caused by a suboptimal intake of Vitamin A." The American brown kelps, Phaeophyta, studied extensively for their vitamin content, were found rich in thiamin, niacin, and riboflavio Norris and others found a thiamin concentration comparable to that of many fruits and vegetables in Alaria valida, various Laminaria species, Porphyra neteocystis, Rhodymenia pertusa, Ulva lactuca, and especially Porphyra perforata. The Vitamin B values of these algse were determined in fats first fed on deficient basal diets for about two weeks, when weight loss and cessation of growth took place. There appeared to be no loss in Vitamin B activity from drying the algae, nor was there any noted relationship to the depth of the water from which the algae were harvested Limu eleele and Lomu lipoa, edible Hawanan seaweeds, were found by Millerat to be poor sources of both Vitamins B and C. Only smajl amounts of Vitamin C were found in dulse (Rhodymenia palmata) How. ever, concentrations comparable to those in lemons have been reported in both purple and green laver Generally the spring and autumn harvests of seaweed are highest in ascorbic acid content Unlike the case with Vitamin B, seaweed collected from littoral zones is sicher in Vitamin C than that obtained from deeper waters Although ascorbic acid is usually destroyed by the drying preparatory to shipment of seaweed to market, coastal populations still derive significant quantities of the vitamin from eating the fresh algae 68,69 The occurrence in seaweed of the Vitamin D precursor, ergosterol, has also been recognized 70 Sauvageau reported marked anti-rachitic benefits in 25 human subjects fed various macroscopic algae 11 In 1952, Ericsons8 reported the presence of Vitamin B12, folic acid, and folinic acid in various brown and red seaweeds from the Baltic and North Seas. The B12 content of some of these was comparable to that in Jever The presence of a-tocopherol 10 seaweed was noted by Brown. 72 Norrises would appear to have been correct when he wrote in 1937. "Seaweeds contain no starch, and the complex carbohydrates which are present are not readily hydrolyzed by the human digestive enzymes. The value of (macroscopic) marine algae as food is due mainly to the in. organic salts present and the vitamins." It is possible, however, that broader chemical studies in the future may disclose more assimilable substances; and it is also conceivable that materials which m zitro presently appear inutile may be shown to be otherwise in t'rro. At any rate, the nutritional compositions and values of the macroscopic algae are in strik- ·MACROSCOPIC ALGAE (SEAWEEDS) 11 ing contrast with those of certain microscopic algae to be discussed in another section. While the direct nutritional utility in humans has been limited, the readily available hundreds of thousands of tons of seaweed might well be more effectively exploited in agriculture and industry Seaweed has been used as a fertilizer in the coastal portions of southeast Asia for many centuries. Even in the fourth century, Pallıdus mentioned the use of Seaweeds in western lands as a partial substitute for manure 73 To this day, particularly in Brittany, Normandy, and the Channel islands, algae are used in soil conservation Marrett** has attributed the small size and small bones of Jersey cow's to a calcum deficiency associated with the high 1odine content of the grass, a state resulting from the application of sea. weed fertilizer to the ground. The rodine content of algae is highest in the fall and lowest in the spring, with marine specimens containing much more than freshwater ones According to Lunde and Closs, 75 10dinewhether derived from seaweed fertilizer or meal causes an increase in the iodine content of the cow's milk, but this changes nesther the quantity nor the quality of the milk That the cattle on the Isle of Jersey are free from bovine tuberculosis, hoof and mouth disease, and brucellosis is attributed by some to the use of algal fertilizers, it seems more reason. able, however, to explain it on the simpler basis of relative isolation from contagion On the Pacfic coast of the United States, Desmarestra, a brown seaweed, is believed to be possonous to cattle, possibly because of the free sulfuric acid present in the plant 76 This has doubtless acted as a deterrent to the wider employment of other algae in that area Many forms of algae have been used as dietary supplements for livestock. Caesar, in his "Commentarses," tells of saving a crucial battle by feeding dred seaweed to his hungry horses when the supply of grain became exhausted a series of reports from 1745 to 1884 indicates that livestock in the northern British Isles and in the North Sea areas were encouraged to feed on algae washed up on the shore; in addition, the naves collected and mixed the seaweed with conventional fodder 65 Large aquatic plants reportedly have also been used as forage for cattle in Yugoslavia for many years * It was not until World War I, however, when the French were confronted with a shortage of oats and other grains, that any strong interest was manifested in the provender value of seaweed The military service then began studies on the nutrient effect of seaweeds on horses and cows.

·12 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE Sauvageau and Moreau?? reported that Fucus serratus and Laminati flexicaulis formed excellent animal aliment, Adrian78 found in animal experiments that seaweed cused an increase in weight proportional to the weight of the material ingested; seaweed was also credited with stimulating the assimilation of conventional nutrients In other studies at the Sorbonne, cats and chickens were well maintained on seaweed feedings Gloess79 found that macroscopic algae, when cleared of their potassium, bromide, and iodide salts, made a satisfactory replacement for oats in horses' diets, they could also be used as a supplement to standard rations for swine and poultry Beckmanso successfully maintained dogs and hens on a baked loaf containing a mixture of ground seaweed, iye, and potato flour Irish moss was also used to feed cows, calves, and pigs Freudenthalos in 1920 reported the use of seaweed as cattle fodder in Iceland During World War II, a dried seaweed meal was prepared in 2 factory in County Clare, Ireland, for use as a stock feed 81 Following a series of studies with pigs, it was concluded that the main value of the meal Tay in improving the total amount of digestible ration. Use of seaweed for fodder has also increased in Sweden as a result of scarcities occurring during the Second World War. To guard against adverse effects from seaweeds used in their entirety, it has been found advantageous to rinse the algae first in fresh water to leach out excess salts e7 In some of the feeding experiments reported. the animais required from several days to a week or more to accommodate their dietary habits to a predominantly seaweed ration. Undernourished animals often responded better to algal diets than did well nourished beasts From this it might be sucnised that seaweed supplements had overcome some existing deficiencies Palsson and Grimssogº have described in seaweed-fed lambs a syndrome demonstrated to be associated with a generalized spotty neural demyelınızation The clinical picture resembles that in the previously reported "swayback," a disease characterized by muscular weakness, incoordination, paralysis, and death *3 Palsson and Grimsson state that supplementing the diet of the ewes with copper during the latter part of pregnancy markedly decreased the inadence, morbidity, and mortality of this malady. They do not make dear why supplemental copper should be erther necessary or helpful, since the offending seaweeds showed a re latively high copper content. It is possible that something in the seaweed (or some contaminant on it) may have interfered with the proper utilization of copper, or perhaps markedly increased the need for it. In en ·MACROSCOPIC ALGAE (SCAWEEDS) 13 interesting comparable situation, Dunlop 19 Western Scotland administered single doses of copper sulfate to lactating cows fed on seaweed and obtained increased butterfat production within a few days.84 B. MEDICINAL USES .. The medicinal purposes to which algae have been put are indeed diverse 45,47,71,85-94 Their origins are often clouded in mesty legend and hearsay, their rationale generally empirical The Chinese credit the therapeutic qualities of algae to Shen Nung, the mythical First Farmer or Father of Husbandry and Medicine, who supposedly lived about 3000 BC. The Materia Medica of the Chinese in the eighth century AD described one alga thus. "Whole plant is officinal Taste butter and salt Nature cold Nonpoisonous . ... the bar isao grows in Tung Hat (Shantung) in ponds and marshes. It is gathered on the seventh day of the seventh month and dried in the sun." of the seaweed Porphyra coccinea, the Chinese Materia Medica said. "This algaf plant is a sort of laver which is green in the fresh state and purple when dry It grows on the seashore and the Fukicnese. · press it into cakes It is not poisonous, but when taken in excess it produces colicky pains, flatulence, and eructations It is recommended in diseases of the throat, especially goiter" A related laver of the Porphyra grouping was the source of a preparation called k'unpu, utilized for the relief of dropsy A similarly named viscous solution, kwanpu or hai lai, derived from a Laminaria, was used by the Chinese fot menstrual difficulties They also used a scaweed concoction known as lung-she-esar for abscesses and cancer. For many years Chinese gourmets have used the nests of some passerine birds (such as the sca swift) to make an exotic soup with reported aphrodisiac powers, and millions of these nests have been imported from Borneo Although they were once thought to be made of a gelatinous seaweed, Gracilaria spinosa, it is now known that the main constituent is dried avian salivary juice! In upper India, an algal prepartion known as gillur-ka-putra served to treat bronchococle Sugar wrack, Laminaria saccharına, was here con sidered useful against goiter, while in the Himalayas it was more prosai. cally employed to treat syphilis. The ancient Polynesians used filamentous algac to make poultices for bruises, cuts, and inflammations The H2%311ans made watery extracts of Centroceros and Hypnta nidifica for constipation ·THE ROLE OF ALGAE AND PLANKTON IN MEDIONE and other gastro-intestinal disturbances. The primitive Maori tribesmen, living in close contact with their domestic animals, frequently acquired their scabies; to treat this, they used Durvillea ("rimuroa"). The latter was also fermented with an extract of the poisonous Tutu shrub to make a cathartic. The Indians of Sitka, Alaska, devised an unique mechanical nostrum for earache Utilizing the tube-like character of the stipe of the bull kelp, they placed the thin end in the victim's ear and the bulb on a hot wet stone, thereby allowing steam to enter the afflicted external auditory Canal' South American Indians are reported to have valued Sargassum baccifer to cure goiter and renal disorders, just as many other primitive tribes considered bladderwrack helpful in inflammations, spruins, and rheumatism In the West, algae were exploited somewhat as in the Orient, but to a much lesser extent The Latin poets Virgil and Horace (first century BC.) held algae in very low esteem Nevertheless the Romans must have attributed to certain algae the property of healing sores; indeed the name Conferene (Class Chlorophyceae) derives directly from the Latin word "confervere," meanmg "to solder" In addition, Pliny recommended certain seaweeds for gout, which the Romans knew only too well Descondes, a Greek physician of the first century AD., recognized absorbent properties in white or medicinal corraline, Muscus corralius, The powder from this alga was highly valued. It was prescribed as an external absorbent for burns, skin abrasions, rashes, and scurvy; and as an internal absorbent for chronic gastritis, heartburn, and diarrheas resulting from hepatic and intestinal disorders. The American Indians, according to Chase, 43 did not believe in the use of salt on food It seems that the Iroquois, having occasionally feasted on the white intruder, learned that he tasted saftier than the red man Stace salt was "white man's style," they avoided using it, firm in the belief that salt would make their hair turn gray and their toes turn up before their time. However, by taking "ses lettuce" or laver each spring, they supplied directly from nature the salt needs of their bodies. The saline tastes of the Iroquois prior to Columbus (or Lef Ericson's) visit are not recorded. In cases of goiter and scrofula, ancient medical men were known to prescribe clane sponges, watercress, and similar substances which condensed and fixed marine clements The Chinese Pentsio recommended all the medianal algae in gostes therapy. The fact that goiter is practically ·MACROSCOPIC ALGAE (SEAWEEDS) 15 unknown amongst the maritime Chinese and Japanese is doubtless due to the abundance of iodine in their seaweed dietary supplements and in their fish foods In striking contrast is the noted prevalence of goiter in Alpine inhabitants, who eat no marine algae, and whose agricultural soil has been leached of iodine by ancient glaciers 95,98 In the Andean regions of South America, the natives indeed learned to chew "goiter sticks"—the stipes or stems of species of Phyllogigan—to counter the high incidence of thyroid enlargement. Likewise because of their iodine content, the therapeutic use of oarweeds and wracks was common. Most popular were Laminaria digitata and saccharina and Fucus vesiculosus, the latter being prepared as an infusion Another favored way of administer. ing iodine was in the form of kelp pills, ash, or charcoal Charcoal kelp was used under the name "Aethiops vegetabilis" for goitrous and scrofulous swellings Coudert, in Geneva, was among the first to utilize lodine by itself as a therapeutic agent. In time it was found more economical to derive iodine from the saltpeter beds of Chile, and soon thereafter the great kelp industrias of northern and western Europe began their rapid decline One of the earliest therapeutic applications of algae by the Chinese and Japanese was that of the derivative agar in gastrointestinal disorders In cases of fever attributed to stomach conditions, their monks recommended Gelidium boiled into a jelly and sprinkled with ginger and sugar For relief from disorders caused by extreme heat, an agar milk paste flavored with sugar and vinegar was given This preparation was particularly favored for children Gracılara lichenoides was also utilized as a demulcent in intestinal and bladder difficulties Ia more recent times, the U S. Dispensatory recommended agar as a laxative, eaten in small pieces like cereal with sugar and cream, or else in chocolate-coated form its value as a laxative is due to its colloidal property of absorbing and holding water without being digested This attribute takes it desirable as intestinal bulk or roughage 97,88 Agar also decreases gastric emptying tune The jelly extract of Irish moss, like agar, was advised in general irritability of the alimentary tract During the past century it was prescribed for gastric ulcer, as well as for diarrhea and dysentery On the eastern Asiastic coast, a red algz, Digenea nmplex, was dsed and sold by Oriental apothecaries; its extract, "helminol," has been em. played in the management of ascariasis and oxyuriasis.** In Chine, a combination of algae, including an Enteromorpba, a Chordana, and seven ·16 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE Floridees, was favored for its vermifugal virtues; it is thus not too surpris. ing that the Chinese credited practically all marine algae with such powers. In Europe, until the end of the eighteenth century, vermifuges prepared from the calcareous algae Lammaria officinal and ruben enjoyed high public approval. In 1775, the Greek physician Stephanopoli discovered the anthelmintic properties of the small red tufted alga Alsidium belmin. thocanton, found at low tide along the rocky Corsican shores. Additional seaweeds of help in deworming include Hypnea musciformis (particularly in Turkey and Greece), two species of Chondria, the green fresh-water alga Rbrzoclonum t'orulare, and dulse Employed by the Maoris of New Zealand was yet another vermifuge, prepared from the bull kelp Durvillex Irish moss, the red seaweed that flourished in Carragheen, was used -after cleansing, cuting, and bleaching—as a demulcent for coughs Its mucus-forming qualities made it popular in the nineteenth century for pulmonary desorders generally, and for "consumption" particularly. The purest Carragheen preparation, Electum albissimum, was used for this purpose It was sometimes prepared as a liver oil emulsion. Carragheen was also used occasionally to disguise the taste of better drugs A related extract of Chondrus crispus found usage in World War I in easing the throat irritations of soldiers who had been gassed Carragheen has also been denatured with whiskey and used liberally for coughs in some Hibernian bars in New York 47 A distinctive French preparation, to serve in place of linseed meal poultices, was cotton wool soaked in a Carragheen concoction, then dried Other seaweeds considered useful in lung diseases and scrofula included Gelıdrum cartilagineum (in Japan), Dictyopteris polypodiodes and Sulophora rhizoides (in the Mediterranean area), and Laminaria saccharını. Irish moss also served as a demulcent in diarrhea and in irritations of the urinary tract in the middle of the eighteenth century, Scottish peasants were said to chew dulse to induce sweating in states of pyrexia The bladderwrack Fucus vesiculosus also enjoyed a role in certain concoctions and "slimming teas" in the management of obesity 100 Its therapeutic merit might be attributed to its iodine content and its effect upon the thyroid gland Agar, by virtue of its bulk and relative in. absorbability, also served in the treatment of problems of overweight. In the surgical field, stipes or "stem pieces" of Laminaria cloustoni, known as Stiprtes lamınarsa, w'ere utilized to widen fistulas and wound openings because of their ability to swell markedly following moistening These stipes were also used in distending the uterine neck in labor.

·MACROSCOPIC ALGAE (SEAWEEDS) 17 Despite the many old traditional uses of algae in therapeusis, the early twentieth century German medical lists included only Carragheen, Agar-agar, Laminaria, and Fucus. The direct medicinal uses of macroscopic algae have clearly undergone a marked decline Certain extracts of seaweed are, however, of current significance in medical and para-medical fields The most important of these are agar, algin, and carragheenın They are extracted from algae by either water or alkali Though they are hydrophilic colloids, they have often been erroneously referred to as seaweed "gums" The term “phycocolloid" has been introduced as a more correct designation Aside from the medical areas, these extracts are also of value in the drug, food, and cosmetic industries because of their propertres as gelling, suspending, emulsifying, thickening, and body-producing agents Shabin and Shapiro101 in Russia have recently reported success in treating trichomonal vaginitis with a Laminaria derivative Monceaux 102 has enthusiastically advocated the use of the cellular plasma of marine algae for a diversity of clinical conditions He advises oral ingestion for systemic diseases and deficiencies, and local application for dermatological afflictions Monceaux values seaweeds so highly because of the naturalistic premnise that they contain many of the therapeutically efficacious minerals and trace elements present in sea water Since its utilization by Koch48 in 1881 for the culture and isolation of pathogenic microorganisms, agar has become an important adjunct of every hospital and bacteriological research laboratory This dried and bleached gelatinous extract of the red seaweed Gelidiaceae es produced primarily by the Japanese; the British derive some agar from Gigartina It possesses the desirable property of absorbing water easily and swelling up to a considerable extent Only when heated to boiling does it dissolve, upon cooling it reverts to a somewhat colorless translucent gel Agar is good as a solid culture medium particularly because it is relatively inert, not digested by most bacteria It has also been used as a sealer and coating for pills, as well as a vehicle for lactic and administered in intestinal infections Algin is the common name for derivatives of alginic acid, such as the sodrum, potassium, and ammonium salts and the propylene glycol estes 103 The discovery of alginc acid and the production of algin therefrom is credited to Stanford in 1885 *7 Alginic acid is the hydrophilic colloidal polymer of anhydro-B-D-mannuronic aad It is extracted from the brown algae, especially the giant kelp Macrocystis pyrifera, Laminaria digitala ·18 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE and hyperbores 104 The action of therapeutic agents can be modified by combining them with algia, which is free of nitrogen and possesses a low allergeniaty It lends itself well to incorporation into medications to be injected, since algınıc solutions are clear and colorless, soluble in hot or cold water, and retentive of their smoothly flowing nature throughout wide temperature ranges By varying the algia concentration only slightly, it is possible to control not only the viscosity, but also the rate of absorp tion and assimilation of medication 105 One example is the delayed-action algin-epinephrine mixture being used in allergic disorders 108 Another is its use in retarding the absorption of penicillin and various myan drugs 107 In this connection, it is interesting to note that Suzuk,109 has been able to produce anaphylaxis in guinea pigs by intraperitoneal injections of saline extracts of seaweeds common in the Japanese diet Ouer, on the other hand, has reported minimal human sensitivity on intradermal injection of propylene glycol algınate in 50 allergic and 50 non-allergic individuals 108 Differing results have also been reported on toxicity and other adverse reactions Mellis and Reed 110 noted that sodium alginate does not interfere with calcium absorption in healthy human adults Gill and Duncan111 likewise observed no electrolytic alterations, nor any significant gastrointestinal disturbances Nilson and Wagner 112 fed sodium alginate and propylene glycol alginate to dogs, mice, and chicks; they considered these substances harmless and nutritionally wholesome, but bulky" Chenoweth 112 reported a high degree of toxicity in cats On intraperitoneal or intravenous injection of sodium alginate, he found large antemortem intracardiac clots, as well as frequent indications of injury to the brain, liver and kidney He asserted, too, that sodium alignate thus givena elevates the erythrocyte sedimentation rate just as do the various gums Chenoweth also found much toxicity with the aitrate derivative, which also greatly prolongs blood clotting time The dialdehyde derivative of algınıc aad was somewhat less toxic Solandt114 reported sodium alginate toxic for rabbits, while Frantz found the same with rats 113 Blaine116 considered that alginates make a good absorbable surgical substance, capable of being used as a gel, foam, or gauze with a minimum of local tissue reaction. He also used lamunarın as a surgical powder in place of talcum 117 Some of these disagreements in reported results may be due to product impurities or to differences in test animals Others may be caused by variation in the source of materials, thus, Solandt and Franz both used Laniinaria as the origin of their alginates, whereas Ouer used Microcystis ·MACROSCOPIC ALGAE (SEAWEEDS) 19 Because alginates are not subject to breakdown by gastric juices, they have been used successfully as a coating for medications designed to have their effective activity in the intestine. Alginates have also seen much application as hydroabsorbent agents in the management of constipation Berger and his colleagues118 considered the water-absorbing and retaining properties of sodium alginate superior to those of natural guns, methyl. cellulose, and carboxymethylcellulose. They also found the acid-binding properties of sodium and calcium alginate comparable to those of carboxymethylcellulose. Gill and Duncan 11 thought that alginates were less effective in this respect than the carboxyl resins Mulinos119 tested the sodium and calcium salts of alginic acid in 60 humans with colonic consti. pation He found them effective, and thought that, even though the algi. nates swelled minimally in water or gastric juice, they increased their original bulk 25 to 35 times in the alkalinity of intestinal secretions Ludwig and his associates 120 compared the alginates favorably with the carboxylic type of cation exchange resins in their effect on intestınal sodium and potassium According to Feldman, 121 human subjects tolerated 45 gm of algınic acid quite well It acted as a mild laxative, and it increased the fecal sodium and potassium to 16-20 per cent of that seen with the common cation exchange resins. Since 1937, when Sears introduced the "reversible hydrocolloid technic" in the preparation of dentures, the use of alginates in dentrstry has increased notably.122 With this method, the dentist is able to secure with one impression reproductions of multiple cavity preparations, to reproduce undercuts without rupture or distortion, and to save chair time for himself and the patient 123 in a similar fashion, alginates are used for fixed bridges 124 and dental hemostasis, 126-128 as well as for various types of medical moulages 123 Alginates have also been put to the following medical uses 1. As an endaural wound dressiog 130 2 As a mold for applying skin grafts after exenteration of the orbit 131 3 As a hemostatic in brain and thoracic surgery, 182.144 especially to close the bronchial tree in pulmonary tuberculosis, and in extrapleural pneumothorax 4 As a decontaminant and treatment in mustard gis poisoning and burns. 185 5 As swabs in the bacteriological examination of eating utensils, ·20 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE since alginate "wools" adsorb more bacteria on wiping than does cotton. 136 6. As a replacement for tragacanth and other natural gums in the manufacture of lubricating jellies 104 These diverse applications of the alginates utilize only a fraction of the phycocolloid production of the United States Most of it is serving the dairy industry By conservative estimate, for example, over half the ice cream made is stabilized with algin to provide smoothness of texture and body Carragheenin has found some utility as a stabilizer in bakery products. 49 An antibiotic property has been attributed to some algal substances The natives of some Pacific islands wrap their fish catches in fronds of certain local seaweeds to guard against rapid decomposition,"? but no speafic antibotic has yet been identified here The antiviral activity of the algal polysaccharides sodrum alginate, lamiaafia, and fucoidın has been tested in chick embryos 107 While no toxic effect upon the embryos was noted, neither was there any upon the viruses. Pratt, Mautner, and their associates 137 in 1951 demonstrated an in vitro inhibition of the growth of Staphylococcus aureus, E coli, and P pyocyaneus by extracts of several species of marine algae collected in California during the spring and fall. The same investigators138 in 1953 obtained in vitro inhibition of the growth of several Gram positive and Gram negative pathogens with ether extracts of the red alga Rhodomela lanıx They assert that the effectiveness of these algal substances us not due to their sodine content, but may be on the basis of some brominated phenolic compound The anticoagulant properties of algal substances have also received some attention Following the identification of heparin as a mucoitin polysulfuric aad ester, and the finding that blood coagulation was inhibited by polysaccharides artificially esterified with sulfuric acid, Elsner 189 in 1937 began testing natural polysaccharide sulfuric acid esters for anticoagulant effect Considerable anticoagulant activity was found in watery extracts of the red algae Gelidium, Chondrus crispus, Irides lamineroides, and Delesserie sanguinea. The aqueous extracts of the brown algse Chor daria flagellsformis and Laminania digitala also had anticoagulant powers, but of a lesser degree 110 The strength of these extracts was found to be dependent upon such variables as seasonal fluctuations, length and extent of drying of the algal fronds, the age of the extraction, and the amount of heat to which the substances were subjected. Meunier, Molho, and their ·MACROSCOPIC ALGAE (SEAWEEDS) 21 Co-workers in France reported similar studies 111,112 with alginic derivatives in 1949 and 1951. Burt, at the Institute for Seaweed Research in Edinburgh, 107 in 1952 reported some heparin-like activity in vitro for laminarın sulfate; others there reported even greater activity the following year for sodium laminarin sulfate used in rabbits and rats. These extracts contained 061 to 2.15 sulfate groups per glucose unit. Even though the evolution of the coumarın compounds has progressed tremendously and overshad. owed most others, it is important to know the anticoagulant potentials of algae Not only might it be possible to perfect better agents in this category, but it could also help assess possible undesirable effects from eating certain algae Blaine, working in Scotland and in Jamaica, studied the potentialıtıes of laminarin as a blood plasma substitute. He used it successfully on dogs in shock, small test doses in humans produced no adverse reactions Wilkerson likewise reported some success in administering lamınarın to rabbits in shock 107 With the advent of the atomic age, concern has been manifested over radioactivity of marine substances exposed to test explosions of nuclear weapons. It has been found that elements of atomic weights 83 to 115, and 127 to 154, are produced in greatest concentration by nuclear fission Studies are reported on the way on the uptake by seaweeds of radioactive lodine, Caesium, Rubidium, Cobalt, and Selenium 107,143 The potential effects of radioactivity on marine organisms are only too obvious. C. ALISCELLANEOUS USES Not only have algae been useful in themselves, but they have also supplied some exotic products which have been highly prized by certain peoples For example, in the distant Siberian province of Kamchatka, the natives employed dulse in the preparation of a potent alcoholic drink Similarly the Alaskan Indians, in making an alcoholic concoction called "hoochenoo" (sic!), used the hollow stipes of Nereocystis--not for its intrinsic content, but for its mechanical attributes as a worm condenser to cool the distilled spirits 144 The Irish, like the Alaskan natives, used dulse-washed, dried, then hand-rolled--as a sort of chewing tobacco During World War II, considerable quantities of dried karengo, a species of Porphyra, were used by Maori soldiers serving in the Middle East, chewing Karengo was asserted to be more thirst-quenching than chewing ·22 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE gum On the island of Grand Manon, off the coast of New Brunswicki, the reddish-purple leaves of dulse are still considered a great chiewing delicacy 145 Pling the Eldes (first century B.C.) considered sezweeds valuable as dyes, as did the Roman ladies who used thema for cosmetic purposes. They availed themselves of a rouge extracted from the Fucus wracks (the word Fucus is in fact derived from the Latin word for touge.) Even today, a similar extract mixed with fish oil is in vogue with Kamchatkan women wanting to redden their cheeks 116 Since the seventeenth century, a seaweed glue called "funori" has been derived by the Japanese from the sed alga Gloiopeltis furcata. Its main utility has been as an adhesive and as a sizing for paper, fiber, or doth. In Europe, "kelp"--the burat ash of certain browo seaweeds-Was used in glazing pottery and manufacturing glass, as well as a source of soda.147 The production of "kelp" was a big industry for a whole ceatury, until the development of the saltpeter beds in Chile made it uneconomic. IV. PLANKTON A. ZOOPLANKTON AS PREVIOUSLY INDICATED, zooplankton is that variety of plankton which consists of minute anumal forms 118-160 It is collectible in a #2 bolting silk net, and it is thereby separable from the main phytoplanktonic organismos (diztons and dinoflagellates), which pass readily through such a net. The chief components of zooplankton are the copepods, prawns of krills, and fish eggs and larvae.151-154 The most significant, in both bulk and nutritional value, are usually the copepods, which feed on phy ·PLANKTON 23 toplankton 135 There have been about 750 species of copepods identified, averaging 20 to 3.0 mm in length The Calanus finmarchicus, for ex ample, is well recognized as an important food for many fish, and is very abundant during the summer around North Atlantic shores The colder northern seas generally contain the largest quantities of copepods, though the Paafic and tropical water offer more varieties The mysids and the euphasiids (krills) are the most noteworthy of the higher crustaceans, they comprise some 400 species They are much farger than the copepods, the euphasirds occasionally reaching a length of 5.0 cm. As is commonly the case, these larger organisms live at deeper levels and come to the surface only during the spawning season They are then so plentiful in Antarctic waters that the blue and fin whales feeding on them can attain a weight of sixty to seventy tons in two years 3 G O Sars is ated by C E Lu30 as having early discovered that most fish have planktonic eggs and larvae Even herring, which lay their eggs on the ocean bed, contribute the planktonic amalgam when their larvae leave ocean floor and float to the surface The use of zooplankton as a nutrient has been discussed with increasing frequency since the latter part of the nineteenth century Although both freshwater and marine zooplankton are potential sources of food, the oceanic varieties have excited greater interest 188-158 A search of the recent literature on zooplankton reveals a repetitive pattern of reference to the same group of articles One of the better summaries is that of Robinson and Bajkop259 in there 1947 survey of marine plankton for the United States Army Air Force W A Herdman160 is frequently credited with the first publication on the eating of zooplankton His single paragraph communication to Nature in 1891 related that eight yachtsmen off the coast of Norway breakfasted well on cooked red copepods, probably Calenus finmarchicus, it also mentioned that the Prince of Monaco had already considered the nutrient potentialities of copepods for shipwrecked sailors Yet even in 1888 Greelyet had reported that men in his 1882 Arctic Expedition ate shrimplike crustaceans so small that 700 weighed only an ounce In 1908, J Johastone162 reported Brand's chemical analysis in 1898 of dried copepod plankton as follows: protein 59 per cent, carbohydrate 20 per cent, fat 7 per cent, chitin 4 7 per cent and ash 94 per cent Shortly before the first World War, Nansen 163 reemphasized the potential value of plankton to castaways In 1939, Seidenfaden 164 wrote that sailors often considered prawas a delicacy.

·24 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE In 1939, too, George L. Clarke, 2 of the Woods Hole Oceanographic Institute, wrote in Saence that he had heard the Germans in Heligoland were studying the use of plankton as a food supplement; he said they considered zooplankton nutritionally equivalent to the best meat, and phytoplankton to rye flour Clarke hypothesized that of all the organic matter in marine zooplankton were assimilable, it might yield four calories per gram Thus 750 gm plankton would be required to produce the 3000 calories needed daily to support a man If, as he figured, one cubic meter of water yields only 0 1 gm dry plankton, then 7500 cubic meters of sea water would have to be filtered to provide daily sustenance. This represents a heroic operation, but Clarke suggested using nets in the tidal flow's of estuarial or interinsular waters. The war situation spurred further interest. In 1940, W. A. Jaschnovios spoke in Russia of the desirability and the feasibility of using planktonic foods His opinion was not based on theory alone. Between 1921 and 1937 he had studied the distribution and concentration of zooplankton in the Barents, White, Kara, Laptev, East Siberian and Chukotsk Seas, as well as in the adjacent polar areas. He examined 2500 samples and made 1300 quantitative determinations From these he concluded that 200- plankton contained over 90 per cent copepods, and that Calonus fin. marchicus made up about 95 per cent of the latter Like others, Jaschnov also found the highest concentration of zooplankton in the autumn He calculated that there was an average of 65 tons of Calanus available annually per square kilometer of sea surface, equivalent to a bromass of 43 tons The total biomass available yearly from the northern seas of the USSR was estimated to surpass 50 million tons-40 million tons from the Barents Sea, 5 million from the Kara Sea, 3 million from the Laptev Sea, 2 million from the East Sibenan Sea, 1 million from the Chukotsk Sea, and one-half million from the White Sea Berutski188 also noted seasonal variations of copepods in the White Lake in Russia Wiborg 167 classified many of the copepods in the Oslo fiord In 1941 Sir John G Kerr suggested in the House of Commons and in the London Times that marine plankton be investigated as a food source 168 Indeed, soon thereafter, Professor A C. Hardya enthusiastically urged in Nature that Clarke's tidal flow filtrations be tried in the planktonsich sea lochs of western Scotland. In the following year, Hardy28s also suggested freshwater plankton as a source of food In 1493, Judyt re. ported in Science on the good aquatic food potentialities of Wisconsin lakes. That these war-inspired enthusiasms might be practical was further ·PLANKTON 23 attested to by Dr. S P. Chu, who wrote to Clarke la 1946 from Yunnan that Chinese peasants had long used marine plankton as food 117 In the same year, too, Sir Hubert Wilkias wrote Bajkov that he had eaten plankton in the Antarctic with no ill effects In 1945, Knowlton and Irving, 170 of the Acro-Medical Laboratory, studied the effect of eating plankton upon the water requirements of men on life rafts Thear investigation was basically a physiolagic analysis of the nutrients of copepods caught in Buzzard's Bay (Massachusetts) by Bajkov, On the basis of determinations made by Dr A. Butler, the chioride concentration of the plankton was calculated as about 037 per cent, as com pared wsth 1.9 per cent in sea water. The protein content was 51 per cent, while the fat was estiraated to vary from 10 to 30 per cent. If 800 ml drinking water art available daily (from solaz still, desalination kit, and captured rain water), they concluded that orer 300 calones could be desved from plankton before the limsts of salt excretion were reached With this amount of planktonic protein metabolized, the urmaty nitrogen produced would be about 124 gm, also a serious consideration when water intake ss lumusted Robinson and Baykov mentioned in their 1947 report168 that about twenty of their furan subjects ate 50 to 100 gm plankton at one real without all effect, and that one individual even consumed a full pound without harm The plankton as these experiments came from Atlantic, Paabic, and Gulf of Mexico waters In most instances, the plankton was considered quste palatable The raw planktoa tasted like raw oysters, while the cooked material resembled cooked shrimp Varsations in the smell of raw plankton were produced by admixtures of certain species of diatoms and diooflagellates in the same report, Robinson and Bajkov give the results of feeding plankton to rats Knowsng that Atlantic plankton could be digested and assiradlated by rats (studies at Woods Hole Manne Laboratory, 1945), they successfully repeated the procedure with fresh water plankton from an Ohio lake, and also proved that it had a speasc growth-produong value Similar results were then obtained with Pacific planktos, Comparative analyses of several types of plankton vere listed (Table II) It is apparent that the composition of plankton varies with geography, demate, and the predominant species of organisms Water comprises 75 to As per cent of manne copepod piankton On a dry weight basis, 45 to 60 per cent may be proteia; 4 to 31 pxet cent, fat, 18 to 23 per cent, car. bohydrate, and os to 20 per cent, ash Chatun is usually present in less ·26 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE TABLE II Proteia Fat Carbohydrace Total Ash. Chitia Atlantic Pacific (Robinson sad Bajkov) 52 6% 490% 42 70 150 11.0 28.2 32.2 Barents Sea Arctic (Vicogradov) 590% 62 65% 7.0 18.30 200 140 3 49 than 5 per cent concentration in copepods, but is more abundant in higher crustaceans. It forms the exoskeleton of copepods. Although chutin is not digestible, it passes through the gastro-intestinal tract harmlessly as bulk That oceanic plankton can truly be an aid to the survival of drifting castaways has been spectacularly highlighted recently by the Kon-Tik1,171 the Heretique, 172 and the Seven Little Sisters173 adventures. Thor Heyerdahl and five companions left Callao, Peru, on April 29, 1947, on the raft Kon-Tiki, seeking to prove that pre-lacan Indians could well have settled the Polynesian Islands after passively drifting westward on prevailing ocean cutrents They landed 102 days later, physically none the worse, on the Raroia Atoll in the Tuarnotu Archipelago Throughout the 4300 mile journey, they collected plankton in a conical silk mesh net, often getting several pounds of porridge in a few hours The catch was greatest at night and when the sea was cold Heyerdah! vividly described the myriads of planktonic organisms netted, stressing especially their colorations and nocturnal fluorescence Four of the six men really liked plankton The taste varied with the composition of the day's catch; it ranged from that of shrimp paste to that of lobster, crab, caviar, and oysters. Dr. Alain Bombard, a French physician and sailing enthusiast, became obsessed with the conviction that victims of shipwreck would have a better chance for survival of they became better acquainted with the resources the sea itself offered. To dramatize his thesis, he sailed his 15-foot rubber raft, the Heretique, alone from Tangier across the South Atlantic to the Barbados in 1992. Although the 65-day voyage was physically somewhat trumatic, Bombard sustained himself on plankton, fish, water derived from fish, occasional rain water, and small amounts of sea water. Even more recently, William Willis, a sailor from the Bronx, fol. loved the Kon-Tiki route as the solitary passenger on the balsa raft the Seren Little Sisters. He left Callio, Peru, on June 22, 1954, and arrived ·PLANKTON 27 115 days later at Samoa, 6000 miles away. Preliminary reports indicate that he, too, sustained himself to a fair extent on plankton B. PHYTOPLANKTON (MICROSCOPIC ALGAE) 1 General Nature Phytoplankton or "plant plankton" conists chiefly of microscopic algae, which, with few exceptions, are aquatic organisms These microscopic growths comprise mainly the Divisions Chlorophyta, Eaglenophyta, Chrysophyta, Pyrrophyta, and Cyanophyta Though thousands of species are represented, the diatoms and unicellular flagellates are predominant They thrive in both fresh and salt water, however, a given species is specific for the one or the other Algae are the only plants that can dourish in the sea They make available the vast amounts of dissolved minerals and other nutrients, thus Decrtic plankton (near the coasts) is richer because of the higher concentrations of salts As previously indicated, phytoplankton provides-- directly or indirectly—the basic food for all the ascending hierarchy of marine animals Freshwater algae play a somewhat similar role in their own habitat The total quantity of phytoplankton perforce exceeds that of zooplankton However, effective collection of phytoplankton is very difficult because the individual organisms are so small As Burlewhl has so strikingly put it, one quart of moderately thin suspension of Chlorella Pyrenosdosa contains twenty billion cells, a number equivalent to eight times the world's human population' The color of the sea directly reflects the particulate matter therein 174 When there is little marine growth, the water is deep blue, since the light penetrates to deep levels, and the yellow and red rays are absorbed When planktonic growth is abundant, the water assumes the yellow, green, or brown shades of the microscopic algae. Seasonal abundance of tain reddish algal growths may produce the "red water" described since ancient times; to this may also be credited the coloration of the Red Sea and the Vermilion Sea The nocturnal glow of some seas is attributable to the phosphorescence of certain planktoarc organisms Reference is made in the literature to microscopic algal studies as carly as 1779, Priestly175 and Ingen-Housz176 described experiments on ·28 THE ROLE OF ALGAB AND PLANKTON IN MEDICINE the photosynthetic gas exchanges of the simple algae which contaminated their laboratory equipment. However, not until the latter part of the nineteenth century was there any strong interest exhibited in the precise nature of microalgae. These investigations were in three general directions: 1. The chemical constituents of algae (carbohydrates, pigments, and -more recently—the nitrogenous and lipid components). 2 Short-term brochemical experiments, eg, photosynthesis 3. The growing of algae in culture Cohen"?? first cultured algae in soil in the middle of the nineteenth century, while Famıntzin178 in 1871 was the first to raise them in aqueous medium. The initial extensive employment of microscopic algse was by Engelmann179 in 1883, in his work on photosynthesis. In 1890, Betjer. inck,180 a Dutch bacteriologist, described the first isolation of algae in pure culture; he obtained Chlorella vulgaris and Scenedesmus arutus frec from bacteria and all other organisms. In 1896 Molisch181 published his work showing the mineral requirements of simple algae to be similar to those of higher plants. The study of Willstätter and Stol[182 in 1913 pioneered our knowledge of algal pigments. In 1919 Warburg188 recommended Chlorella as a system in which photospathesis might be very simply studied, so thorough was he that in 1931 he received the Nobel prize for his observations on cellular respiration, much of them made on Chlorella viridans Since then, Chlorella has become a standard organisma for studies on photosynthesis It has also pointed the way for many diverse uses of the byproducts of photosynthesis, as will be discussed later By 1920 many algae had been isolated pure culture. In 1926, Roach,184 investigating the effects of carbon compounds on soil algae, first introduced quantitative methods in the study of algal growth. Another landmark was Barker's report185 in 1935 on oxidative pro cesses in the colorless alga Protothera Pearsall and Loose188 demonstrated in 1937 that the metabolic changes in Chlorella growth are basically similar to those occurring in the leaf of a higher plant. The history of algal culture techniques was well summarized in 1946 by Priogsheim 107 Only during the past two decades have the diverse studies of algal metabolism been unified into a cohesive entity. The first truly compre: heasive reviews of algal physiology and chemistry were those of Myers 268 aad of Blinks in 1951 in 1953, Schussaig's handbook on plant biochemistry180 summarized many physiological data on phytoplankton, while ·PLANKTON 29 Fogg's monograph offered an invaluable compilation of facts on the metabolism of algae In the same year, the Carnegie Institution issued "Algal Culture: from laboratory to pilot plant”41; this volume, collected and edited by Burlew, made available extensive data on algal metabolism and culture. Additional data on the metabolic aspect of algal growth was provided in the "Growth of Protozoa, "190 issued in 1953 by the New York Academy of Sciences. 2 Metabolism, Culture, and Nutritional Values Even more so than with zooplankton, the recent intensive interest in phytoplankton has been sparked by the growing disparity between world population and food supply.191,182 The attractiveness of phytoplankton as a food is enhanced by several inherent qualities. Practically all of it is digestible, in contrast with higher plants, which contain a large proportion of cellulose The protein content is higher than in field plants, it is of low molecular weight, and it contains most of the required amino acids. Under controlled conditions, an excellent Ispıd yield can be achieved. Vitamins and minerals are present in abundance And, best of all, algae can be cultivated and almost always be maintained at a maximal growing rate, whereas land crops utilize sunlight most efficiently only just before harvest Interesting comparative figures are offered in Table III Microscopic algae have a basically simple cellular structure 103-188 Like other pigmented plants, the algal cell usually comprises a nucleus, one or more chloroplasts, and a cytoplasm containing various formed structures The pigments are generally concentrated in the chloroplasts, the "solar engines" which are now considered the main sites of photosynthetic activity.197 Here the pigment enables algal cells to convert inorganic to organic substances by utilizing radiant energy.108 The key pigments involved in algal photosyathesis (as condensed from Fogg') are shown at the top of page 31 The chlorophylls and carotenoids are both fat soluble. The former embody a cyclic tetrapyrrolic qucleus with magnesium, the latter generally contain about 40 carbon atoms. The carotenes are hydrocarbons, while the Xanthopbylls are oxygen derivatives of the carotenes The phycobilins differ from the other pigments in being soluble in water but not in fat solvents. They resemble the chlorophylls in possessing a tetrapyrrolic nucleus, but the latter is lineat instead of cyclic, and is linked to a globulin instead of to a metal.

·30 Ulva sp. 367 . .. Alfalfa .... Castor. TABLE III.--Elementary Analysis and Calculated Composition of Some Morini Algar, Leaves of Land Plants, and Freshwater Algae FORM ASI с N R-saluet PROT. CBH LIPIDS * ash-free dry wt % ash-free dry wt % dry at by analysis calculated composition MANNE ALGAE Gigurtins Agardhu 17 82 42 90 6 27 583 29 85 364 63.6 trace 18 72 44 36 6 09 4 87 30 33 304 696 trace Amphipkura rutilans 46 24 46 52 6 48 33 67 354 62 8 18 Macrocystis pyrifera 37.66 4602 682 660 32 24 41 2 576 Naricula sorgwatum 35 10 48 45 672 8 80 36 22 55.0 43.7 1.3 Egregria Menziesi 31.74 49 97 6 29 3 39 36 27 33 7 57.2 91 LAND PLANT LEAVES Corn 5.39 46 69 604 2 02 31 88 126 820 5.4 Begona 12 92 45 73 6.39 3 04 32 00 190 776 34 Pine 313 SI 83 652 2 40 37 74 150 65 7 19.3 10 31 49 90 6 86 7 39 38 00 462 449 89 13 62 51 12 638 563 38 OZ 35 2 51.9 129 Sunfower 14 09 51 04 629 987 38 37 61 7 339 4.4 Nasturtun 9 58 50 46 6 84 38 45 428 459 11.3 11 80 50 87 6 78 770 38 77 481 41 7 10.2 12 25 51 91 701 7.11 40.08 444 40 & 148 Cedar. 5 53 53 58 708 291 40 75 18.2 560 258 FRESHWATER ALGAE: Chlamydomonas sp 4 74 47 46 682 581 35 27 36 3 58.2 3.5 Anabaenopsis sp.. 935 49 37 700 7 28 37 88 45.5 45 6 89 Oikomonds fermo. 3 08 $1 76 760 5 36 40 85 335 45.8 20 7 Strebococcus baallaris 650 5265 699 996 41 44 62 3 258 119 Suncbococcus baallarts. 11.24 57.14 8 12 361 46 51 226 38.3 389 Chlarella pyrenardosa... 3 45 49 51 678 9 31 37.92 580 37 45 Cblorella pyrinodosa 3.46 70 17 10.53 1.43 63 33 87 57 85.6 * Synthesized from several tables ated by Milnerti R-value, us conceived by Spoche and Milnet, is the level of reduction of the total organic mattet contained in plant material. The scale of R-values ranges from o for CO: to 100 for methane. the proportions of protein, carbohydrate, and lipid Protein 15 6 25 x nitrogen From the R value calculated on the basis of carbon, hydrogen, and oxygen analysis, and from the nitrogen content, one can estimate 691 Tobacco Flax.. THE ROLE OF ALGAE AND PLANKTON IN MEDICINE ·PLANKTON 31 KEY PIGMENTS IN ALGAL PHOTOSYNTHESIS CHLOROPHYLLS Types a, b, c, d, CAROTENOIDS Carotenes Types A, B, C Xanthophylls Lutein Siphonetn Dradinoxanthin Zeaxanthin Siphonoxanthin Dinoxanchin Violaxanthin Fucoxanthin Neodinoxanthin Flavoxanthin Neofucoxanthin Peridinia Neovanthin Diatoxanthin Myxoxanthin Myxoxanthophyll PHYCOBILINS rand - Phycoerythrins rind c Phycocyanios The distribution of these pigments in algae is rather variable The commonest chlorophyll is the type a, just as in gher plants Chlorophyll b, also present in higher plants, occurs only in the Chlorophyceae and the Euglenincae The other classes contain chlorophylls c, d, and e Of the carotenes, the B-type is most abundant The a-carotene, typical Chlorophyceae and the higher plants, is not found in some algae In Bacıllarıophy. ceae, for example, it is supplanted by e carotene Each algal class also has its own characteristic xanthophyll, as may be deduced from the num. ber that have been identified Phycobilins, on the other hand, occur only in the Rhodophyceae and the Myxophyceae, the former containing mostly phycoerythrin, the latter phycocyanın, the origin of these related pigments in either Rhodophyceae or Myxophyceae is indicated by the designation r of C, respectively. Granick and his associates reported 199,200 a 1953 on the protoporphyrin precursors produced by Chlorella mutants They have been particularly interested in the relationshup between chlorophyll and heroin, seeking information on porphyria and blood formation in human berogs Della Ros2,201 using 14, demonstrated that Chlorella vulgaris was able to synthesize chlorophyll from acetate and glycine; he also showed that chlorophylls a and b are not interconvertible Jorgensen"1 in Venezuela demonstrated to one of us (MS) the red blood wiggler Corronomus growing on the sides of his Chlorella culture containers; he suggested that there might here be a conversion of chlorophyll to here French and Young202 indicated that phycocyanin may be an intermediate in the resonance transfer of energy from phycoerythrin to chlorophyll.

·32 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE The photosynthetic role of the various pigments can be estimated by amparing the absorption spectrum with the "action spectrum" or photosynthetic capacity of light at various wave lengths 10,41 Photosynthetic efficiency is usually measured in terms of oxygen production. The chlorophyll absorption bands are chiefly in the blue, blue-green, red, and infrared ranges The yellow-orange color of the carotenoids results from absorption in the blue and green. The chlorophylls and phycobilins are often strongly fluorescent in solution. Tucker203 has even used pigment extraction with acetone as a method of quantitating phytoplankton. Despite all the emphasis on photosyothetic capacities, it must be noted that not all algae contain pigment Beijerinck180 long ago noted the natural occurrence of yellow and colorless colonies of Chlorella variegata on culture media Bogorad and Granick200 produced colorless, pale yellow, and light green mutant colonies of Chlorella vulgaris by radiation Butler206 exposed Chlorella pyrenordosa to ultraviolet light and obtained colorless mutants without chlorophyll. He also noted that similar perm. anently colorless forms of Prototbeca have been observed in nature, and that they were probably mutants from Chlorophyceae. Very recently Tolbert and Zill203 reported photosynthetic studies with protoplasm ex truded from Chara and Nitella. They found activity 12-15 per cent that of the whole cell, as measured by C, fixation. Both protoplasm and cut cells reduced carbon dioxide in light to sucrose and hexose phosphate. Dark cootrois fixed Cza into products associated with plaat respiration, but it is not clear whether they represented true photosynthetic pigments. In fact, the whole sequence of biochemical synthesis in the dark or without pigmeat remains rather obscure This is particularly so in the light of Duysens' daim208 that light energy is convertible only through chlorophylla or b (he maintains that in diatonis, for example, carotene must be converted to chlorophyll a.) Radiation and natural mutation are not the only methods of producing non-pigmented algac Dubiezot produced chlorophyll-deficient Chlorella t'ulgaris with streptomycia And even strong radiation does not always effect alterations Blinks208 reported there was no significant change in coroposition or function of marine algae one year after the atomic explo sions in the Bikini area The only exception was an increase in catalase, possibly due to an increased hydrogen peroxide concentration secondary to radiation. Algal cells generally do not grow old and die; iastead, they produce more cells by binary fission However, not all algae grow by cellular ·PLANKTON 33 division It has not been widely realized that sex may play a role even in this remote corner of the cosmos' Sexual union of two gametes to produce a zygote does occur, particularly among the Phaeophyta and Chlorophyta (Chlorella and Scenedesmus are notable exceptions.) Either or both gametes may be Aagellated, or mating may be achieved by amebord motion Gilbert M. Smith209 notes that algal sexuality was initially de. scribed in 1858 in Oedogonium. In 1926, Jollos demonstrated that the heterothallic green alga Dasycladus excretes sexual substances which in. fluence the movement of gametes toward each other Since then, much experimentation has been carried on with Chlamydomonas eugametos, a unicellular green alga Motility is absent in cells grown anaerobtcally in the dark, but it is revived by filtrates from cells grown in light The substance responsible is reported to be crocin, an ester of the sugar genttobiose combined with the carotenoid pigment crocelin There are also substances affecting maleness and femaleness; these are activated by exposure to light, weakened by dilution The vırılızıng factor appears to be safranal, the feminizing one a gentiobiose of safranal. There are all degrees of sexual activity, depending upon the concentration of activating substances and exposure to leght There are also varying degrees of maleness and femaleness, even intermediary forms, depending upon the ratio of opposing substances. Similar findings were reported by Sager and Granıck211 in 1954 They noted that depleting the nitrogen supply in culture media of Chlamy. domonas reinhardı encourages the differentiation of ordinary cells to gametes If nitrogen is added, the process is reversed Light apparently acts only indirectly via photosynthesis, and it is not obligatory for zygote formation of dark-grown nitrogen-depleted cells Much has been learned about the metabolic characteristics of algae by observations of cultures Chlorella has been widely used in photosynthetic experiments especially because it can grow under such varied environ. mental conditions The rate of algal growth is often measured by "generation time"—the time required to double the number of cells in a culture 41 There are two major phases of growth, one chiefly of cellular multiplication, the other of storage of reserve materials in the first phase, protecas predominate, in the second, fats and carbohydrates The main factors influencing algal growth are. available radiant energy, carbon dioxide concentration, temperature, and composition of nutrient substrates The exact channeling of the various nutrients may depend upon the amount of available nitrogen, or upon the presence of enzyme systems. Chlorella ·34 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE extracts, for example, have been shown to contain transaminase systems which aid the transfer of amino groups to either pyruvic or a-keto-glutarıc aad 212 The most active enzyme systems in algae are those of aspartic and glutarıc acids, and alanıne. Spoehr and Milnertl demonstrated that the composition of Chlorella pyrenoidosa could be varied specifically simply by controlling cultural conditions In over 300 pure cultures, it has been possible to produce the following wide range of variations in composition and R-value Protein 73 to 88 0% Lipid 45 to 860% Carbohydrate. 57 to 38 0% R-value. 37 92 to 63.33 The amount of radiant energy present markedly influences the rate of growth of a culture, either directly or by increasing the nitrogen absorption The latter is certainly of critical importance, since a Chlorella cell can no longer undergo fission when the nitrogen content falls below 16x 10-13 gm As has already been indicated, the protein content of microscopic algae may vary widely; its level may be grossly estimated from the nitrogen content There is a definite similarity in the protein content of Chlorella and the leaves of many higher plaats Under optimal cultural conditions, a protein level can be achieved in Chlorella which exceeds that of the best vegetable substances used as animal feed This includes such materials as Brewer's yeast, torulx yeast, soybean meal, dried skimmed mulk, and whest Not very much is known of the specific nature of algal proteras, except that there is often a resemblance between their amino acid patterns and those of higher plants 213-215 The nutritional value of Chlorella is based not merely on its high protein content, but also on the fact that it contains all the essential amino aads Methionine is present in only small amounts, but this deficiency also occurs in many vegetable proteins. The essential amino aad Biological Index of Chlorella vulgaris has been calar. lated to be 62 in a pilot plant assay, as compared with an arbitrary value of 100 for whole egs protein The Index of most animal proteins canges from 80 to 90, while cereal proteins are usually between 60 and 80. Thus Chlorella is roughly comparable to white flour, corn gluten, and peanut meal. An amino aad assay of dried Chlorella is given in Table IV. Although most algae are able to utilize wide varieties of basic inorganic nutrients and to synthesize from them the more complex metabolites re quired, some are not so versatile. They may not be able to synthesize all ·PLANKTON 35 TABLE IV-Aino Acid Assay of Dried Chlorella Pilet Plant Laboratory Natrrent Sampk Sample Torula Yeast Crude protein 44 00% 40 00% Arginine 2 06 2 39 3 61% Histidine. 062 065 1 31 Isoleucine 1.75 169 3 75 Leucine 3 79 199 3 57 Lysine 206 243 4 14 Methionine 0 36 0 57 084 Phenylalanine 1 81 2 14 241 Threonine 2 12 191 2 58 Tryptophane 080 0 41 0 66 Valine 2,47 2 67 298 Glycine 2 20 0 22 the amino acids from ammonta and the requisite carbon fractions, and they must thus depend at least in part upon an exogenous source of some amino acids The deficiency is not of the amino acid per se, but of the capacity to form amino groups from ammonia Thus, for proper growth, Euglena deses requires aspartic acid, while Chlamydomonas chlamydogama needs histidine and aspartic acid.40 Nucleic acid has been demonstrated by staining reactions to be present in algal cells Jeener216 reported that Polytomella caeca had 6 to 10 per cent of its protein as ribonucleic acid in actively growing cells He indscated that there was great variability in the synthesis of nucleic acid, and that this was unrelated to the rate of general protein synthesis and cellular multiplication Szafarz and Brachet217 indicated that formation of ribo nucleic acid can proceed independently of the nucleus in Acetabularia mediterranea Studies by Goryunova218 have indicated the presence of mucins in Oscillatorid The assimilation of nitrogen by algae is important not only with respect to amino acids, but also in the synthesis of various carbon compounds As in the case of certain bacteria, many autotrophic algae are characteristically capable of nitrogen fixation Nitrogen may be absorbed as the element, or as outrate of ammonia Frank,219 in 1889, first reported the possible nitrogen-fixing properties of certain algae, but this was not proven until 1928 220 Since then, further corroboration has come from Kjeldahl determinations and tracer studies using nitrogen isotopes.18 So far, over 20 species of Myxophyceae have been proven capable of fixing nitrogen. Aspartic aad, glutamic acid, succinamide, asparagine, and glutamine can ·36 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE serve as the sole sources of nitrogen for various green algae via dezmina. tion or transamination. It is probable that nitrogen enters the general metabolic pool most often in this manner. Nitrogen-fixing algae can contribute to the fertility of the soil, especially in tropical areas. A notable example is the utilization of blue-green algae in the Usar area of northern India to decrease the excessive alkalinity of the soil 15 The algae thrive during the rainy season and decrease the pH of the soil from 90 to as low as 70, and thus make possible the growing of field crops during the dry scason This resembles somewhat the employment elsewhere of large seaweeds as direct fertilizer. The cultivation of rice is also aided by microalgae The latter flourish in the wet rice paddies and, by fixing atmospheric nitrogen, enable perennial cultivation without additional fertilizer. Nitrogen fixation here may be greatly enhanced if bacteria are also present and contribute their fixation activities In addition, the algae, by producing oxygen, help to aerate the rice plants Watanabe221 reported in 1991 that 13 species of 643 blue-green algae from Far East and South Sea area nice fields were capable of atmospheric aitrogen fixation, chiefly in warmer climates. Most of them belonged to the genera Tolypothrix and Nostoc, some to Schizothnix, Caloshrix, Anabaenopsis, and Plectonema Aspartic and glutamic aads and alanıne were the systems mostly involved a similar function was reported by Douin,222 who noted a symbiosis between the Cycadares plant and Anabaena cycadeae, which fixes oitrogen for its host Much less is known about the casbohydrates of microscopic algae than of the macroscopic 10,11 Whereas the latter may contain large quantities of carbohydrates in the cell walls (chiefly algınıc complexes), the microalgae have relatively scanty structural materials, and their storage products are lipid rather than starchy Milner has isolated statch and sucrose from Chlorella pyrenoidosa Broun found 17 8–20.2 per cent (dry weight) total carbohydrate in Scenedesmus obliquus The substances included mainly insoluble polysaccharide, there were also small amounts of free reducing sugar, sucrose and other oligosaccharides, and water-soluble polysaccharides Healthy algal cells, particularly in older cultures, liberate various organic substances into their fud environment These consist primarily of nitrogenous substances and pentosans. It is probable that more specific knowledge of microalgal casbohydrates will emerge when higher carbohydrate contents are achieved by varying cultural cavironmental condi. tions, ·PLANKTON 37 The lipids represent one of the most important components of microscopic algae 23 Many investigators have confirmed Beijerinck's observation in 1921 that fat droplets accumulate in diatoms as the cultures become old and the available fixed nutrogen decreases 40,41 This occurs, as We have indicated earlier, because nitrogen deficiency limits growth and favors the building of reserve materials The ratio of lipid to carbohydrate is variable, it depends far less on algal genetic trarts than it does on environmental factors such as nitrogen supply and enzyme systems Also noteworthy is the fact that algal fat metabolism is but little reflected in gas exchanges The lipid content differs not only from one species to another (the blue-greens have very little), but also may vary greatly within the same species with changes in cultural conditions Chlorella generally contains 20-25 per cent fat, yet nitrogen starvation can increase the amount to as high as 86 per cent Even during World War II, the Germans, culturing Chlorella, Nitzschia, and Scenedesmus as potential sources of needed fat, attained lipid yields of 40-70 per cent These comprised chrefly the triglycerides of stearic, oleic, and linoleic acids Most of the fat in phytoplankton is unsaturated, with a high proportion in the 16 and 18 carbon series 41 Detailed analyses of Chlorella, for example, show 54–67 per cent unsaturated C-18 and 18-29 per cent C-16 acıds Unsaturation ranges from 1 60 to 2 25 double bonds per molecule Palmıtıc is the main saturated acid, while stearic constitutes less than 4 per cent of the total fatty acid As the lipid content of Chlorella rises, the unsaponifiable fraction decreases and the fatty acid portion increases, at the same time there is a significant fall in the degree of unsaturation of the fatty acids Chlorella fat resembles other unsaturated plant lipids and falls into the chemical classification of "drying oils" Sterols have been obtained from most of the algal groups, excepting the blue-greens Klosty and Bergmann224 reported in 1952 that chondal lasterol is the principal sterol in Scenedesmus obliquus, while ergosterol predominates in Cblorella pyrenoidosa. Chondrillasterol may be remembered as having been favorably considered a few years ago as a starting point for cortisone synthesis Table V offers analyses of the various lipids in several samples of Chlorella Phytoplankton, like the macroscopic seaweeds, has a faigh vitamin content It has been determined that most of the known vitamins are present in Chlorella and related microalgae The quantities are eminently adequate for the algae to be used as food (for man or animals), but not if ·38 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE 1 4 75 31 868 33 120 TABLE V-Analysis of the Lipids of Chlorella Analysis Chlorella lot no 2 3 1 Tocal lipid (6 of Chlorella) 23 37 33 17 62 96 Composition of rocal lipid 2 Fatty acids ($ of lipid) 28 0 49 5 830 3 Unsaponſable fraction ($ of lipid) 7.7 33 4 Water-solable, after saponification (% of lipid) 600 428 13 7 5 Calculared fat (% of Chlorella) 685 Analysis of total facty acids 6 Palmitic acid, C16 (%) 16.6 10.9 79 7 Stearic acid, C18 (%) 04 41 39 8 C16 unsaturated acids (%) 29.1 18 3 27.2 9 C18 unsaturated acids (%) 539 667 609 99 696 172 347 11 4 180 67 1 1000 999 1000 272 7 2695 163 1 273 6 143 8 274 1 125 3 4 4H -41H -4 SH 10 Tocal fatty acids (%) 1000 11 Equivalent weight 12 Iodine number 1436 Degree of unsaturation 13 C16 plus C18 unsaturated acids -44H -36H -37H - 32H 14 C16 unsaturated acids 15 C18 unsaturated acids -34H they are intended purely as a source of vitamin concentrates In the latter case, synthetic methods are more economical. Microscopic algae are the ultimate basic source of vitamins in fish, and the finding of large quantities of Vitamin A in fish livers was indeed the stimulus for much of the study of algal vitamins 225 Several decades ago, it was shown that Nitzschia closterium and other phytoplankton were good sources of Vitamin A and its carotene precursors 726 in the early 1930's, Jorgensen in Scandinavia considered using algal carotene to feed cows and to fortify margarine 11 Later, in Venezuela, he used dried plankton from Lake Maracaibo as a source of carotene and related unidentified substances as a highly efficacious food supplement in the therapy of lepers. Vitamin B, is generally necessary for algal growth, and it has been demonstrated in most species in which it has been sought.so Especially large amounts are found in Chlorella Young cultures with activelydividing cells contain more thiamun than do the older fat-storing ones Thus, von Witsch obtained yields of 1.8 to 180 gamma per gram of dried substance from cultures of varying ages While Chlorella and related algae contain dehydrogenase for direct ·PLANKTON 39 oxidation of pyruvic acid, there is also some alternative participation in the Krebs tricarboxylic acid cycle, as in vertebrate tissues For this reaction, the co-carboxylase thiamın pyrophosphate must be present An example is the great enhancement of pyruvic acid oxidation by the addition of thiamin to the thiamin'deficient Protothera If Vitamin B, is not present as such in the aqueous nutrient, it must be synthesized by the algae Most are capable of so doing, but some cannot synthesize the thiazole, others the pyrımıdıne, radicles of thiamin Thus the thiazole portion must be supplied to Polytomella ocellata, and the pyrimidine component to Euglena gracilis and to a radiation mutant of Chlamydomonas moeuws The pyrimidine-thiamin requirement of the Euglena can be markedly decreased by the addition of glutamate. Both thrazole and pyrimidine must be supplied exogenously for the growth of Polytomella caefa, Chilomonas paramoecium, and Protothera zopfi. Riboflavin, like thiamin, has been demonstrated in practically all algae in which it has been sought It is a component of the prosthetic group of the favo-proteins, but its essentiality to algal growth has not been shown Para-aminobenzoic acid is known to be necessary for the growth of only one alga, a mutant of Chlamydomonas moewus! With the latter and with Nitzschia, para-aminobenzoic acid has the same antagonism to sulfonamide inhibition of growth as with many bacteria Aniline may replace para-aminobenzoic acid as a growth factor for Chlamydomonas with an efficiency of one per cent, but it does not neutralize the sulfonamide effect 40,227 Plankton paste contains considerable quantities of cyanocobalamin or Vitamin B 2 40,41,228 Traces of this substance are necessary for the growth of various Euglenneae as well as other classes, such as Chlamydomonas chlamydogama of the Chlorophyceae Vitamin Bı, may be concerned in the synthesis of desoxyribosenucleic acid Ascorbic aad is present in phytoplankton (especially Chlorella) in amounts comparable to those in lemon juice, however most of it is lost in drying and storage. Although little is said about the Vitamin D content of microalagae, feeding experiments have indicated good antirachitic activity. Indirect evidence comes from the potency of fish liver oils, which ultimately derive their Vitamin D from planktonic sources,229 As with seaweeds, the vitamin activity of phytoplankton is based upon ergosterol content. Substances with coagulant effect have been demonstrated by Dam, who determined the Vitamin K content of Chlorella as six gamma per gram of dried material 230 Phytoplankton has a great potential as a source of vitamins in human ·40 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE nutrition Though most of the studies have been with Chlorella, other microalgae are comparable. It has been calculated that a quarter pound of Chlorella will supply all the daily minimal human vitamin needs except for ascorbic acid Table VI, taken from Burlew, ti details the vitamin content of two samples of Chlorella TABLE VI -- Vitamin Assay of Dried Chlorella Pilos Plant Laboratory Vitamin Sample Sample Carotene (mg /Ib ) 2180 Thiamin (mg/lb ) 110 43 Ribofavin (mg /11 ) 26 2 163 Niacin (mg /lb ) 540 109 0 Pyridoxne (mg /16 ) 10 4 Pantothenic acid (mg /lb ) 36 91 Choline (mg/lb ). 13700 Biotin (mcg /Ib ) 67 0 Vitamin B, (mcg/lb ) 450 100 Lipoic acid (acecate units/1b ) 15 Algae require basically the same nutrients as higher plants-itrogen, phosphorus, potassium, sulfur, magnesium, and iron Unlike the higher plants, all except a few algae can dispense with calcium Potassium and magnesium (and occasionally calcrum) are important in the photosynthetic process because their bicarbonates can make available supplemental carbon dioxide In addition, algae also have need for certain trace elements or micronutrients Myerstl has specified algal requirements for manganese, zinc, calcium, boron, and copper The exact metabolic role of these has not been elucidated Neither has it been for the following substances, which have also been identified in algal cells antimony, arsenic, beryllium, cobalt, germanium, jodine, lead, manganese, molybdenum, nickel, rubidium, silver, strontrun, tin, titanium, tungsten, uranium, vanadium, and yttrium (Does not this listing of elements sound more like a roll-call of our present-day electronics and metallurgical industries, and are not the biologic and the mechanistic worlds drawing closer?) It has been determined that, because of selective absorption, the cellular content of manga. nese, strontium, uranium, and yttrium is above that of the surrounding media. The only known role of molybdenum is that of fixing sitrogen in Anabaena Even though their functions are obscure, the addition of trace elements to algal culture media has become standard operating procedure. An excellent means for providing continuousiy adequate micronutrients are ·PLANKTON 41 the chelating of complexing agents. These are biologically inert cyclic organic compounds ("inner complex salts”) resulting from the attachment of a group at two points to the same metallic atom They prevent excessive absorption and precipitation Ethylenediamine tetraacetic acid (EDTA) is the chelating agent most commonly used When a given amount is added to a nutrient medium, it will permit the release of enough ions through mass action to provide for cellular needs By forming easily reversible complexes of trace elements, it provides tonic concentrations at preferred levels throughout the life of a culture Besides the trace elements, there are other unidentified factors sometimes necessary for algal growth It is common observation that some algae grow with ease an impure culture, yet do not fourrshma bacteria-free media Gymnodinium cannot be continuously subcultured without an organic factor extracted from soil Similarly Cryptomonas oralt, Synura orella, and Gloeotrichia echinulata demand the presence of certain soil factors Also, Dityllium bng elle requires, for good growth in artificial seawater, two organic substances from natural seawater One of these is a specific sulfur grouping, the other a substance also found in yeast or algal extracts 40 The exact nature of these various needed supplementary compounds will doubtless become clearer with more experience in algal culture Although mass culture of phytoplankton for food and special organic material has long been the subject of considerable discussional enthusiasm, little specific investigation was done until World War 11 231 Most of the experiments have been performed with some species of Chlorella, Scenedesmus, or Nitzschia because they grow so fast and tolerate a diversity of cultural conditions Chlorella pyrenoidosa has been an espeaal favorite because of its previous wide use in studies on photosynthesis As listed by Burlew in "Algal Culture, from Laboratory to Pilot Plant,"41 the elements of a plant for growing algae on a large scale are relatively simple 1. A container with a transparent upper surface 2 A means of arculating the culture medrum, so that the algae do not settle 3 A means of controlling temperature 4 A means of introducing carbon dioxide and other nutrients continuously. 5 A means of harvesting the algae almost conttauously. 6 A means of preserving the harvest until used ·42 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE A great variety of apparatus has been employed to achieve the mechanical requisites. In 1951, Jorgensen and Convit in Venezuela grew freshwater algae in unglazed baked sed clay bowls exposed to the open air and hot sun The nutriment consisted of a commercial fertilizer suspended in water. Evaporation from the porous sides of the bowls kept the temperature at 26°C even in direct tropical sunlight Von Witsch and Harder in Germany utilized nutrient solutions in glass tubes 3 x 30 cm and 6 x 150 cm, these were illuminated by 300-watt water-cooled lamps and supplemented with an air stream containıng 05 per cent carbon dioxide. At the Carnegie Institution in Washington, greenhouse culture of Chlorell: was carried on in five-gallon botties in 1947, illumination, temperature, and carbon dioxide supply were controlled This study led the following year to activation of a successful pilot plant for continuous Chlorella culture at the Stanford Research Institute In 1949, Geoghegan in England tried out a variety of culture vessels cylinders 15 x 175 inches, large tubes 4 5 feet x 2.75 inches; aspirator bottles 18 x 10 5 inches, and a tall outdoor plastic tank 4 5 feet high, 15 feet long, and 0 33 feet wide He concluded that shallow horizontal tubes or troughs were preferable, and that sunlight is more economical than artificial illumination Wassink and a group at the Agricultural University at Wageningen in Holland experimented with mass Chlorella culture from 1948 to 1950 Instially they used inverted half-liter glassstoppered bottles with incandescent light Later they tried outdoor sunlit square meter tanks with 300 liter capacity. The tanks were covered with glass to minimize contamination, carbon dioxide was supplied from a cylinder, and agitation was accomplished by a motorized stirser Algae were harvested after 5 to 7 days of growth Subsequently they set up similar concrete tanks in a dark room and illuminated them with either daylıght fluorescent bulbs or high-wattage incandescent lamps They obtamned a 12-20 per cent efficiency of light-energy conversion, and there. from concluded that excessive illumination could decrease algal growth under either artificial or natural conditions Gummert and his colleagues, working in Essen in 1950-51, hoped to demonstrate that local conditions were suitable for large-scale outdoor and greenhouse culture that could take advantage of the huge amounts of waste carbon dioxide in the industrial Ruhr. They succeeded for the most part, but their deliberately nonsterile procedures resulted in troublesome contamination with foreign algae and protozou. This could be overcome by using protozoa-resistant Scenedesmus, by altering the nutrient media, ·PLANKTON 43 by increasing the amount of illumination, or, in fact, by doing anything that promoted faster growth of the desired algae Much was accomplished in America from 1950 to 1953 under the guidance of the Camegie Institution of Washington They helped sponsor sach investigators as Spoehr, Milner, Davis, Myers, and Krauss, with much of the work done in the Department of Plant Biology at Stanford Here studies were performed not only with various types of algae and nutri. ments, but also with differing forms of culture equipment The latter included large bottles, rocking trays, and plastic and glass tubing The culmination of the Carnegie studies was the establishment of a large pilot plant for Chlorella culture in Cambridge, Massachusetts, by Arthur D Lettle, Inc, a consulting research and development organization Three large-scale culture units were developed and studied The last, and best, consisted of a massive plastic tubing in U-shape, set up on a roof-top with free exposure to suplight it was equipped with a centrifugal pump to circulate the culture, a measured inflow of carbon dioxide (5 per cent in air), a heat exchanger for cooling the culture, a harvesting system, and apparatus to freeze or spray dry the harvested material from this study, it was concluded that with this type of pilot plant, 20 gm /sq. meter/day could be realized, equivalent to 17 5 tons Chlorella per acre per year Pilot plant studies were also carried on in Israel and in Japan The former were begun in 1951 at the Hebrew University in Jerusalem under government sponsorship The latter were done in Tokyo in 1952 at the Tokugawa Institute for Biological Research, Tamiya, the director of this group, spent some time as guest worker in the Carnegie Institution labora. tories in Stanford through a fellowship supported by the United States Office of Naval Research Experiments on mass culture of algae are also being carried on by segments of private industry An example is the reported work of Pruess and his colleagues232 for a pha naceutical concern on algal culture in carboys and in deep tank fermentations Kindred to the latter is the growing of in 50-called "oxidation ponds," with the double purpose of sewage disposal and formation of algal protein suitable for feeding Renn233 has estimated that domestic sewage from a community of 10,000 could pro. duce nearly 1,400 pounds of protein daily of properly treated Oswald and his colleagues251 at the University of California are also enthusiastic proponents of this method. One eager college professor-º has already designed a house with an algal pond on its flat roof. No comment is offered on the monotony of having to stick to one brand of sewage!

·44 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE The enthusiasm for culturing phytoplankton for feeding is based on reports that it is innocuous and that it possesses great nutritionat value These have been well summarized by Fisher and Burles.41 Mention has already been made of Jorgensen's feeding plankton soups to lepers in 1941, with a resultant improvement in weight and energy. These comprised cultures of Chroocottus, Homocyshimea, Oocystis, Ankistrodesmus, Chlorelld, and Scenedesmus. More recently, Jorgensen told one of us (M.S.) that an algal broth, when fed to mice with skin cancer, brought about an appreciable decrease in the size of the lessons; inexplicably, however, test animals maintained on such a dict did not have a long life span Currently Jorgensen is feeding his phytoplankton in rather palatable compressed pills Another early feeding experiment was that in 1949 at the Stanford Research Institute Rats fed on a diet containing a third Chlorella pyrenaidosa had a weight gain only one-third that of a control group This has been attributed either to the unpalątable flavor or to the mechanical difficulty resulting from the hygroscopic nature of the food preparation Henry, at the National Institute for Research in Darrying, in Engiand, fed freezedried Chlorella prepared by Geogbegan to young rats No adverse effects resulted after four months on a 17 per cent Chlorella diet In a four-week comparative study, the protein efficiency of Chlorella appeared slightly superior to that of dried brewer's yeast and peanut meal, but inferior to that of skim milk in 1951, Combs,235 at the University of Maryland, tested New Hampshare chickens on Chlorella provided by the Carnegie Institution. A marked increase in growth and an improvement on feed efficiency resulted from adding 10 per cent Chlorella to a basal diet in place of soybean meal. This was attributed to the high vitamin content of the alga However, a growth depressant effect was noted after extended feeding because of impacted beak conditions This occurred much sooner with 20 per cent Chlorella feedings Similar results were obtained in 1952 with chicks when vacuum-dried Chlorella was tried by a feed supply concern Most of the difficulties encountered were apparently due to excessive water absorption by the algae Other methods of preparation would seem to be indicated Tamıya22,41 has included substantral announts of Chlorella in preparing highly acceptable bread, noodles, soup, tea, and ice cream. He has also used hydrolysates of dried algae as a substitute for soy sauce Hes students and even his colleagues admitted no ill effects whatever from repeated ingestion of these concoctions.

·PLANKTON 45 The palatability and acceptability of phytoplankton as a human food are dependent upon many factors These include especially appearance and taste These depend, in turn, upon whether the material is fresh or dried, also, whether it 15 eaten as such, or mixed (and diluted) with more familiar foods Doubtless much improvement can and will be made sa all these respects for a greater acceptability Certainly these items must be worked out before plankton is really the "food of the future "*238 3 Medical Aspects a Water Supply and Sewage Disposal Despite all the hopeful reports on their merrts, it must be emphasized that phytoplanktonic organisms can exert adverse effects upon the waters in which they flourish These include not only changes in appearance, odor, and taste of the water, but also changes in other aquatic organisms 287 249 Even more important are the varying degrees of intoxication and reaction in animals and humans who ingest or come in contact with these waters The potential economic losses are self-evident Similar to the striking colorations of the seas previously mentioned (eg, Red Sea, Vermilion Sea) are the alterations in appearance of fresh water bodies "Water bloom" is present when extensive growths of microscopic algae blanket the surface of a lake, pond, or stream In Europe the phenomenon has been variously called "Wasserbluthe," "deur d'eau," and "flos aquae", in some North Central parts of the United States, the ancestral Dutch "vasserbloon" designation is used Under favorable crcumstances, microscopic algae, especially the Myxophyceae, can grow so rapidly that the water assumes the color of the organism And these colors are indeed variegated The Oscillatoria and Cylindrospermium produce a green slime and a bluish soup, respectively. Chlamydomonas and Pleuro. corrus are a brilliant green Ceratrum bitundinella presents a rusty color and a powerful stench-perhaps the basis of the Biblical story (Exodus 7) of the turning of the Nile into blood and the death of the fish therein "Red water" and "red snow" in Australia are due primarily to Myxophyceae, but desmids (Conjugata) may be a factor Both "red tide” and "yellow tide" have been reported along the coast of the Gulf of Mexico, while two major outbreaks of "red water” occurred in Gokasho Bay, Japan, in 1933 and 1934, from Gymnodımım miktmotor. Whereas the colorations are interesting and occasionally even weird, the odors arising from aquatic algal growths are more important, both esthetically and economically They affect water supply and sewage dispo- ·46 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE sal Odors and tastes may arise from the oils of living algae, from products of active photosynthesis, from products of decomposition; and from the death of fish and other aquatic organisms The oils of living algae are often unpleasantly odorous, each oil being characteristic of the gene. rative organism The smells may be fishy, asomatis os grassy. Their disagreeable character may be heightened after planktonic death, when the frail oil globules are more widely dispersed. The odorous photosynthetic substances liberated during very rapid growth are ill-defined, but the offending materials present after algal decomposition are better known They include hydrogen sulfide and other sulfurous compounds, phosphorous and nitrogenous substances, methane, ethereal sulfates, and other volatıle gases Fish and other aquatic organisms may die from suffocation, or from toxic substances such as hydroxylamine resulting from the breakdown of algal proteins These dead animals contribute a liberal stench to the already odoriferous situation Often enough, the chemical treatments applied to kill off offending algae aggravate the situation even further The treatment, such as chlorination, kills the organism and Jiber. ates the taste and odor-producing materials, and chemical combination with chlorine may favor the formation of additional smelly compounds. The most troublesome algae in lakes and reservoirs are the blue-green ones, which grow rapidly and float high in the water, forming surface Greatest growth takes place in waters which are shallow and warm, and in those rich in nitrogen, phosphorus, and loosely-bound carbo. hydrates The principal causes of bad odors and tastes are the Myxophycene and Diatomales, which are less potent oxygenators than are other algae When unpleasant odors affect ordinary lakes or ponds they merely detract from the beauty of nature. But when they occur in reservoirs of water intended for daily drinking, they present a trenchant problem Even mild alterations in smell or taste may be psychologically unacceptable to the average person As Howard and Berry239 have pointed out, odors caused directly or indirectly by living plankton by the decomposition of vegetable growths are immediately ascribed the Jayman to con. tamination with sewage This is not too strange, since the decomposing plankton floats on the water, then is thrown onto the shore, where it forms odorous, blackish, discolored masses It is difficult to convince people that the odor itself signifies no health hazard When Irving algae are present in the drinking water, there may be a mild initial taste, but a bitter aftertaste; when the organisms are dead, there is a strong first taste, but praaiscums ·PLANKTON 47 cally no aftertaste Measures to control algal growth241-248 in reservoirs include periodic chemical treatment (with algicides such as copper sulfate), removal of accumulations from the shores before decomposition occurs, and restricting the food supply of the algae by removal of tree branches, leaves, and other floating debris While removal or eradication of algae is desirable 1a reservoirs, quite the opposite is true in sewage disposal units Oxidation ponds have been widely used for secondary sewage treatment As summarized by Oswald and his colleagues, 234 oxidative sewage treatment is based upon the fact that organic matter decomposes rapidly in water When in solution, it is readily available to microorganisms and their enzymes These convert complex organic compounds into simple substances such as carbon dioxide and ammonia Because oxygen is the ultimate hydrogen acceptor in these rapid conversions, it may become so depleted that aerobic oxidation is halted Thus the activated sludge and the trickling filter processes are so desig that atmospheric oxygen is forced into the liquid phase at an accelerated rate Algal photosynthesis offers a completely different method of supplying oxygen Here the availability of oxygen is independent of the physical laws normally governing aeration from atmospheric sources Bodies of water containing green algae may attain supersaturation with dissolved oxygen up to three or four times the normal maximum of water in equilibrium with air The organic waste material is changed by the gclic activity of algae and aerobic bacteria into more algal cells These are then disposed into natural streams or allowed to settle and then be digested anaerobically as bottom sludge The particular merits of this method lie in the minimal odor, nuisance, and operating cost It is therefore considered ideal for small communities situated in mild climates Particularly in Texas and Southern California has there been reliance on the use of sewage lagoons and oxidation ponds for adequate and economical treatment of sewage Algal purification of drinking water is also sometimes employed, on the other hand, uninvited algae often clog standard sand Giltration plants As previously indicated (section on Culture), Redna33 has recommended that oxidation ponds be used not merely for sewage disposal, but also to convert nitrogenous wastes into food protein This can be accomplished by encouraging the growth of algae (especially Chlorella, Scenedesmus, and Chlamydomonas) which yield a high quality protein A relatively new problem in sewage disposal concerns atomic wastes.251 Coopey252 demonstrated radioactivity in all forms of life in the Columbia ·48 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE River below the Hanford Atomic Energy plant in Washington. He estimated that about 570 tons of wet weight plankton passed the plant daily, with a resulting planktonic radioactivity about 0.6 curies per day. Analyses indicated that 5-30 per cent of the plankton's radioactivity came from isotopes with half-lives near to or longer than P32, while the remainder was of shorter half-life Some radioactivity persisted beyond 600 days' decay Since algae can transmit their radioactive toxicity right up through the nutritional pyramid to higher food animals, it is clear that more than conventional disposal methods must be devised for atomic wastes. An indirect application of algal attributes relates to mosquito control Matheson-19 noticed that mosquitoes did not breed in certain pools #here Chara fragilis thrived, he and Hinman then postulated that the large amount of oxygen from the algae either interferes with the larval food supply or gives the mosquitoes indigestion However, Biswas, 250 over a decade later, insisted that algal flora provide not only food for the mosquito, but also shelter! b. Animal Intoxications The microalgal scums, odors, and tastes described in the last section produce chiefly esthetic problems of far greater import are the many harmful reactions reported to have occurred in both animals and humans 253-273 Freshwater phytoplankton has been implicated since 1878 in the intoxication and death of domesticated animals, as well as water fowl, shorebirds, and their mammalian or avian predators In Table VII we present a chronological summation of 38 incidents of animal intoxications by phytoplankton as culled from the literature In most cases attacks occurred after the animals had drunk from lakes or ponds contaioing heavy algal growth, usually during hot weather They have variously been suspected of being due to botulism, anthrax, or worm infestations, or to poisons such as lead, senis, copper, cyanide, or alkali; or, as in South Africa, to gallsickness (carried by ticks), Jamsiekte, and poisonog by plants such as senecio, goustekte, and giſblaar. Also confused with algal poisoning has been the African geel dikkop of sheep, produced by photosensitizing toxins The reported symptomatology of algal intoxications has varied, but the most striking dinically have been the involvements of the neuromuscular and respiratory systems As described by Francis, 253 in the Australian outbreak due to Nodularia spumigeria, "the animals dereloped stupor and unconsciousness, falling and remaining quiet, as if asleep unless touched, ·PLANKTON 49 when convulsions came on, with the head and neck drawn back by rigid spasm which subsided before death." Similar manifestations have been reported from Alberta,269 Manitoba 262 North Dakota, 286 Minnesota, 254 Ontario, 289 and South Africa 258-280 In several of these, the animals also exhibited extreme photosensitivity, 266 with severe blestering of the skin,258 260 and also jaundice 258 Fitch"54 tested laboratory animals on the Microcystis and Anabaena which had killed cattle in Minnesota The characteristic symptoms in guinea pigs included restlessness, incontinence, deep breathing, sneezing, coughing, salivation, lacrimation, weakness in the hind quarters, clonic spasms, and death Rabbits presented the same picture plus opisthotonus Pigeons also developed opisthotonus, as well as rapid blinking and swallowing Mason and Wheeler270 injected Microcystis aeruginosa extract into mice, rats, guinea pigs, and cats. After a latent period of 20–180 minutes, there appeared pallor, hypotension, tachycardia, hypothermia, hyperglycemia, respiratory difficulty, and death Steyn258-260 in South Africa reported general paralysis of strychnine-like convulsions following consumption of large quantities of Microcystis by cows, smaller amounts resulted in constipation, drop in milk yield, generalized weakness, and severe photosensitivity of the skin If the animals survived longer, many developed jaundice and ascites Some of Steyn's laboratory animals manifested symptoms after simply being drenched with the contaminated water Smit271 injected or fed Microcystis to rabbits In acute cases, the animals developed anernja, restlessness, dyspnea, progressive paralysis, comna, and death within one-half to four hours In chronic cases, the outstanding finding was cirrhosis of the liver, with restlessness, atonia, and ascites McLeod and Bondar282 in Manitoba found that Microcystis administered to cats and muce caused a loss of equilibrium and progressive paralysis, later there were clonic muscular spasms, dyspnea, and cyanosis, then death within twenty hours The pathological findings in phytoplanktonic intoxications have been most interesting Autopsies made 11 1930 and 1933 in Minnesota showed "no gross pathology"254, cultures, smears, guinea-pig inoculations, and tests for copper and cyanide were also negative On the other hand, Mason and Wheeler, 370 after injecting Microcystis extracts, observed markedly congested livers and dilated right hearts, but little generalized venous congestion Their terminal dinical findings included severe anemia and reduction in total serum proteras Small amounts of the toxic extract ·so TABLE VI - Animal Intoxications by Phytoplankton Source Year Locale Victim sheep, horses, dogs, Pigs cattle, horses, hogs cattle cattle 1 Francis 2 Porter 3 Porter, Arthur, Stalker 4 Ballad 5 Cotton 6 Fitch 7 Howard, Berry 8 Wilmot Enterprise 9 Gillam 10 Gillam 11 Gillam 1878 1882-83 1884 1900 1914 1918 1924 1925 1925 1925 1925 Australia Minnesota Minnesota Minnesota Michigan Minnesota Ontario South Dakota Alberta Alberta Alberca steer and cow sheep, hogs, chickens cattle stock hogs hogs horses, cattle, hogs, poultry, wild birds livestock sheep, cattle carile Organis Nodularia spumigeria Blue-green algae Glocotrachia echmulara Aphanizomenon Anabarna Corlosphaerium, Anabaena Anabaeng Blue-green algae Blue green algae Blue green algae Blue-green algae 12. Woodcock. 13 Steyn 14. Fitch 15 Fitch. 16, Fich 17. Fitch 1925 1927 1930 1930 1933 1933 Michigan South Africa Minnesota Minnesota Minnesota Minnestoa Blue-green algae Blue-greca algae Macrocystis Blue green algae Alicrocystis Microcystis, Anabaena, Aphani" zomn0n Microcystis, Anabara Blue green algae THE ROLE OF ALGAE AND PLANKTON IN MEDICINE pigs sheep, lambs, chickens turkeys, ducks, geese, cows, 18 Titch 19 Ilinderion Pigs, horses cattle 1933 1933 Michigan Finland stock ·SONTCE Year Locale Victims 20 Deem, Thorpe 21. McLeod 22 McLeod 23 Steyn PLANKTON 1939 1938-40 1939 1942 Colorado Manitoba Alberta South Africa Organism Anabaena, Microcystis Blue-green algae Blue-green algae Blue-green algae fish, birds, horses animals animals horses, catcle, sheep, mules, hares, water birds cattle, livestock sheep sheep, cattle horses, calves, Pts, cat cattle WAI IN Macrocystos Blue-gecca algae Blue-green algae Blue-green algae Blue-green algae Date... 24 Louw 23 Quin (Remer) 26. Ophel 27. McLeod, Bondar 28 Brandenburg 29 Mullor, Wachs 30 Prescott 31 McLeod 32 Stewart, Barnum, Henderson 33 Olson 34 Olson 35 Olson 36 O'Donoghue 37 O'Donoghue 1942-43 1943 1945 1945 1946 1948 1948 1949 1949 1950 1948-51 1945-50 1950 1950 South Africa Montana Australia Manitoba North Dakota Argentina Bermuda Manitoba Ontario Canada Minnesota Manitoba Alberta Alberta cattle animals cactle ducks livestock Livestock JAIPUR. O Price......... Accn. No... .. Aphanizomenon Blue green algae Microcystis, Anabarna Blue-green algae Blue green algae Blue green algae Microrystus Macrocystis 3 Ayyuan COW Pigs, turkeys, geese, dogs, chickens, horses, gulls, ducks, blue-sinch horse, spaniels 39. McLeod, Bondar 1951 Manitoba Apbanizomenon, Anabaena, Microcystis 51 ·52 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE caused systolic arrest of the isolated perfused frog heart, but the uterus of the virgin pig was unaffected. Oxygen consumption of various mat tissuc slices also remained unchanged Steyn259-260 reported marked blistering of photosensitized animal skins, as well as extensive involvement of the liver and the central nervous system Post-mortem examinations showed the lungs full of blood, the spleen enlarged; and the liver engorged and dark red to black In chronic cases, the liver was either yellow and soft, or dark and brittle Ashworth and Mason272 were able to produce generalized cellular damage by administering Alecrocystis extract to rats. Especially severe injury was noted in the parenchymal cells of the liver. Succes sive stages included acute parenchymatous, hydropic, and fatty degeneration, followed by central lobular necrosis If death did not ensue, complete restoration of normal lobular architecture --without fibrosistook place in thirty days The toxic alterations were comparable to those produced by chloroform, phosphorus, and epidemic hepatitis Similar acute hydropic and necrotic changes were also seen in the heart and kidneys, while microscopic hemorrhages and edema sometimes occurred in the lungs Ashworth and Mason felt that rapid death was due to shock and arculatory collapse, while delayed demise was consequent to hepatic insufficiency, renal failure, and diffuse cellular damage Stewart and his colleagues 268 reported fresh pulmonary hemorrhages and hepatic and cardiac lesions in Ontario cattle dying after ingestion of Anabaena and Microcystis scum They found the same material lethal to pigeons and rats Pathological findings from Alberta were "dilatation of the heart" and "mottled Irver" As mentioned earlier, many non-algal poisons have been wrongly suspected in several cases, botulinus antitoxin was administered, to no avail Solutions of sodium nitrite and sodium thiosulfate given intravenously assertedly produced recovery of some poisoned animals Although the various outbreaks are now known to have been algal in origin, the specific nature of the toxins involved has certainly not been clear Fitch234 determined that the substances from Aficrocystis and Anabaena are nonvolatile, heat-stable when dry, stable in solution at ordinary temperatures, but unstable when heated to 100°C They are not electrolytes, alkaloids, or toxalbumins They are organic componds of low molecular weight, some what resistant to add and alkaline solutions The substances apparently disappear when the algae putrefy Mason and Whecler270 nere able to inactivate Alicrocystis extracts partially by heating with acid or alkali. Steynco dlamied that growing algae are poisonous, that they discharge their poisons into the water when they dic, that the decomposition of ·PLANKTON 53 masses of dead algae decreases the toxicity, and that boiling the contaminated water does not reduce its toxicity He thought that Microcystis contains two active principles a potent fucoin-type hepato-neurotoxin, and a fluorescent photosensitizing pigment, phycocyan Wheeler, Lackey, and Schott273 determined that Microcystis is less porsonous when fresh than when frozen or dried They found the toxin dialyzable, soluble in alcohol, and able to withstand autoclaving in neutral solution It also survives, unfortunately, the laboratory equivalent of a water purification processalum coagulation, chlorination, and filtration It is adsorbable on carbon, but only when this is used in comparatively large amounts According to McLeod and Bondar,282 in a Manitoba outbreak associated chiefly with Aphanizomenon (and some Microcystis and Anabaena), the toxic material was in the plants and not in the water. They found the toxin stable in the plant, not destroyed by air drying at 37°C, by freezing, or by ultraviolet irradiation Shelubsky257 claimed in 1951 that the Microcystis toxin us all-labile In contrast with other investigators, Louw has reported isolation of two alkaloids from Microcystis The first is an in. active one with a probable formula of GoH1NO, The other, without stated molecular structure, was isolated as a picrate with a melting point of 165°C It is supposedly the active hepatotoxın, and it has been effective in experimental rats Somewhat similat findings with marine algae from the South Pacific have recently been described by Habekost, 274 who 150- lated substances toxic to mice C Human Intoxications Algal poisoning has not been limited to animals During the past 25 years, phytoplankton has also been incriminated in human reactions These have presented as three general clinical pictures dysenterial disorders, systemic allergic reactions, and local allergic eruptions Table VIII sum. mnarizes the reported human cases 275-281 (Poisonings from ingestion of plankton-nourished fish will be discussed separately later ) The epidemic intestinal disorders of the early 1930's involved many thousands of people, in areas as widely separated as the National Parks in the Northwestern United States 275 and Charleston, West Virginia, 276 Syınptomatology consisted of nausea, vomiting, diarrhea, and sometimes abdominal pain. It was self-lemitrag and usually resolved in a few days. These outbreaks were of no known speafic etiology, but, significantly cnough, were always concomitant with the presence of extensive algal blooms in the local water supply Doubtless many other similar outbreaks have been neither studied not reported ·54 TABLE VIII - Human Reactions Associated unh Phytoplankton Auribor Year No Cases Locale Etalogy 1. Spencer 1930 95 Symptomatology Nausea, vomiting, duarthea, abdominal pain Same algat bloom 2 Tisdale 1930 8,000-10,000 algal bloom 3. Olson 1930-31 National Parks, Northwest US Charleston, West Virginia Anacostua Reservoir arca near Washington, DC North Lake, Wau. kesha County, Wis Dysentery-like epidemic Igal bloom 4 Horse 1934-35 1944 57 yr old man Oscillatoraceae 3. Hesse 2945-46 old 39 1 woman Lake Keesin, Wau. kesha County, Wis Oscillatoraceae 6. Sams 1947 67 Itching, conjunctivitis, blocked nares, bronchial astham Swollen eyelids, blocked nose, generalized urticarial eruption Acute urticanal-papular "sesbachers' cruption" Acute urticarial.papular eruption Positive skın tests Erythematous papulovesiculas dermatitis of exposed arcus Southern coast of Fonda Gulf of Mexico THE ROLE OF ALGAE AND PLANKTON IN MEDICINE manne algae 7. Ayres . 1948 1 "red tide" 8. Heise 9 Cohen. 1951 1953 Office patients 6 ye. old child Lake Carey. Pa Microcytes Anabarna (phycocyanın) ·PLANKTON 55 Heise277 described two cases in which systemic hyperergic pictures appeared following swimming in a Wisconsin lake In the one, the chief manifestations were itching, conjunctivitis, blocked nares, and bronchal asthma; in the other, swollen eyelids, nasal stuffiness, and a severe generalized urticarial eruption were the presenting findings in both cases, Oscillatoraceae (Myxophyceae) were proven to be the causative organisms Herse also obtained many positive skin reactions to Microcystis among his office patients Local allergic phenomena have been recorded more frequently of late Sams278 described an acute urticarial papular eruption-locally known as "seabathers” eruption"—in swimmers off the southern coast of Florida Ayres, 280 in 1948, had a patient who swam in the Gulf of Mexico during the "red tide," then developed an acute urticarial papular eruption And as recently as 1953, Cohen and Reif281 reported phycocyanın, the blue pigment in Anabaena, as the cause of an erythematous papulo-vesicular contact dermatitis in a six-year old child who had bathed in a Pennsylvania lake Microscopic algae have also been assigned a more indirect role in some human disease states Marian,282 reported on his and Redaelli's work with Protosheca portoricensis. This alga, isolated from the stools of tropical sprue patients in San Domingo, produced a transitory local caseous granulomatosis in guinea pigs when injected intramuscularly or intraperitoneally No specific sensitization or immunization could be achieved even with repeated injections A related alga, Blastocystis enterocola, was isolated by Newiadomskı from human, tat, and mouse feces, and was considered by him capable of passing through a Seitz filter Marianı could not confirm the filtrability of Blastocystis, nor did he prove any pathogenicity Neither could he corroborate Szendy's assertion that algae contributed to infections in metapneumothorax Although algae have been isolated from the skins of animals, none has yet been reported resident on the intact human epidermis 283 However, in 1946, de Almeida and his colleagues284 in Brazıl isolated Chlorella from lesions of three patients with mycoses The first had perirenal actinomycosis; the second, pulmonary and lingual paracoccidiosis (brasiliensis), and the third, lymphatic and dermal paracocadrosis. The Chlorella was not considered a contaminant, but the authors debated whether it might be pathogenic either alone or in association with the fuag! They wondered, too, whether the pathogenicity of the fungi might not even have been Iessened by the action of "chlorellin," an antibiotic substance isolated from ·56 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE Chlorella The activity of this material (effective against Gram positive and negative bacteria) has been shown to result from photo-oxidation and splitting of long carbon chains of the unsaturated fatty acids in Chlorella,285 The antibacterial substance dubbed "phyceine" by Lesage®2 is probably similar to "chlorellin." Related antibiotic agents in Stocbocorcus and Protosyphon have been reported by Harder and Opperman 286 An interesting medico-legal application of planktonic distribution relates to the diagnosis of death by drowning 287-200 Attempts have been made to prove death resulted from submersion by demonstrating algae in the lungs or the gastro-intestinal tract It has even been proposed that the time and exact place of the drowning might be determined from the nature and type of algae present It is clear that inherent faults exist in this method, since the algae may be dificult to identify, and since algal distribution in any area may vary from hour to hour. A striking illustration of indirect algal effect is found in human intoxications resulting from ingestion of certain fish Since 1530, there have been recorded many episodes of fish poisoning involving thousands of people 291-301 Fifty-eight of the best documented have been summarized in Table IX (pages 57-59) The most familiar intoxications are the acute "paralytic" shellfish poisonings resulting from eating mussels and clams The manifestations are primarily neurologic. prickling and numbness of the mouth and fingers, ataxıc gait, muscular incoordination, ascending paralysis and ultimately respiratory failure Death may take place in two to twelve hours Outbreaks have occurred in many parts of the world, predominantly during the summer months For years there was considerable groping-superstitious, epidemiological and toxicological—for the underlying cause of causes It was not until 1928 that Lindner208 suggested they might reside in the food supply of the fish This has indeed been proven since then, especially through the work of the groups at La Jolla and the Hooper Foundation 292,294, 295,297 They showed that intoxications result only when shellfish ingest large quantities of the marine dinoflagellate Gon. yaulax (Dinophyceae) These algae can grow so profusely during warm weather that the sea turns a deep rusty red during the day and brightly phosphorescent at ought. It has been demonstrated that the Gonyaulax contains an alkaloidal type of poison resembling in effect such substances as strychnine, muscarine, and aconıtıne. It is heat-stable in acid or neutral solution, but it is slowly destroyed by boiling with alkalı It is readily soluble in water and alcohol, but insolublc in ether or chloroform One-millionth of a gram will kill a ·TABLE IX-luman Personing Follouing Ingestion of Fish PLANKTON Time of Year No Sick No Dead Awtbor 1 Martyr 2. Kaempfer 3 Anderson 4, Vancouver 3 Kraus et al Year 1330 1690 1776 1793 1799 Fish Implicated Coguatera group Tetraoden (puffer) Mussels Mussels 3 200 1 150 Locale Virgin Is Japan Central Pacific British Columbia Penil Way, near Sitha, Alaska Leith, Scotland Victoria Dock, England Tralee, Ireland Tralce, Ireland Liverpool, England Mendocino, Calif June August 1 6 Combe 7. British Medical Journal 8 Crumpe 9 Crumpe 10 British Medical Journal 11 Meyer ec al 1827 1897 1872 1872 1872 1870(?) Mussels Mussels Mussels Mussels Mussels Mussels 30 (?) 3 1 2 ܝ ܝܢ ܚ ܐ ܝܕ Oct August "Band" of Indians 4 Mussels Mussels Mussels Mussels 1 19 3 3 7 1 5 12 Varchow & Wolf 13 Schmidtmann 14 Permewan 15 Cameron 18 Todd 17 Thesen 18 Carr 19. Rolfe 20 Necter & Ribadeau.Dumas 21 Neale 1885 1887 1888 1890 1891 1901 1903(?) 1904 1907 1909 Wilhelmshavea, Germany Occ Wilhelmshaven, Germany Sept Liverpool, England August Dublin, Ireland June Sept Christiania, Norway Timbe: Grove, Calif Ayonmouth, England June Calaos, France May Basty, South Wales August 1 May Mussels 2 > Mussels 5 12 2 13 19 1 2 1 57 ·TABLE IX (continued) - luman Poisoning Pollowing Ingestion of Fish 58 Tone of Year NO Sick No Dead Locale Author 22. Taharu et al. Yray 1911(?) Japan Implicated Pufferfish (Fugu Poison) 1915 Santa Cruz, Calif 4 0 23. Meyer er 15 24 California Stare Board of Health 25. Kofoid & Vaughan 26 Olson 13 2 1917 191817) 1924 Mussels Mussels Eels & Fish Livers 1 800 appr 6 27. Sommer & Meyer 1925(?) 28 Sommer & Meyer 29 Sommer & Meyer 30 Samme & Meyer 31 Sommes & Meyer, 32, Olson 1927 1929 1930 1932 1932-33 Santa Cruz, Colf July La Jolla, Calif. September Koenigsberg Haff, East Winter Prussta, Germany Autumn Subtropical American Waters Central Calif Coast July Central Calif Coast Summer Central Calif Coast July Central Calif. Coast June & July Koenigsberg Haff, East Prussia, Germany Gokasho Bay, Tobz, Nov, Dec Japan Jan., Feb Central & North Calif & July & Sep- Oregon tember Oregon October Alaska July Southern Calif Coast May Bay of Fundy, Nova July Scotia Local Fish (Ciguatera) Mussels Mussels & Clams Mussels Mussels & Cams Iish or Fish Livers 102 56 2 37 125 THE ROLE OF ALGAE AND PLANKTON IN MEDIANE 33. Sommer & Meyer 1933-34 34. Sommer & Meyer 1933 Mussels 19 1 33 Sommer & Meyer 36. Sommer & Meyer 37. Sommer & Meyer 38 Murphy 1933 1934 1936 1936 Mussels & Clams Mussels Mussels Mussels 3 12 3 39. Halstead.. 1938 Amberjack 1 ·Pish Implicated Time of Year Summer No Sick No Dead Author 40 Wolczynska & Conrad Year 1939 Locak Bruges-Zeebrugge Canal Belgium Calif Coast Lake Ysmen, Sweden PLANKTON 41 Olson 42. Olson 1939 1940 Eels and other fish livers 43. Halstead 44 Lively 45 Lewis 46 Halstead 1941-45 1944 1944 1944 Small epidemic 400 4 6 38 || Ili Septernber August November & December Red Snapper Red Snapper Sea Bass 47 Von Fraenkel & Krick 49 Medcoff 49. McCann 50 Cohen et al 51 Ross Micronesia Gilbert Is, Pacific Line Is, Pacific Honolulu, Hawaii, Mid. way & Christmas Is Mariana Is, Pacific New Brunswick, Canada Palmyra Islands, Pacific Mariana Is, Pacific Fanning Is, Pacific 1945 1945 1945-46 1946 1946-47 Barracuda Mussels Red Snapper 30 28 32 51 95 14 Fish-type not stated 32 Medcoff 1948 2 2 53 Sapeika 54. Olson Le Boules, Quebec, St Lawrence River, Can Capetown, South Africa Lahe Ysmen, Sweden 1948 1948 Many 11 1 Mussels Eels and other fish lovers Black Moray Eel 2 59 Halstead $6 Martin » Halstead 58 Tennant et al 1949 Saipan, Pacific 1950-51 Johnston Is, Pacific 1953 Marshall Is, Pacific 1954 Metus Beach, St Lawrence River, Can May Summer March Black Moray Ecl Mussels 57 20 6 7 2 65 ·60 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE mouse on injection, while a few milligrams orally can be fatal to a human being, especially when taken on an empty stomach It has been estimated that 4,000 algal organisms are required to produce human toxicity. The shellfish store the poison chiefly in their "livers" or digestive organs; thus the "dark meat" of the infested bivalves is much more toxic than the "white" portions The fish themselves are unaffected by the toxin, and they will in fact excrete it in several weeks if Gongaulax disappears from the macine diet.297,298,302,303 Post-mortem examinations of animals and human beings who have died of acute paralytic shellfish poisoning have, surprisingly enough, revealed no structural changes However, in chronically poisoned experimental animals, destinct alterations are noted in the medullary ganglion cells and in the large cells of the central horn of the spinal cord.802 The Golgi apparatus of small and medium-sized spinal ganglionic cells is also damaged Neural mitochondria remain normal, but those in the convoluted tubules of the kidney show definite damage In concert with the latter finding are the clinical nephropathies reported by the French in human shellfish poisoning 394,305 It is probable that other algae besides Gonyaulax contain toxins Some have been associated with Ceratium and Prorocentrum, but na direct extraction or isolation has yet been accomplished. It is also quite logical to expect that fish other than muussels and clams could have improper planktonic relations and thus transmit their toxins to predators This is indeed so, and-except for the puffer fish and possibly barracuda--it is likely that poisonousness is directly dependent upon periodic ingestion of noxious plankton, cather than being due to inherent toxicity Planktogenic fish toxicity can occur during any time of the year, while puffer and barracuda may be most lethal during the reproductive season Epidemios have been unrelated to spoilage or bacterial infection Poisonous fish are found through the world. They are most numerous in the warm seas (especially in the Caribbean and Central and South Pacific), but even certain arctic sharks have been reported to be toxic Duro ing World War II, fish poisoning represented quite a problem, since epidemics were encountered wherever American forces were stationed in the Pacific theater The Japanese, too, estimated that over 400 of their military men died in Micronesia from cating toxic fish Halstead and Lively300 have made a partial lasting of fish most often found to be poison- Qus: surgeonfish, triggerfish, pompano, porcupine fish, wrasse, snapper, filcfish, surmullet or goatfish, moray cel, parrot fish, sta bass or groupet, ·PLANKTON 61 barracuda, and puffer They have divided fish poisoning into four clinical groups, the last three of which are probably planktogenic: 1. Tetraodon (puffer) poisoning 2 Gymnothorax (Moray eel) poisoning 3 Cigualera 4 Scombroid poisoning Tetraodontoxin is the only ichthyosarcotoxin about which anything specific is known Japanese studies indicate it is a white hygroscopic powder, soluble in water but not in the ordinary organic solvents Although it is thought to have a formula of C16H1N016, little else is known except that it is not an alkaloid or a protein or a protamıne It is a most potent poison, with symptoms occurring within thirty minutes of ingestion Insteally there are numbness and tingling of the face and extremsties, then nausea, vomiting, headache, dizziness and overwhelming weakness Next come speech impairment, dyspnea, and generalized muscular paralysis Death from respiratory paralysis may occur in 1-24 hours. The mortality rate is about 60 per cent In Gymnothorax poisoning, similar neurotoxic manifestations are seen at first, but then the patient develops motor incoordination and violent convulsions, laryngeal spasm, coma, and respiratory paralysis Here the mortality rate is only about 10 per cent, and, in the non-fatal cases, major symtomatology disappears in about ten days With Cigualera (originally considered only Caribbean), the attacks are milder than with Gymnothorax and usually come on more slowly The chief symptoms include sensory disturbances, myalgias, arthralgias, and severe weakness Only 2-3 per cent of the victims die, but recovery may take many months Scombroid poisoning results from eating various tropical tuna-type fish The manifestations are mostly hestamine-like headache, flushing of the face, con. junctivitis, giant hives, erythema, and gastric upset Recovery usually takes place in 8-12 hours Haffkrankbeit-erroneously designated "Haff's Disease" by some enthusiastic eponymıst--was first described in 1924295 among fishermen la Koenigsberg Haft ("harbor") in East Prussia It follows contact us ingestion of fish (primardy eels) and fish livers It is characterized by a sudden onset of weakness, severe myalgia, myoglobiquria, and some digestive upset Pathological findings indude fatty degeneration of the liver and kidneys, and darnage to the anterior horn cells of the spinal cord Over 1,000 people have been reported afilicted, some fatally. Similar episodes have been recorded at Lake Ysmen in Swedea Originally Haft- ·62 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE krankheit was considered to be caused by arsines in sewage. However, the correlation of outbreaks with climatic changes, and the predominantly neurotoxic nature of the manifestations, suggests that this disease, like the several other categories of fish poisoning, is dependent upon periodic planktonic intoxication No method of testing suspected fish is known, other than feeding it to animals (kittens have been a favorite) Neither does any specific antidote exist Treatment has consisted primarily of rapidly emptying the gastrointestinal tract by mechanical means, apomorphine, and laxatives Administration of charcoal as an adsorbent may be helpful Also important are supportive measures such as stimulants, parenteral fluids, oxygen, and artificial respiration Intubation and tracheotomy may be necessary with respiratory distress Anti-convulsants and opiates may have their indica. tions, while some assign a special merit to intravenous calcium gluconate To us it would seem logical also to employ the adrenal steroids, either to combat anaphylactoid reactions or to diminish end-organ response to toxins Anti-histamine agents, too, might play a part in treating the Scombroid type of poisoning Doubtless the wisest management of fish poisoning consists of avoiding those fish with a known toxic potential, and, in the case of shellfish, avoiding them when the Gonyaulax or other suspected algae are flourishing V. DISCUSSION AFTER COMBING THOUSANDS of articles on algae and plankton, we are in firm agreement with Conway Zırkle, e 306 who laments the fact that our age of specialization has produced a "splintered learning" He points out that many important developments are hopelessly buried because of the compartmentalization of knowledge, the spe. cialized jargons of the various sciences, and the overwhelming volume of new publications Because of a limited lateral diffusion of information, many workers remain unaware of pertinent findings even in kındred fields This is especially true in the realm of medicine with the tremendous mass of studies that physicians must continually digest, their meager knowi: edge of algae and plankton is readily understandable The rare medical ·DISCUSSION 63 articles on these subjects have usually dealt only with some special point of interest or a courtesy seference has appeared in a botanical, nutritional, or public health journal The present assemblage of data brags mto focus much diversified material, a good portion of which deserves closer integration into the Geld of mediane Interest in algae and plankton has varied over the years from ancient times, the predominant emphasis was on the searreeds, at the sum of this century, it shifted to zooplankton, and, since World War II, it has centered chiefly about phytoplankton. The traditional medianal use of seaweed has declined sharpiy, as has its role as a human nutrient Its value today resides primarily in the contained minerals and vitamits, and its greatest utility is as fertilizer and animal provender There is also much merit in the alginates, and a good potential exists for other seaweed derivatives such as the antibiotics and anticoagulants Consideration of plankton has for the most part been focused on its use as food 807-809 Zooplankton is nutritionally far richer than seaweed, but it does not possess the significance its more enthusiastic proponents confer upon it Despite the fact that the Thats?? have reportedly collected 5,000 tons sa one year, it should not be forgotten that there are tremendous variations in quantity and quality of yield in different oceans and at different seasons While zooplankton may be considered an excellent emergency ration and food supplement, its extensive use as a food staple awaits the development of far more efficient collection techniques Since it is still much too early to envision extensive in viro photosynthesis 197 without the intermediary of plants, it is the phytoplanktonic organisms which offer the most exciting prospect for nutritional guin They not only supply controllable proportions of protein, fat, vitamins, minerals, and calories they can also be cultured selectively on a large scale basis The marvelous adaptability of algae like Chlorella gives man another round in the cracial Malthusian battle of population versus food supply in the past three centunes, the world population has risca from a half billion to two and one half billion 23 if fission and fusion do not permanently put an end to both mankind and its problems, the population will be four billion in 1980, and eight billion in 2050. Selective fresh. waler mass culture of organisms (we suggest the term "aquaculture" as more descriptive than "hydroponics" and less restrictive than "masi. culture"») like Chlorella in speaalis built plants hold great promise for Jarge quantities of valuable foodstuffs it is soore practical than "minuring” the sea with salts and minerais"three-dimensional farmung"-o increase ·64 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE the growth of phytoplankton, whether one hopes to harvest the phytoplank. ton as such, or whether it is planned to support a greater fish population The latter is, in fact, a wasteful process, since it has been calculated that 2 pound of codfish ultimately represents 100,000 pounds of marine phytoplankton 17 Before phytoplanktonic feeding becomes commonplace proced. ure, there will have to be long-term human experiments, with careful observation of possible toxicities and biochemical changes, especially with respect to electrolytes, hemic elements, and proteins It is also a certainty that a re-education of food tastes will have to take place before the acceptability of planktonic foods reaches a high level 310 Some other applications of phytoplankton reach almost into the realm of science fiction Bassham, 311 at the University of California, has been working for the US Navy on adapting microscopic algal cultures to control the oxygen and carbon dioxide in atomic submarines which would be submerged for long periods It has been calculated that one kg fresh weight of Chlorella (equal to 100 liters of growing algal suspension) can easily supply the 25 liters per hour oxygen required by a 70 kg man, as well as utilize the carbon dioxide exhaled by him in a similar fashion, the Department of Space Medicine of the Air Force 2 is considering using algae in space ships, with the intriguing extra notion of complete recycling of biological elements the algae are also to be used as food by the space. men, while their excreta will serve as nutriment for the algae! From the strictly medical standpoint, the number of disease syndromes attributable to algae comes rather as a surprise and raises some interesting questions The most obvious afflictions are the allergic dermatitides resulting from bathing in water contaminated with algae Also under. standable are the respiratory irritations from water-borne or inhaled algae; in fact, many cases of allergic rhinitis (and its corollary, chronic sinusitis) in coastal areas mught be traceable to algae instead of being blamed on "damp nate" or that handy favorite of allergists, "dust." Then there are the nephropathies, and the striking epidemic gastroenteritides, so blandly ascribed to the 24-hour or 48-hour virus" By all odds, the most dramatic algal intoxications are the acute neurotoxic and myotoxic ones typified by paralytic shellfish poisoning and some other ichthyosarcotoxicoses. Although pitifully meager long-range studies have been made, it seems incredible that such potent toxins should not produce related chronic sequelae in victims who have survived acute attacks, or who have ingested smaller amounts of toxin over prolonged periods One cannot help but wonder hether chronic algal intoxication ·DISCUSSION 65 may not play a part in causing some of our idiopathic neuropathies such as multiple sclerosis, the muscular dystrophies, and amyotrophic lateral sclerosis Interestingly enough, very recently a US Public Health Service team has reported312 an extensive survey of amyotrophic lateral sclerosis among the Chamorros of Guam and other islands of the Mariana group They made the study because the incidence of the disease there is 420 per 100,000 population, a rate 100 times higher than in the rest of the world, at any one time, one per cent of the adults are affected, and eight to ten per cent of adult deaths are produced by it No cause was discovered, except perhaps some tenuous genetic relationship to the Chamorro tribe The investigators have apparently given little consideration to the water contacts or fish-eating habits of these people it may be more than co. incidence that epidemics of fish poisoning—likely due to toxic phytoplankton-have been particularly plentiful in the Marianas That algae have not been implicated in the nerve afflictions may be due to the difficulty in demonstrating them, or simply to their not even having been considered It is interesting, if hazardous, to speculate also on a possible algal role in another neurologic disease-acute poliomyelitis The viral etiology is certainly proven beyond any doubt, yet the epidemiology is anything but clear, Is it possible that a symbiotic virus-alga state exists, favoring viral transport or even viral multiplaction? The thought is suggested by the fact that the highest incidence of poliomyelitis is during the warm summer months, when algal blooms are most prolific, also, that infection occurs frequently following swimming In the latter case, the victim, besides ingesting virus, might also decrease local tissue resistance by contact with algal toxins However, we are not yet prepared to suggest Chlorella instead of monkey kidney as a medium for culturing polio virus! A more definite chronic entity is "swayback,"82,83 the neurologic disorder seen in lambs fed on seaweed Whether a toxin disturbing copper metabolism resides in the seaweed itself, or whether it originates in con. taminating microscopic ae, has not been established Also worth considering are the hepatitis and cirrhosis seen in animals suffering from "waterbloom" toxicity Their occurrence raises the question of whether all human hepatitis classified as "viral" is really so Certainly these various diseases indicate that much might be gained from studying the results of controlled administration of toxic algae to animals ·BIBLIOGRAPHY 1. Allen, WE The primary food supply of the sea, Quart. 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Ribonucleic acid content of cells and growth rate, Biochim et biophys acta 8.125, 1952. 217 Szafarz, D, and Brachet, J Le rôle du noyau dans le métabolisme de l'acide sibonucléique chez "Acetabularia mediterranea," Arch, internat. physiol. 62 154 (Feb ) 1954 218 Goryunova, S V Slızıstye veshchestra (Macias of Oscillatoria) Izvestiga Akad nauk SSSR, Ser bio! no 1, P 29, 1950 (In Ressian) 219 Frank, B Uber den experimentellea Nachweis der Assimilation freien Stickstoffs durch erdbewohpende Algen, Berichte Deutsche Botan Gesellsch 7 34, 1889 220 Drewes, K Uber die Assimilation des Luftstickstoffs durch Blaualgen, Zen tralbl f Bakt Abt 2, 76 88, 1928 221 Watanabe, A Production in cultural solution of some amino acids by the atmospheric attrogen-Exing blue-green algae, Arch Biochem 34 50, 1951. 222 Douin, R Sur la fixation de l'azote libre par les myxophycées endophytes des cycadacées, Compt rend Acad dst. 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·B18LIOGRAPHY 77 235, Combs, G. P. Algac (Chlorella) 35 « tource of putrientes for the fuck, Science 116 453 Oct 24) 1932. 236 Food of the furure, 1.AMA 156 21 (Advert Oxt 30) 1954 237. Chamberlan, y } Effects of algac on water supply, Unsversaty of Queense land, Brisbane Department of Chemistry Papers 1, no 29, 1948 80 P 60 plates 238 Howard, N), and Hessy, A E Algal nursances surface waters, Cansd Pub Health 24-377, 1933 239 Upspenskaya, V 1: Izuchenie fiziology pelanus Oscillatona spleodida Grew Oscillatoria agardhu Gossy slazı obrazovaniems sakepleniem zapukhov i prevkusov vody | Studes os physiology of nuttatus of Oscillatona splendide Giew and Osallatonia agardhu Gom and its relation to fosaation 25d concentration of odors and taste of water I, Mikroblogiya 22(3) 517, 1993 240 Puerta Palacios, MC freeoorganismos acuáticos, problemas técnicos y sansarios causados por su presencia en los abastecimientos de agua potable, 1949, Editorial Granamerica, Medellin 163 2413 Prescott, GW Algae of the western Great Lakes ssea, exclusive of desmids and diatoms, 1931, Cranbrook last of Science, Bloomiseld Hills, Michs 946 P (Cranbrook Inst of Science Busi no 31 ) 2416 Forest, HS Handbook of algae with special reference to Tennessee and the southeastern Vosted States, 1934, University of Tennessee Press, Knoxville 467 p 242 Standard methods for the examination of water, sewage, and seadustrial wastes. prepared and published jointly by American Public Health Association, American Water Works Association, Federation of Sewage & Industrial Wastes Association, 10 ed 1955, Am Pub Health Assn, New Yozk 522 p 243 Smith, G M Cryptogamıc botany, vol 1, Algae and fungi, ? 1935, AlcGraw Hill, New York 546 244 Ingrasi, W M, and Tarzwell, C M Selected distrugraphy of publicacions relating to uadestradle offects upox aquatic life by algrader, insekticidas, weedecides, Wash, US Govt Print # 1954 p 35 (US Pble Health Service Publ 4003 (US Public Health Service Pubic health bibliography Serres 13) 243 Hassas, J E Report on 13-fouting stsearch, 1942-44, Iron & Steel last (London) 194100 2) 297, 1946 246 Pyefiach, K A Methods of assessment of ann fouling compongonis, Iron and Steel Inst (London) 132(60 2) 229, 1945 (Paper no 16/1946. Car- FOSION Corriepittee) 247 Zehnder, A Uber den Esnuss aarsbiotischer Stoffe auf das Wachaturs on Grünalgen, Experientia (Basel) 7 99, 1931. 248 Foter, MJ. Palmer, CM, and Malones, T I Antialgal properties of VARIOUS satsbiotics, Antibiobes & Chemothes 3 305. 1933 249 Matheson, R. and Hinman E H A seasonal study of the planctos of Spring fed Chura pool verses that of a las pocate to semi permanent wood land poot is relation to mosquito beredandi Am Hy8 11 174, 3930 250 Boswit, K Algal Aott and mosquito $294*, Repriat, Science and culture 8 189, 2942-2533 ·78 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE 251 Effects of radioactive substances on the anacrobic digestion of sewage sludges, US Atomic Energy Commission. Technical Information Service Report No Wash—129, Sept 11, 1952. P 63 (Sanitary Engineering Conference, South District Filtration Plant, City of Chicago). 252 Coopey, RW Radioactive plankton from the Columbia River, Tr Am Microscop Soc 72 315, 1953 233 Francis, G Poisonous Australian Jake, Nature (London & New York) 18 11 (May 2) 1878 254 Fitch, CP, and others "Water bloom" as a cause of poisoning in domestic animals, Cornell Veterinarian 24 30, 1934 253 Cotton, H L Algae poisoning. Am J Vet Med 9 903, 1914 256 Gullam, WG The effect on livestock of water contaminated with fresh water algae, J Am Vet Med Assoc 20 780, 1925 257. Grant, GA, and Hughes, E O Development of toxicity in blue green algae, Canad J Pub Health 44 334, 1953 258 Steyn, G Poisoning of animals by algae on dams and pans, Farming in South Africa 18 489, 1943 259 Steyn, G Vergifuging deur slyk (algae) op damme en panne, South African M J 18 378, 1944 260 Steyn, G. Poisoning of animals and human beings by algae, South African J SC 41 243, 1945 261 Deem, AW, and Thorp, F Toxic algae in Colorado. J Am Vet Med Assoc 95 542, 1939 262 McLeod, J A, and Bondar, G F A case of suspected algal poisoning in Manitoba, Canad J Pub Health 43 347, 1952 263 Louw, PGJ The active constituent of the poisonous algae, Mycrocystis toxica Stephens, South African Industrial Chemist 4 62 (April) 1950 264 Quin, A H Sheep poisoned by algae, J Am Vet Med Assoc 102 299, 1943 265 Olson, T. A Toxic plankton, Water & Sewage Works 99 75 (Feb ) 1952. 266 Brandenburg, T 0, and Shugley, F M "Water bloon" as a cause of poison. ing in livestock in North Dakota, J Am Vet Med Assoc 110 384, 1947 267 Prescott, G W Objectionable algae with reference to the killing of fish and other animals, Hydrobiologia, acta hydrobiologica (The Hague) 11, 1948 268 Stewart, AG, Bamum, DA, and Henderson, J A. Algal poisoning in Ontario, Canad J Comp Med 14 197, 1950 269 Donoghue, J G, and Wilton, GS Algai poisoning in Alberta, Canad J Comp Med 13 193, 1951 270 Mason, MF, and Wheeler, RE Observations upon the toxicity of blue green algae, (Am Soc Biol Chemists 36 Ann meeting. Boston) Federa. tion Proc 1(12) 124, 1942 271. Smi's ID Experimental cases of algae poisoning in small animals, South African Industrial Chemist 4 66 (April) 1950 272. Ashworth, CT, and Mason, M F. Observations on the pathological changes produced by a toxic substance present in blue-green algac (Afycrocystis aeruginosa), Am J Path 22 369, 1946 273 Wheeler, R E, Lackey, J B, and Scholl, S A contabution on the toxicity of algae, Pub Health Rep 57 1695 (Nov 6) 1942 ·BIBLIOGRAPHY 79 274 Habekost, RC, Fraser, 1 M, and Halstead, B W Observations on toxic marine algae, J Washington Acad Sc 45 101, 1955 275 Spencer, R R Uausually mild securring epidemic simulating food infection, Pub Health Rep 45 2867 (Nov 21) 1930 276 Tisdale, E S Epidemic of intestinal disorders in Charleston, W Va, occur. ring simultaneously with unprecedented water supply conditions, Am J Pub Health 21 198, 1931 277 Heise, H A Symptoms of hay fever caused by algae, J Allergy 20 383, 1949 278 Heise, H A Symptoms of hay fever caused by algae, II Mycrocystis, Ann Allergy 9 100, 1951 279 Sams, W M Seabather's eruption, Arch Dermnat & Syph 60 227, 1949 280 Ayres, S Discussion of W M Sams' Seabather's eruption, Arch Dermat & Syph 60 236, 1949 281 Cohen, S G, and Reif, C B Cutaneous sensitization to blue-green algae, J Allergy 24 452, 1953 282 Mariani, PL Ricerche sperimentali intorno ad alcune algae parassite dell'uomo, Boll d Soc ital di microbiol 14 113 (Sept - Dec ) 1942 283 Harper, RM Algae on animals a biblzographical note, Ecology 31 303, 1950 284 de Almeida, F, and da Silva Lacaz, C Considerações sobre três casos de micoses humanas, de cujas lesões foram isoladas ao lado dos cogumelos responsavers, algas provavelmente do gênero Chlorella, An Fac de med da Univ de São Paulo 22 295, 1946 285 Pratt, R, and others Chlorellin, an antibacterial substance from Chlorella, Science 99 351 (Apr 28) 1944 286 Harder, R, and Oppermann, A Ueber antibiotische Stoffe bei den Gruenalgen Stichococcus bacıllatis und Protosyphon botyroides, Arch f Mikrobiol 19 398, 1953 287 Kasparek, B Beitrage zur Diagnose des Ertrınkungstodes durch den Nachweis von Planktonorganismen in Lunge und Duodenum, Deutsche Ztschr fd ges gerichtl Med 27 132, 1936 288 Merkel, H Betr Planktonbefunde in Wasserleichen und deren praktische Auswertung, Deutsche Ztschr i dges gerichtl Med 31 211, 1939 289 Tomonaga, T Nochmals ueber den Nachweis von Plankton in den Ertraenkungsduessigkeiten, Jap J M SC, VII, Social Med & Hyg 3 197* (Dec ) 1940 290 Incze, G Die Bedeutung der Phytoplankton Resorptron beim Ertrınkungstod, Acta morphol (Budapest) 1 421, 1951 291 Halstead, B W Ichthyotoxism a neglected medical problem, Medical Arts and Sciences (College of Medical Evangelists) S 125, 1951 292 Meyer, K F, Sommer, H, and Schoenholz, P. Mussel poisoning. J Prev Med 2 369, 1928 293 Sherwood, H P Some aspects of shellfish sanitation in Britain, J Roy San Inst (London) 72 671, 1952 294 Sommer, H, and Meyer, K F Mussel porsoning, California & West Med 42 423, 1935 ·80 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE 293. Schittenheim, A Die Haffkrankheit, In Handbuch der inneren Medizin (Mohr & Stachelin) 3 ed 1934, v 1, p. 1018, J Springer, Berlin 296 Sommer, H, Whedon, WF, Kofod, C. A., and Stohler, R. Relation of paralytic shell-fish poison to certain plankton organisms of the genus Gonyaulax, Arch Path 24 537, 1937 297 Sommer, H, and Meyer, K F Patalytic shell-fish poisoning. Arch. Path 24 560, 1937 298 Sommer, H., and Meyer, K P Mussel poisoning, Hygeia 19 620, 1941 299 Tennant, AD, Naubert, J. and Corbeil, HE An outbreak of paralytic shellfish poisoning, Canad M A J 72 436, 1955 300 Halstead, Bw, and Lively, W M Poisonous fishes and achthyosarcotoxism, US Armed Forces M J S 157, 1954 301 Sapeika, N Mussel poisoning, South African M J 22 337, 1948 302 Covell, W P, and Whedon, W F Effects of the paralytic shell-fish poison on netve cells, Arch Path 24 411, 1937 303 Sapeika, N Actions of mussel Poison, Arch internat. de pharmacodyn et de thérap 93 135, 1953 304 Porge, J F. Les néphropathies d'origine alimentaire, Bull et mém Soc méd Paris 156 39, 1952 305 Dérot, M La néphrite après ingestion des moules, rdle de l'allergie, Presse méd 60 316 (March 5) 1952 306 Zerkle, C Our splintered learning and the status of scientists, Science 121 513 (Apr. 15) 1955 307 Kaempffect, W Plankton from the sea is already a food but it is unlikely to solve a world problem (Science in Review), New York Times, March 14, 1954 308 Ocean discounted as a food factory, New York Times, June 24, 1994 309 Sea's food riches scarcely tapped, New York Times, Nov 14, 1954 310 Kaerapffert, W Strange foods some people like, New York Times, March 14, 1954 311 Bassham, J A Atmospheric control by photosynthesis, Rescarch Reviews, Office of Naval Research, Wash Nov 1954, p 7. 312 Amyotrophic lateral sclerosis (progressive motor neutone disease) an epdemiological study, an exhibit at the Meeting of the American Medical Association, Atlantic City, June 6-10, 1955, by US Department of Health, Education, and Welfare, Public Health Service, National Insututes of Health, National Institute of Neurological Diseases and Blindaess, (1955, Govt Print Of. 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·INDEX Abrasions 14 Abscesses 13 Absorption spectrum 32 Absorbeats 14, 15, 17, 18 Aratabularia 33 "Action spectrum ' 32 Agar 6, 15, 16, 17 Agar Agar 8, 17 Alaoine 8, 34, 36 Alanta 9, 9, 10 Alcoholic drinks 16, 21 Algae 1, 2, 3, 4, 3, 6, 7, 8, 9, 10, 11, 12, 13, 14, 13. 16, 17, 20, 21, 27, 28, 29, 32, 33, 34, 35, 36, 37, 38, 39 40, 41, 42, 43, 44, 46. 47 48 52, 53, 54, 55, 56, 60. 62 63, 64, 65 Also set Macroscopic algac, Microscopic algae, Scaweed, Plack ton, Phytoplankton Algal bloom 54 Algal broth 44 Algal poisoning see Toxicity Algicide 47 Algin B, 17, 18 Alginate 18, 19, 20, 63 2010 binding property 19 deosal usage 19 effect on intestinal sodium 10d potassium 19 hemostatic 19 hydroabsorbent Agent, 10 constipation 19 ingestion 18 Injection 18 lubricating jelly 20 medication contag 19 mold for skin graft 19 moulage 19 mustard gas decontamioane 19 surgical absorbept 18 Amylase B Amyocrophic lateral sclerosis 65 Ansbatna 36, 40, 49, 50, 51, 52, 53, 54, 55 Anabaenopsis 30 36 Anaphylaxis 18 Anual aliment 11, 12, 23, 34, 63 Animal plackton, Jet Zooplankton Animal toxicity, see Toxicity Ankorodemus 41 Antibiotics 20, 55, 56, 63 Anticoagulants 20, 21, 63 Anthelmatics 16 Antimony 40 Aphanszomenon 50, 51, 33 Aphrodisiacs 13 Aquaculture 63 Arginine B. 33 Arsenic 40 Ascariasis 15 Ascices 49 Ascorbic acid 10, 39 40 Ash 9, 22, 25, 26, 30 Aspatagine 35 Aspartic acid 8, 34, 35, 36 Aromic submarine 64 Aconic wastes 47. 48 Autotrophic sigac } Calcium 8, 9, 11 18, 19, 40 Calotbrx 36 Cancer 13 14 Carbohydrate 2, 7, 8, 25, 26, 28 30, 33, 34, 36, 37, 46 Carbon 28, 29, 30, 35, 37, 53 Carbon dioxide 2 32, 33, 40, 41, 42 43, 47, 64 Carotene 3 29, 31 32, 38 40 Caroccords 29, 31, 32, 33 Carrag been 6, 16, 17 Carragheenin B, 17 20 Catalase 32 Cathartics 24. also see Larative Canon exchange residus 19 Cell wall componcars 2, 3 7. B, 36 Cellular plasma 17 Cellular respiration 26 Also see Photosyachesis Cellulose 2,3 8, 29 Centrocres 13 Ceralism 45, 60 Cbara 32, 48 Chelating agents 41 Chemistry 23 25, 28, 30 39 Chewing Caterial 21 22 Cbrlomonds 39 Cbironomer 31 Chibn 8, 25 26 Chlamydomon 3, 30, 33, 33, 39, 45, 47 Chlorella ), 27, 28, 30, 31, 32, 33, 34, 35, 36, 37, 38, 39, 40, 41, 42, 43, 44 47, 35 36 63 64 65 Chlorella 35, 36 Chlorination 46 Chlorine 9 25, 46 Chlorophyceae 14, 31, 32 Chlorophyll 3 29 31 32 Chlorophyta 2, 3, 27, 33 Chloroplast 29 дорого Cboxine 16 Choodrillesterol 37 Cbendras 3, 6, 7, 16, 20 Cberdasa 13, 20 Céreororcas 3, 44 Chrysop byta 3, 21 Cogueira 57, 58, 61 Bacillariophyceae 31 Bacteria 4 17 20 28 35 36 39, 41, 47, 56 Benthal growth Berrylum 40 Biochemistry 28, 32 Biomass 24 Bladderwrack 14 Bisnt mange? Blastocyst 35 Blood formation 31 Blood plasma substitute 21 Blue green gre 36, 37, 46, 50, 31 Boron 40 Bread 6 7, 12 Bromine 9. 12. 20 Bumns 14, 19 swabs 19 wound dressing 19 Alginic acid 6, 17, 19, 36 Allergy 16, 53, 54, 5, 61, 64 Hladam 16 Anaton 6 Aging acids 8, 29, 34, 35 Ammoba 17, 39, 47 Amporplaara 30 Celdrar 23, 24 Calcsteous age 16 81 ·82 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE Currhosis 49, 65 Cladophors 3 Clams 56, 58, 60 Classes of algae 2, 3 Classification 2, 3 Coagulant 39 Cobale 39, 40, Alse Cyano. cobalamin Co-carboxylase 39 Codium 9 Coclospbarem 50 Color classification of algie 2 Colorless algae 28. 32 Complexing agent, 1 Chelar ing agent Conferrar 14 Conjugais 45 Constparon 13. 19. 19 Copepods 4, 22, 23, 24, 25, 26 Copper 12, 13, 40. 47, 6) Corralına 3, 14 Cortisone synthesis 37 Counctics 17, 22 Cough 16 Crannost 6 Croceton 33 Crocin 33 Crustaccae 4. 25. 26 Crypromonai 41 Culrure 17 28 29. 32, 33, 34, 36, 37, 38, 41, 42, 43, 44, 47, 63, 64 Culture apparatus 41, 42, 43 Custard powder 7 Cyanocobalamın 10, 39, 40 Also se Cobalt Cyanophyta 3, 27 Cycadaces 36 Cylindrespermswn 45 Diacoms 3. 4, 23, 25, 27, 32, 37 Dictyopteris 16 Digenta 15 Digesubility 7, 28, 29 Daolagellace 4, 22, 23, 2ì. 56 Dinophyceae 56 Dietary shortage 1 Dietary supplement 1, 11, 26, 44, 63 64 Dilluk 6 Dullam 41 Divisions of algte 2, 3 Dorset uert 6 Drinking wacer concimination 46, 47 Dropsy 13 Drowning, diagnosis of 56 Drying oul 37 Duckweed") Dulcitolo Durmillea 14, 16 Dulse 6, 10, 16, 21, 22 Dysentery 15, 53, 54 Flondee 16 Floridoside 8 Fluorescence, 8 Phosphores cence "Flos quse" 43 Folic sad 10 Folinic acid 10 Food, algae and plankron as 1, 4, 5, 7, 13, 24, 25, 27, 29. 37, 38, 41, 44, 45, 47, 49, 63. 64 Freshwater algac 5, 27, 30, 42 Freshwater phytoplankton , 48 Freshwater zooplankton 23, 24 Fried seaweed 7 Fucoidia 8, 20 Furns 3, 9, 12, 15, 16, 17, 22 Fugu poison 38 Fungi 55 Funor 22 Furcellarte 9 Earache 14 EDTA 41 Ecls 58, 59, 60, 61 Egregia 30 Enteromorpha 15 Enzyme systems 33. 34, 37. 47 Ergoscerol 10, 37, 39 Englena 3. 35. 39 Euglenineae 31. 39 Euglenophyta 2, 3, 27 Euphasıds 23 Also see Krill Galactosides 8 Gameres 33 Gastroincestinal disorders 14, 15, 53, 64 Gelidaceae 6, 17 Gelidium 3. 6, 8, 15, 16, 20 Genera of algae 2. 3. 36 "Generacion time' 33 Gentiobionr 33 Germanyum 40 Gigartina 3. 17, 30 Glocoincbie 41, 50 Glaropeltis 22 Glucopyranoside 8 Glucose 7, 8 Glutamate 39 Glutamic acid 8, 33, 36 Glutamine 33 Gluranc acid 34 Glycine B, 35 Gorter 13, 14, 15 Gonyarlax 3, 56, 60, 62 Gout 14 Gracilarta 6, 13, 13 Gule most 7 Growth of algae and plaokron 4, 33, 37, 38, 19 Gymredinim 3. 41, 45 Gymnotborex poisoning 61 Dasgeladas 33 Deamination 36 Decomposition of algae 1 Warer 46 Definitions 2,4 22 Dehydrated algae 7 Debydrogeoise 18 Dulestria 20 Demulcent 15, 16 Dermacolagic disorders, Skin disorders Desmareste 11 Desmid 45 Desoxyribosecucleic acid 39 Duartbea 14, 15, 16, 33, 34 Diatomales 46 Families of algae 2, 3 Fars 7, 9, 25, 26, 33, 37, 38, 63 Fatty acids 9, 37, 38, 56 Feeding experiments 10, 11, 12, 18, 23, 39, 44, 49, 64. 65 Femaleness 33 Atso si Scivality Fertilizer 11, 36, 42, 63 Fash 4, 15, 23, 26, 39, 45, 46. 33. 36, 57, 58, 59, 60, 61, 67, 64 Fish poisoning 53, 56, 57. 58. 59, 60, 61, 62, 65 Flavoproccio 19 "Fleur d eau 45 Floride un scarcb & Heftruckbud 61. 62 Hansea 1 disease 33, 44 Harvesting 41, 43 ·INDEX 83 Knull 4, 22, 23 Kwashiorkor 1 Heartburn 14 Hekssroplankron" 4 Helmiol' 15 Heme 31 Hemicellulose B Hemia 31 Hepatitis 14, 32, 65 Hepacotoxins 49, 52, 53 Heretagne 26 Hetose 32 Histidine 35 Homorystinea 44 Human toxicity, sur Toxicity Hydrocarbons 9, 29 Hydrocolloids 19 Hydrogea 30. 47 Hydrogen peroxide 32 Kydrophylıc colloids 17 Hydroponics 63 Hydroxylamine 46 Hypse 13, 16 Hypovitaminosis 1 Minerals 2, 17, 27, 28, 29, 63 Molluscs 4 Molybdenum 40 Mosquito control 48 Macin 35 Muscus 14 Mussels 36, 37, 38, 39, 60 Mucants 31, 32, 39 Myroxuc manifestations 61, 64 Mysıds 23 Myxophyctae 31, 35, 45 46 35 Labor 16 Laminaria 3 6, 7, 8, 9, 10. 12, 13, 15, 16, 17, 18, 20 Laminarın 8, 20, 21 Land plants 29, 30, 34, 40 Laver 6, 10, 13, 14 Laxative 13. 19 Also see Cacharic Lead 40 Leprosy, Re Hansen s disease Leucine 8 33 "Limactic growth 5 Linoleic acid 37 Lipids 2, 9, 28, 29, 30, 34, 36, 37, 38 Lipoic acid 40 Lipoid substantes 9 "Littoral growth 3 Lysine 8 35 Ice cream stabilizer 20 Ichthyosarcotoxin 61, 64 Also Jer Fash poisoning Illumination of cultures 42, 43 Inflammation treatment of 14 Ingestion of algae and plank ton 4, 5, 6. 7, 25, 44, 48. 49, 12 Injection of algae and plankton 18, 49, 50, 52 54, 55 * Inner complex sales 41 Intestinal disorders 14, 15, 17, 53 Intracellular storage macter 7 8, 33, 36, 37 Iodiac 9, 11, 12, 15, 16, 20, 21, 38, 40 Index 20 Irish moss 6, 12, 15 16 Iron 40 Isoleucine 39 Isotopes 35, 18 Macrocystis 3, 17, 30 Macroscopic algae 3, 5, 7, B. 9, 10, 12, 17, 36 Also see Seaweed Magnesium 8, 9, 29, 40 Maleness 33 Also Jee Seruality Manganese 40 Mannican 7 Mannito! 7, 8 Mannose 7 "Maduring of the sea 63 Maouronic acid B, 17 Maricuicure 63 Marine algae 2 > 15. 16. 17. 20, 30 54 Marine plankton 2, 3, 4, 23 24, 25 Marne sence 6 farine zooplankton 23, 24 Medicinal usage 13, 14, 13. 16, 17, 39, 44, 45 Menstrual disturbances 13 Metabolisen 2, 29 29, 32, 37 Methionine 34, 35 Afacrorys 3, 19, 49, 50 51, 32, 33, 34, 35 Micronutriepos, Trace ele. ments Microscopic algae 3, 4, 7, 11, 27, 28, 29, 34, 36, 37, 38, 39, 40, 41, 49, 33. Ale ne Phytoplankton Nanaoplankton 4 Newcala 50 Nephropathies 60, 64 Nerroryst 3, 21 Nenicic plackcon 27 Neurologic manifescations 12, 14, 48, 49, 56, 60, 61, 64. 65 Neurotoxins 49, 52, 53, 61, 62 Niaca 10 40 Nickel 40 Nitella 3, 12 Nitrogen 2, 8, 18, 25, 28, 30, 33, 34, 35, 36, 37, 40, 46, 47 Nitrogen Gitation 35, 36, 10 Nitzschra 3. 37, 38, 39, 41 Nodulars 48, 50 Nontor 36 Nucleic acid 9, 35 Nutricos 12, z3 27 33, 34. 40, 41, 42, 43, 63, 64 Nuttitool uses 2. 3. 10. 11, 29 39 44, 63, 64 Darweed 13 Obesiry creatment of 16 Odour 45, 46, 47 48 Ord ogonsum 3. 33 Oikomora 30 Oleic acid 37 Oligosaccharide 36 Oorto 44 Orders of algae 2, 3 Oscillaroraceae *. » Oscillalonia 35. 45, 44, 39 "Oxıd1000 ponds" 43, 46. 47 Ondore processes 28 Oxygen 29 92,36 46 47, 49, 52, 64 Oryuriasis 17 Oils 10 Jaundice 49 Jelly mos7 Kex for 6 Kareago 21 Kelp 2 6, 8, 9, 10, 14, 15, 16, 17, 22 Kombro Kon Tik 26 Krebs ticarboxylic acid cycle 39 ·84 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE Poulaces 13, 16 Prawns 22, 23 Proline 8 Prorocentre 3. 60 Protein 1, 2, 7, 8, 25, 26, 29, 30, 33. 34, 35, 43, 44, 46, 47, 63 Procoporphyrin precursors 31 Protosyphon 56 Prototbeca ), 26. 33, 39, 33 Protozoa 4, 29, 42 Puffer fish 57, 58, 31 Pulmonary disorders, creatment of 16 Pyrexia 15, 16 Pyridoxine 40 Pyrrophyta 3. 27 Pyrimidine 39 Pyruvic acid 34, 38, 39 Safranel 33 Salt 12, 14, 25, 27. 63 Sargassum 14 Sausage castags 7 Scabies 14 Scenedusmus 3, 28, 33, 36, 37 41, 42, 44, 47 Schrotbrix 36 Scombrord poroning 61, 62 Scrofula 14, 15, 16 Scurry 14 Sea, color of 27. 45 Sua lele 6 ''Sea lettuce" 6, 9, 14 "Seabachers' eruption" 54, 55 Seaweed 2. 3. 5, 6, 7, 8. 9. 10, 11, 12, 13, 14, 16. 17. 20, 21, 22, 36, 37, 39. 63- 65 Also se Macroscopic algae bread 7, IZ dye 22 Blue 22 Palmitic acid 9, 37, 38 Pantothenic acid 40 Para aminobenzoic acid 39 Passeroc burds 13 Prarl moss 7 Pectins 2, 3,8 Pencoses 7 Pentosan 36 Pepade 8, 9 Phacophyceae 7, 8 Phaeophyta 3, 10.33 Phenylalanine 8. 33 Phosphorescence 26, 27, 32, 56 Phosphorus 40, 46, 48, 52 Phoroscositivicy 48, 49, 52, 53 Photosynthesis 2, 28, 29, 31, 32. 33, 40, 46, 47. 63 "Phyceine 56 Phycobilin 5, 29, 31, 32 Phycocolloids 17, 20 Phycocyanın 3. 31, 53, 54, 55 Phycoerythrin 3. 31 Phycology 1 Phylloggor 15 Physiology of algae 28 Phytoplankton 3, 4, 23, 24, 27 28, 29, 32, 37, 38, 39. 41, 44, 45, 48. 50, 51, 54, 63. 64 Alrose Microscopac algae Phytoplankton poisoning. HI Toxiciсу Pigments 2.9 28, 29, 31, 33. 13.55 Polor plane studies 34, 35, 40, 41, 42, 43 Plankton 1 3. 4. 5 22, 24, 25, 26, 27, 39, 45, 46, 48, 56, 62, 63 Alro sa Phytoplankton, Zooplankton, M. croscopic algac foods 26. 44 45, 63, 64 poisoning te TONICITY Planc plankton, ** Phyro plankton Plectonema 36 Pleurococcus 45 Poisoning, tri Toxicity Polysaccharides 8. 20, 36 Poliomyelios 65 Polytomella 3. 33, 39 "Pondweed's Porpema 3, 6, 10, 13, 21 Porphyria 31 Pocassium 9, 12, 17, 19, 40 R-value 30, 34 Radiant cnergy 29, 33, 34 Radiation 33, 39, 47. 48 Radioactivity 21, 47, 49 Rashes 14 Also 11 Skin disorders Red Sea 27, 45 Red snow" 45 Red ude 43, 54, 55 Red water. 27, 45, 56 Reducing sugars 7, 36 Renal disorders 14 32, 60, 64 Also su Urinary disorders Reproduction 2, 32, 33. 35 Also see Scrualery Respiratory manifestations 48. 49, 56 61, 64 Reversible hydrocollord rech. DIC 19 Rheumatism 14 Rbitorionum 16 Rbodomela 20 Rhodophyceae 8, 31 Rhodophyta), 6 Rbody meria 3, 6, 8, 10 Riboflavin 10, 39 40 Ribonucleic acid 33 Rice cularacion 36 Rakers 9, 10, 29 Rock moss 7 Rorfer4 Roughage 19 Rubidium 40 "gums 17 ISOnglass 6 Sensuvery to algae 18 Also sre Allergy Serine 8 Seren Lonle Sustes 26 Sewage disposal 43, 45, 46, 47 Sewage lagoons 47 Servalery 33 Also Are Repro duction She 16sh poisoning 56, 60, 62, 64 Shipwreck survivors 23, 2), 26. 63 Shock 21 Shrimp 4, 25, 26 Silica 3 Silver 40 Skin disorders 14, 17, 19, 44, 49, 52, 53, 54, 5, 64 "Slimming teus 16 Sodium 9, 17, 19 Soil, alkalinity of 36 Soil conservation 11, 36 Solar engines 29 Sorbitol Space ships 64 Species of algae 2, 27 Sprains 16 Starch 8, 36 Stearic acid 37, 38 Sterol. 2, 9. 37 Ssebacecew), 30, 36 Sacrbara19 ·INDEX BS Urinary disorders 15, 16 Also Ter Reol disorders Stilapbors 16 Suputas 16 Stipes 16 Strontium 40 Succio um de 33 Sucrose 32. 36 Sugar wrick 13 Sulfates 9. 46 Sulfur 40. 41 Surgical powder 18 Swayback" 12, 65 Sweating, prevention of 16 Sykurd 4t Syphilis :3 Valne 8, 35 Vanadiom 40 "Vasserbloom" 45 Vermifuges 16 Vermilion Sea 27. 43 Vitamins 2. 7, 9, 10, 29, 37, 38, 39, 40, 44, 63 Tin 40 Tirasiun 40 Tocopherol 10 Telypotbrix 36 TOXICIT 11, 19, 45, 48, 49, 50, 31, 32, 33, 34, 56. 57, 58, 59, 60, 61, 62, 64, 65 clinical bodiagt 48, 49, 52, 54. 61, 64 differencal diagnosis 49, 52, 54, 64, 65 pathologic findings 49, 32, 60. 61 symptomatologs 54. 64 treatmcot 62 Toxins, chemistry of 52, 56, 60, 61 Trace elements 17 40, 41 Transaminase systems 34 Transamination 36 Trehalose 8 Trichomonal Vigiaus, treatment of 17 Triglycerides 37 Tryptophane 35 Tungsten 40 Wacer, coloration of 27, 49 panficauon of 33 supply 45, 53 Laste 45, 46, 48 'Warerbloom* 45. 65 "Wasserbluche 43 Whales, food of 4, 23 World population 1, 27, 29. 63 Wrucks 13. 22 Taste of algae sod plankton 45, 46, 47, 48 Temperature, effects on cul tures 41. 42 Tetraodor 57.61 Tetraodotoxin 61 Therapeutic value of algae 13, 14. 38, 44 Thiamin 10, 38, 39, 40 Tbiarole 39 Three dimensional furung" 63 Throac disorders 15, 16 Thtronine 8. 35 Thyroid gland 15. 16 Alsa Jer Goicec, Iodine Xylan 8 Xylose 8 "Yellow tide 45 Yttrium 40 Zane 40 "Ultraplankton" 4 Ulva 3, 6, 9, 10, 30 Unsaturated fatty aids 37. 39 Uradium 40 Zooplankton 3. 4. 22, 23, 24, 27, 63 ·84 THE ROLE OF ALGAE AND PLANKTON IN MEDICINE Palmitic sad 9. 37. 38 Pantothenic acid 40 Para-aminobenzoic acid 39 Passerine birds 13 Pearl mais 7 Pertins 2. 3.8 Penroses 7 Pentosaos 36 Pepride 8. 9 Phacophyctat 7. 8 Phaeophyta 3. 10, 33 Phenylalanine 8, 39 Phosphorescence 26, 27, 32, 56 Phosphorus 40, 46, 48, 52 Phorosensitivity 48, 49, 52, 53 Photosynthesis 2, 28, 29, 31, 32, 33, 40, 46, 47, 63 "Phyceine 56 Phycobilin 5, 29, 31, 32 Phycocolloids 17, 20 Phycocyanın 3. 31, 53, 54, 55 Phycocrythrin 3. 31 Phycology 1 Pbyllogrson 15 Physiology of algae 28 Phytoplankton 3. 4. 23. 24, 27, 28, 29, 32, 37, 38, 39. 41, 44, 45, 48, 50 51, 54, 6), 64 Also Je Micto scopic algae Phytoplankton poisoning. He Toxicity Pigments 2, 9, 28, 29, 31, 33, 33, 35 Piloc plant studies 34 35. 40. 41, 42, 43 Plankton 1. 34, 3, 22, 24, 25, 26, 27, 19, 45, 46, 48, 56, 62, 63 1110 st Phyto. plankton Zooplankton, MAcroscopic algse foods 26, 44, 45, 63, 64 porsoang, see Toxicity Plant plankton, $* Phyco plankton Plectonema 36 Plavrococcus 45 Poisoning. #+ TOXICITY Polysaccharides 8, 20, 36 Poliomyelits 65 Polytomella 3, 3. 39 "Pondweed 5 Porphyra 3, 6, 10, 15, 21 Porphytua 31 Potassium 9, 12, 17, 19, 40 Poultices 13, 16 Safrans 33 Prawns 22. 23 Sale 12, 14, 25, 27, 63 Prolne B Sargass** 14 Protorerm 3. 60 Sausage casingt 7 Protein 1, 2, 7, 4, 25, 26, Scabies 14 29, 30, 33, 34, 35, 43, 44, Scenedesmas 3, 28. 33. 36, 37, 46, 47, 63 41, 42, 44, 47 Protoporphyria precursors 31 Schizobry 36 Protosypbon 56 Scombroid Pollowing 61. 62 Protorbere 3, 28, 32, 39.35 Scrofula 14, 15, 16 Protozoa 4, 29 42 Scurvy 14 Puffer fish 57, 58, 31 Sea, color of 27. 45 Pulmonary disorders, treatment Sea Iale 6 of 16 "Sca lettuce 6, 9, 14 Pyrexia 15, 16 *Seabachers' eruption" 54.33 Pyridoxine 40 Seaweed 2, 3. 3. 6. 7. 8. 9. Pyrrophyta 3, 27 10, 11, 12, 13, 14, 16, 17, Pyrimidine 39 20, 21, 22, 36, 37, 39. 63. Pyruvic acid 34, 38, 39 65. Also se Macroscopic slgae R value 30, 34 bread 7. 12 Radiant energy 29, 33, 34 dye 22 Radiation 32, 39, 47, 48 glue 22 Radioactivity 21, 47. 48 "gums 17 Rashes 14 Also see Skin sanglass 6 disorders Sensitivity to sigue 18 Alto Red Sea 27. 45 14 Allergy Serine 8 Red snow 45 Seven Lamle Suster 26 Red tide 45, 54, 55 Red Warer" 27, 45, 56 Sewige disposal 43, 45, 46, 47 Reducing sugars 7. 36 Sewige lagoons 47 Reaal disorders 14, 32, 60, 64. Sexualory 33 Also s Repro duction Also 1 Unnary disorders Shellfish poisoning 56, 60, 62, Reproduction 2, 32, 33, 35 64 Also we Sexuality Shipwreck Survivors 23. 25. Respiratory manifestations 48. 26. 63 49, 56, 61, 64 Shock 21 Reversible hydrocolloid tech Shtimp 4, 25, 26 dic 19 Silica 3 Rheumatism 14 Silver 40 Rbrzoclona 16 Skin disorders 14, 17, 19, 44, Rbodomela 20 49, 32, 33, 34, 5, 64 Rhodophyceae 8, 31 "Slimming reas" 16 Rhodophyta 3, Sodium 9, 11, 19 Rbodymenia 3, 6, 8, 10 Soul kaliatry of 36 Riboflavia 10. 39. 40 Soil conservation 11, 36 Ribonucleic acid 35 **Solar engines' 29 Ruse cultivaton 36 Sorbitol 8 Rickets 9, 10, 29 Space ships 64 Reck Moss 7 Species of algue 2, 27 Rotiers 4 Spraias 14 Roughage 15 Search 8, 36 Rubidium 40 Stearic sad 37. 38 Sterol, 2. 9. 37 Sacharina 9 5th borettes %. 30, 56 ·83 INDEX Ungary disorders is, 16 Also se Repul disorders Strlapbore 16 Stupet 16 Sapes 16 Stronuam 40 Succinamide 35 Sucrose 32. 36 Sugar wrack 15 Sulfates 9, 46 Sulfur 40, 41 Surgical powder 18 "Suyback 12, 65 Sweating, prevention of 16 Sak 41 Syphilis 13 Valupe B, 39 Vanadium 40 Vasserbloom 45 Vermitages 16 Vertalion Sea 27, 45 Vitamins 2, 7, 9, 10, 29, 37, 38, 39, 40, 44, 63 Ta 40 Titanium 40 Tocopherol 10 Tolyporbres 36 Toxicity 11, 19, 45, 48, 19. 50, 51, 52, 53, 54, 56, 57, 39. 59, 60, 61, 62, 64, 65 Clinical findings 48, 49, 52, 54, 61, 64 differential diagnosis 48, 52, 34, 64. 65 pathologie findings 49, 52, 60, 61 symptomatology 14, 64 creatmcot 62 Toxins chemistry of 52, 56, 60, 61 Trace clements 17. 40, 41 Transaminase systems 34 Transamination 36 Trehalose B Trichomonal vigintus, frear ment of 17 Triglycendes 37 Tryptophane 33 Tungsten 40 Water, coloracion of 21.45 parification of 33 supply 45, 54 taste 45, 46, 48 'Waterbloom 45, 65 Wasserbluche 1) Whales food of 4, 23 World populacoa 1, 27, 29, 63 Wracks 15, 22 Taste of algae and plankcon 45, 46, 47, 48 Temperature. effects on cultures 41, 42 Tetraodor 57, 61 Tetraodontoxta 61 Therapeuuc value of algae 13. 14, 38, 44 Thiamin 10, 38, 39, 40 Thiazole 39 "Three dimensional farmiog" 63 Thross disorders 13. 16 Threonine 6,35 Thyroid gland 15. 16. Also Kr Goster, lodise Xylan 8 Xylose 8 Yellow Ude 45 Ytrium 40 Zinc 40 "Ultraplankron't Ulve 3. 6, 9, 10, 30 Uosaturated facry acids 37, 38 Uranium 40 Zooplankton 3, 4, 22, 23, 24, 27 63

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