Handbook of Microalgal Mass Culture (1986) [1 ed.] 9781138505933, 9780203712405, 9781351362696, 9781351362689, 9781351362702, 9781138559646

Table of contents :

1. The Production of Biomass: A Challenge to Our Society 2. A Historical Outline of Applied Algology 3. Photosynthesis and Ultrastructure in Microalgae 4. Cell response to Environmental Factors 5. Productivity of Algae Under Natural Conditions 6. Laboratory Techniques for the Cultivation of Microalgae 7. Algal Nutrition 8. Microalgae of Economic Potential 9. Technological Aspects of Mass Cultivation – A General Outline 10. Elements of Pond Design and Construction 11. Outdoor Mass Cultures of Microalgae 12. Algae in Wastewater Oxidation Ponds 13. Nutritional Properties of Microalgae: Potentials and Constraints 14. Products from Microalgae 15. Blue-Green Algae as Biofertilizer 16. Economic Aspects of the Management of Algal Production 17. Future Prospects Index

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CRC Handbook of Microalgal Mass Culture Editor

Amos Richmond, Ph.D. Cahn Professor in Economic Botany in Arid Zones The Microalgal Biotechnology Lab The Jacob Blaustein Institute for Desert Research Ben-Gurion University of the Negev Sede Boqer, Israel

First published 1986 by CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 Reissued 2018 by CRC Press © 1986 by Taylor & Francis CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the validity of all materials or the consequences of their use. The authors and publishers have attempted to trace the copyright holders of all material reproduced in this publication and apologize to copyright holders if permission to publish in this form has not been obtained. If any copyright material has not been acknowledged please write and let us know so we may rectify in any future reprint. Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying, microfilming, and recording, or in any information storage or retrieval system, without written permission from the publishers. For permission to photocopy or use material electronically from this work, please access www.copyright.com (http://www.copyright.com/) or contact the Copyright Clearance Center, Inc. (CCC), 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400. CCC is a not-for-profit organiza-tion that provides licenses and registration for a variety of users. For organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. A Library of Congress record exists under LC control number: 85014679 Publisher’s Note The publisher has gone to great lengths to ensure the quality of this reprint but points out that some imperfections in the original copies may be apparent. Disclaimer The publisher has made every effort to trace copyright holders and welcomes correspondence from those they have been unable to contact. ISBN 13: 978-1-138-50593-3 (hbk) ISBN 13: 978-0-203-71240-5 (ebk) Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com

PREFACE Two groups of algae may be readily distinguished from the standpoint of the technology involved in their cultivation. harvesting, and processing. One group is the microalgae, the morphological features of which may be resolved only with the aid of a microscope. The other group consists of macroalgae, macroscopic seaweeds. In general, seaweeds are attached to rocky substrates on the sea bottom and may reach several meters in length, e.g., the Laminaria species. In contrast, minute planktonic organisms, measuring only a few microns in diameter, like Synechococcus or Chlorella or the filamentous Spirulina are typical mi­ croalgae. These are the primary basis of aquatic food chains and account for ca. 40% of the total photosynthesis on earth. In nature, planktonic microalgae are so dispersed that they may be seen by the naked eye only if blooms, very dense populations of cells, are formed. The occurence of blooms, however, is unpredictable and their harvesting thus cannot support industrial production, which can be achieved only in artificial systems. This handbook is devoted to the mass production of microalgae, and in my part, is based on some 10 years of experience in growing and studying microalgal cultures maintained at high population densities under laboratory conditions and in outdoor ponds. I was attracted to this field because of the exciting promise in growing unicellular algae on local resources as an alternative source of food and feeds for hungry or malnourished populations. At that time, the report of the United Nations’ Advisory Committee on International Action to Avert an Impending Protein Crisis had just been published (see Chapter 2). The message of that report was clear — the world’s population was increasing faster than the expansion of conventional agricultural production and a protein crisis affecting hundreds of millions of people the world over was looming. According to another report much referred to at that time, by the year 2000, the world supply of protein would fall short by some 39 X 106 tons. The United Nations’ committee recommended that conventional agricultural crops be supplemented with high-protein foods of unconventional origin, such as microorganisms. On this context, microalgae were natural candidates for several reasons: 1. 2. 3.

4.

Essentially the entire plant body has nutritional value since only a minor portion of the cell is indigestible. These single-celled organisms contain a substantial amount of protein, up to 65 or even 70%, i.e., some 2 to 4 times that of seeds and leaves of vascular plants. Microalgae have a very high yield potential, since the conditions for the culture may be readily controlled and the population density adjusted to that at which the highest efficiency of solar energy conversion per unit area is obtained. Perhaps the most attractive aspect of algal mass culture as a source of food is that brackish and sea water can be used for their production.

This may be the key for establishing high bioproductivity in regions which lack sweet water and would otherwise remain poorly productive. Today however, algaculture is still far from providing a source of inexpensive food on chemicals. Mass production of microalgae outdoors is a formidable task. Much more remains to be learned about the biology involved in this biotechnology and many technological details must be improved. Production procedures have to be simplified and the average annual yields have to increase severalfold before algaculture can become a significant agricultural endeavor. Nevertheless, the promise of cultured algae, particularly as a salt-tolerant crop in warm arid lands, is real. This is amplified by the growing realization that the deprivation and hunger of many hundreds of millions of people in developing areas of the world cannot be correctly and permanently relieved by importing food and materials from the developed

countries. This is the consensus of all who are involved in analyzing the problems of hunger and poverty in these lands. The only meaningful solution rests in local development, which would depend on imported knowhow and capital at first, but which would aim at promoting economic independence. In this frame of reference, efficient utilization of saline-water for agricultural production is of particular importance. Indeed, the resources of sweet-water are dwindling the world over and shall often have valuable alternative uses. The aim of devel­ oping local resources therefore would include in many cases the production of salt-tolerant crops, such as selected species of algae that could be grown for various economic purposes. Thus, even though the concept of mass production of microalgae to feed a hungry world has not yet been realized, it is still valid as a long-range goal. The concept should be expanded to include microalgae for biosynthesis of chemicals and special products, the emphasis being placed on production based on local resources and skills. Much more research will have to be carried out and detailed experience will have to be gained before cultured microalgae come of age as an industrial crop. I hope this handbook will be useful towards achievement of this goal. It is a pleasant duty to thank Dr. Marj Tiefert for improving the English of Chapters 3, 6, 7, 9. 11, 14, and 17. Also, I am very much indebted to Ms. liana Brina for her most competent and devoted assistance in taking care of the many technical and administrative chores involved in preparing this book. Amos Richmond Sede Boqer, July 1984

THE EDITOR Amos Richmond is Professor of Biology at the Jacob Blaustein Institute for Desert Research of Ben Gurion University of the Negev, the Sede-Boqer Campus. Dr. Richmond was born in 1931 in Tel-Aviv, Israel and received his B.Sc. in 1954 at the California State Polytechnic in St. Louis Obispo, California. He obtained his M.Sc. from the University of California at Los Angeles in 1956 and the Ph.D. in 1963 from Michigan State University at East Lansing, Michigan. He was thereafter a research-fellow at Purdue University in 1964 and at the University of California at Los Angeles in 1965-1966. He served as lecturer and senior lecturer in Botany at the Institute for Higher Education in the Negev from 1966 until 1971, and as chairman of the department of Biology of the Ben Gurion University of the Negev in 1970 to 1971. From 1971 to 1974 he was associate professor of Biology and Dean of the School of Natural Sciences at that university. He became Professor of Biology in 1975 and served as the first director of the Jacob Blaustein Institute for Desert Research at Sede-Boker from its inception in 1974 to 1983. In 1974 Dr. Richmond established at the Desert Research Institute the Micro-Algal Biotechnology Laboratory, which he had been directing since. Dr. Richmond received the Bergmann prize for distinction in applied research in 1984 and is the incumbent of the Miles and Lillian Cahn Chair in Economic Botany at B.G. University. He is the recipient of many research grants, among which from the German government, from the Israel Council for Research and Development, the Bi-national Ag­ ricultural Research Fund and the Solar Energy Research Institute in the U.S. He is the author of more than 70 publications and his current research interests are forcused on the growth physiology of salt-tolerant microalgae with particular reference to the physiology of growth in mass cultures aimed to yield products of economic potential.

CONTRIBUTORS Sheldon Aaronson, Ph.D. Professor Department of Biology Queens College Flushing, New York Aharon Abeliovich, Ph.D. Senior Lecturer The Laboratory of Environmental Microbiology The Jacob Blaustein Institute for Desert Research Ben-Gurion University of the Negev Sede Boqer. Israel Yael J. Avissar, Ph.D. Lecturer Department of Biology Ben-Gurion University Beer-Sheva, Israel E. W. Becker, Ph.D. Senior Scientist Institute of Chemical Plant Physiology University of Tubingen Tubingen, Federal Republic of Germany

David O. Hall, Ph.D. Professor Department of Plant Sciences King’s College London London, United Kingdom Eithan Hochman, Ph.D. Professor Department of Economics Ben-Gurion University of the Negev Beer-Sheva, Israel Drora Kaplan, Ph.D. Lecturer The Laboratory of Environmental Microbiology The Jacob Blaustein Institute for Desert Research Ben-Gurion University of the Negev Sede Boqer, Israel S. Herman Lips, Ph.D. Professor Department of Biology Beer-Sheva and the Salinity and Plant Physiology Lab at Sede-Boqer Ben-Gurion University of the Negev Israel

Zvi Cohen, Ph.D. Lecturer Microalgal Biotechnology Laboratory The Jacob Blaustein Institute for Desert Research Ben-Gurion University Sede Boqer, Israel

Amos Richmond, Ph.D. Professor Microalgal Biotechnology Laboratory The Jacob Blaustein Institute for Desert Research Ben-Gurion University of the Negev Sede Boqer, Israel

Joseph C. Dodd, Ph.D. Consulting Engineer Microalgae Technology Brawley, California

Carl J. Soeder, Ph.D. Professor Institute for Biotechnology Kernforschungsanlage GmbH Julich, Federal Republic of Germany

Zvi Dubinsky, Ph.D. Senior Lecturer Department of Life Sciences Bar-Ilan University Ramat-Gan, Israel

Y. Tsur, Ph.D. Professor Department of Economics Ben-Gurion University of the Negev Beer-Sheva, Israel

L. V. Venkataraman, Ph.D. Scientist and Area Coordinator Autotrophic Cell Culture Discipline Central Food Technology Research Institute Mysore, India

Avigad Vonshak, Ph.D. Senior Scientist Microalgal Biotechnology Laboratory The Jacob Blaustein Institute for Desert Research Ben-Gurion University of the Negev Sede Boqer, Israel

T A B LE OF C O N T E N T S The Production of Biomass: A Challenge to Our Society .................................................... 1 A Historical Outline of Applied Algology..............................................................................25 Photosynthesis and Ultrastructure in Microalgae..................................................................... 45 Cell Response to Environmental F acto rs................................................................................. 69 Productivity of Algae Under Natural Conditions................................................................ 107 Laboratory Techniques for the Cultivation of M icroalgae...................................................117 Algal Nutrition............................................................................................................................147 Microalgae of Economic Potential........................................................................................... 199 Technological Aspects of Mass Cultivation — A General Outline ................................... 245 Elements of Pond Design and Construction..........................................................................265 Outdoor Mass Cultures of Microalgae.................................................................................... 285 Algae in Wastewater Oxidation Ponds.................................................................................... 331 Nutritional Properties of Microalgae: Potentials and Constraints.................................................................................................................................. 339 Products from M icroalgae........................................................................................................ 421 Blue-Green Algae as Biofertilizer...........................................................................................455 Economic Aspects of the Management of Algal Production...............................................473 Future Prospects......................................................................................................................... 485 Index............................................................................................................................................ 489

1

THE PRODUCTION OF BIOMASS: A CHALLENGE TO OUR SOCIETY D. O. Hall

SUMMARY The overuse and undersupply of biomass is currently a serious problem and potentially a greater long-term danger than lack of food. Today 14% of the world’s primary energy is derived from biomass — equivalent to 20 million barrels of oil per day. Its predominant use is in the rural areas of developing countries where half the world’s population lives, e.g., Nepal derives nearly 100%; Kenya, 75%; India, 50%; China, 33%; Brazil, 25%; and Egypt and Morocco 20% of their total energy from biomass. A number of developed countries also derive a considerable amount of energy from biomass: Sweden, 9%; Canada, 7%; and the U.S. and Australia 3% each. A number of European-wide studies have shown that about 5 to 10% of Europe’s energy requirements could be met from biomass by the end of the 20th century. An especially valuable contribution could be in the form of liquid fuels which have become prone to fluctuating price and supply. The resources available, the effect of large agricultural surpluses (especially in North America and Europe), and the factors which will influence biomass energy schemes around the world are issues which, at present, are hotly debated. Worldwide government expenditure on biomass energy systems is over $2 billion a year while the costs of surplus food production is over $60 billion a year. However, biomass energy is not necessarily the panacea for any country’s energy problems, though currently the process of photosynthesis produces an amount of stored energy in the form of biomass which is almost 10 times the world’s annual use of energy. Additionally, the productivity of biomass-for-energy species can be dramatically increased as has already been shown. Such improvements have been accomplished with a number of agricultural species which are well known such as maize, wheat, and rice. The world produces 10 to 20% more food than is required to feed its 4.5 billion people an adequate diet. In North America and Europe the main problem with food is its easy overproduction and general over-consumption; however, there are an estimated 450 million undernourished people, mostly in Asia and Africa. Simplistically, if available food produc­ tion was increased by 1.5% (equivalent to about 25 t of grains), and if this food was distributed equitably to those who need it, there would be no undernourished people in the world. The same argument applies if only 10% of the developed countries’ grain production was diverted away from animals to humans. Health authorities have recommended lower meat and sugar consumption in the U .S . and U. K., and such changes are already occurring in some developed countries. These diet and other biotechnological changes will have long-term socioeconomic consequences. The question is how to achieve both food and biomass fuel production locally on a sustainable basis. Both are required, thus planning and provision of the appropriate infra­ structure and incentives must be provided. Increased support of research and development, training, and firm establishment of top priority to agriculture and forestry are essential in many countries of the world, if necessary, with significant help from abroad.

INTRODUCTION 1 -24 Biomass contributes a significant part of the world’s energy, being an important provider of energy to a great number of people. Decisions that are made over the next few years will significantly influence the level of biomass energy use in the future. How much biomass

2

Handbook of Microalgal Mass Culture

will contribute over the long term will depend very much on decisions that are made both at the local level and at the national level, in addition to international policy making, especially for energy and food. It is not within the scope of this chapter to cover any of the numerous contributions that algal biomass can make to food and energy chains. Nevertheless, it should always be borne in mind that when dealing with aquatic compared to terrestrial species there are usually totally different problems to be solved, whether it is for food, energy, chemicals, etc. Terrestrial plants are easier to grow and harvest but they often use valuable land. “ Food vs. fuel?” This is the wrong question. It should rather be, “ How can we equitably distribute the existing ample supply of food?” The problem of uneven distribution of the world’s adequate supply of food is well known and has been the subject of many reports over the last 50 years. This author is not competent to extend or improve on these studies. What I do, however, wish to point out is that both food and fuel are crucial limiting factors in development and must be locally available on a sustainable basis. Appropriate infra­ structure incentives and planning related to agriculture and forestry enterprises certainly help alleviate shortages in order to allow socioeconomic development to proceed. Again this is not novel, but unless real recognition of the immediate importance of agriculture and forestry in many countries of the world is forthcoming, these developing nations will continue to suffer national shortages of food and/or fuel which may have a debilitating effect on their growth (both personal and political). About 10% of the world’s people are undernourished, but world food production is more than 10% greater than is required overall and per capita food production has been increasing at a compound rate of 0.5% in developing countries and 0.8% worldwide since 1950. Although the world population has increased by 2/3 in the last 30 years (increased from 2.5 to 4.5 billion), the production of food has doubled. The amount of grains needed to solve undernourishment in the developing countries is only about 25 t annually — this represents only 1.7% of the total production worldwide and only 12% of the world’s trade in grains. Biomass energy produced annually is about 10 times greater than the total amount of energy used worldwide; still many people in developing countries suffer chronic shortages of fuelwood and its overuse is creating serious environmental consequences. Today, 2/3 of the world’s people depend on plants for nearly all their food and for the majority of their energy. The question should again be not “ food vs. fuel?” , but “ how can we increase the productivity of plant-based agriculture (and forestry) in order to provide the required levels of both food and fuel at the national or regional level?” Since only less than 10% of the world’s food and hardly any of the world’s biomass fuel enters transnational trade it is local production of plant products which is all-important. The average person in the rural areas of the developing world uses the total equivalent of about 1 t of wood per annum. This is used mainly for cooking and heating, but also for small-scale industry, agriculture, food processing, etc. The use of wood and charcoal in urban areas and for industry is often much greater than is realized. This author does not think many people appreciate the importance of biomass energy use because the statistics are not available to show this significance, and the consequences of biomass overuse are not readily evident. Until a few years ago world energy supply statistics listed biomass at 3 to 5%, if at all. It is now known that over half of all the trees cut down in the world today are used for cooking and heating. The problem of deforestation with its consequent flooding, desertification, and agricultural problems is not solely due to over-cutting of trees for cooking and heating. There are obviously other factors involved such as land clearing for agriculture, commercial and illegal cutting, absence of replanting, and so forth. Recently there have been several good papers published on studies in Southern and East Africa which show that in an average family of six or seven, one person’s sole job is to collect firewood and they will often have to walk great distances; this, of course, has other

3 deleterious consequences. In urban environments, households can spend up to 40% of their income on fuelwood and charcoal. Another aspect which has been highlighted in Tanzania is the curing of tobacco; for each hectare of tobacco the wood from 1 ha of savannah woodland needs to be incinerated. There are many examples showing that it is not only domestic, but also agricultural, urban, and small-scale industrial fuelwood overuse which is having serious long-term consequences. Several attempts are being made by a number of international and national groups to try and reverse the deforestation problem by vigorously promoting reforestation, village fuelwood lots, or community forestry and agro-forestry. The World Bank in 1980 concluded that if one was to reverse the deforestation problem one would need to spend $6.75 billion over the next 5 years in order to start reforesting 50 million ha. There is little hope that this will happen, but it was what realistically was thought to be needed. There are a number of reasons why this will not be possible, but only one shall be mentioned here. It is the very low status that foresters have in developing (and also developed) countries. Consequently, it is all very well to promulgate reforestation schemes to help solve the energy crisis in various parts of the world, but unless one has the people on the ground with the experience and knowledge, implementation of these schemes is impossible. The majority of people in the world exist by growing plants and processing their products. The main issue in developing countries is that of scarcity and the problem of trying to maintain, or possibly even to increase, the present level of use without harming agricultural or forestry and ecological systems. More efficient use of existing biomass and possible substitutes for biomass use, e.g., solar and wind-based technology and indigenous fossil fuels, should be considered and implemented as quickly as possible to reverse the trend of excessive biomass use, as is already occurring in many countries. In the developed world the expertise exists and is already being used to implement biomass energy programs from the standpoint of potential technology and economics. Biomass can provide a source of energy now and in the future; just how much it can contribute to the overall provision of energy will depend on existing local and national circumstances. It is imperative that each country establish its energy use patterns and the potential of biomass energy. This is not easy to accomplish quickly, but needs to be done as soon as possible. How much photosynthetic production of biomass actually occurs on the earth? It has been shown that the world’s total annual use of energy is only one tenth of the annual photosynthetic energy storage, i.e., photosynthesis already stores ten times as much energy as the world needs. The problem is getting it to the people who need it. Second, the energy content of stored biomass on the earth’s surface today, which is about 90% in trees, is equivalent to our proven fossil fuel reserves. In other words the energy content of trees is equivalent to the commercially extractable oil, coal, and gas. Third, during the Carboniferous Era quite large quantities of photosynthetic products were stored, but in fact they only represent 100 years of net photosynthesis. The overall photosynthetic efficiency during the Carboniferous Era was less than 0.002%, thus, our total possible fossil fuel resources only represent a few years of net photosynthesis. Fourth, there is the problem of C 0 2 cycling in the atmosphere. Many people are rightly concerned about this buildup of C 0 2 if we continue to bum fossil fuels. It is a problem of cycling between two or three pools of carbon. The amount of carbon stored in the biomass is approximately the same as the atmospheric C 0 2 and the same as the C 0 2 in the ocean surface layers; there are three equivalent pools. The concern is over how the C 0 2 is distributed between these pools and how fast it equilibrates into the deep ocean layers. However, we should appreciate that increasing C 0 2 concentrations in the atmosphere may be good for plants (since C 0 2 is a limiting factor in photosynthesis and plants have better water and fertilizer use efficiency at higher C 0 2 concentrations). Plants could also act as C 0 2 sinks if photochemical means for fixing C 0 2 were not available to alleviate the problem in the future.

4

Handbook of Microalgal Mass Culture Table 1 SOME ADVANTAGES AND PROBLEMS FORESEEN IN BIOMASS FOR ENERGY SCHEMES Advantages

1. 2. 3.

5. 6.

Stores energy Renewable Versatile conversion and products; some products with high energy content Dependent on technology already available with minimum capital input; available to all income levels Can be developed with present manpower and material resources Large biological and engineering development potential

7.

Creates employment and develops skills

4.

8. Reasonably priced in many instances 9. Ecologically inoffensive and safe 10. Does not increase atmospheric C 0 2

Problems

2. 3.

Land and water use competition Land areas required Supply uncertainty in initial phases

4.

Costs often uncertain

1

.

5. 6.

Fertilizer, soil, and water requirements Existing agricultural, forestry, and so­ cial practices 7. Bulky resource; transport and storage can be a problem 8. Subject to climatic variability 9. Low conversion efficiencies 10. Seasonal (sometimes)

The oil/energy problem of the 1970s had three clear effects on biomass energy and development. First, in a number of developed countries large research and development programs have been instituted which have sought to establish the potential, costs, and methods of implementation of energy from biomass. The prospects look far more promising than was thought even 3 years ago. Demonstrations, commercial trials, and industrial projects are being implemented. Estimated current expenditure is over $1 billion per annum in North America and Europe. Second, in at least two countries, Brazil (which currently spends over half of its foreign currency on oil imports) and China (with over 7 million biogas digesters), large-scale biomass energy schemes are being implemented. The current investment is over $1 billion per annum in Brazil. Third, in the developing countries as a whole there has been an accelerating use of biomass because oil products have become too expensive and/or unavailable. Biomass as a source of energy has its pros and cons. Like all other energy sources one must realize that it is not the universal panacea. Some advantages and disadvantages are listed in Table 1. This author wishes to emphasize the large biological and engineering development potential which is available for biomass. Because no advances in research have been made for many years, the efficiency of production and use of biomass as a source of energy has not progressed in the way agricultural yields for food have increased. Agricultural research has paid off very well (Table 2) and this may also be the case for biomass research and development as has been shown in recent fuel wood schemes. Thus there is an undoubted potential to increase biomass energy yields. Another advantage is the versatility of the biomass production and conversion technologies such that anyone involved in a particular project can select the routes most suited to the prevailing conditions and requirements of that area. The most obvious problems that immediately come to mind are land use in competition with food production. Existing agricultural, forestry, and social practices are also certainly a hindrance to promoting biomass as a source of energy, whether in a developing or an already developed country. In North America and Europe the problem in agriculture and nutrition is overproduction, excessive consumption of animal products via feeding of grains, and surpluses affecting world trade, especially in relation to commodity prices for many developing countries’ products. Obviously this is too simplistic, but the medium-term trends are important as they are likely to be aggravated by increased productivity, influence of new biotechnological processes, and changes in diet. Thus, for many countries the question should be how to

5 Table 2 STUDIES OF AGRICULTURAL RESEARCH PRODUCTIVITY — DIRECT COST-BENEFIT TYPE STUDIES25

Commodity Hybrid com Hybrid sorghum Poultry Sugar cane Wheat Maize Cotton Tomato harvester Maize Rice Rice Rice Soybeans Wheat Cotton Aggregate

Country U.S.

u.s. U.S. South Africa Mexico Mexico Brazil U.S. Peru Japan Japan Colombia Colombia Colombia Colombia U.S.

Time period

Annual internal rate of return (%)

1940—55 1940—57 1915—60 1945— 62 1943—63 1943— 63 1924— 67 1958—69 1954— 67 1915— 50 1930—61 1957— 72 1960— 71 1953— 73 1953— 72 1937—42 1947— 52 1957—62

35—40 20 21— 25 40 90 35 77 + 37—46 35—40 25— 27 73— 75 60— 82 79— 96 11— 12 None 50 51 49

achieve both food and biomass fuel production locally on a sustainable basis. Both are required, thus, planning and provision of the appropriate infrastructure and incentives must be provided. Increased support of research and development, training, and firm establishment of top priority to agriculture and forestry are essential in many countries of the world, if necessary, with significant help from abroad. Overall, biomass energy will not be a simple solution to the energy problems of developing countries. Biomass systems will not necessarily be inexpensive, nor will they be implemented easily, without a major commitment from governments and a considerable amount of political will at the national and international levels. Only a relatively small amount of food (25 t of cereal out of a world production of 1500 tonnes at present) is needed to solve the world’s hunger problem which condemns about 450 million people to suffer nearly perpetual undemutrition. How can this amount of food be produced and distributed so that those that need it actually receive it? This problem has vexed many and has been the subject of lengthy reports and will no doubt continue to do so in the future if, or until, the problem is solved. The gist of my message, which does not claim to be novel, is that it is possible to produce sufficient food and some fuel on the land using good agricultural and forestry practices, today and in the future. To what degree of self-sufficiency a country or region decides to achieve in food and fuel depends very much on local conditions, which are not easy to accurately ascertain from superficial surveys. If we really want to solve the undemourishment/poverty problem at the peasant and village level in developing countries (where most but by no means all of the world’s poor and malnourished people live) we must understand what the limiting factors are in food and biomass production at this level. Is it lack of water, fertilizers, pesticides, credit facilities, extension services, transport, storage, etc., or factors such as lack of human and animal power at crucial times of the year, tax incentives, high energy costs, too low product costs, and so on? My contention is that with a reasonable degree of management, combined with the nec-

6

Handbook of Microalgal Mass Culture Table 3 SOURCES OF BIOMASS FOR CONVERSION TO FUELS Wastes Manures Slurry Domestic rubbish Food wastes Sewage

Residues Wood residues Cane tops Straw Husks Citrus peel Bagasse Molasses Land crops (ligno-cellulose) Trees Eucalyptus Poplar Luceana Casuarina

Starch crops Maize Cassava Sugar crops Cane Beet

Aquatic plants Algae Chlorella Scenedesmus Navicula Multicellular Kelp Water weed Water hyacinth Water reeds/rushes

essary incentives and infrastructures many regions of the world could be self-sufficient in their basic food requirements, and in addition provide reasonable amounts of energy from natural or specifically grown biomass. BIOMASS26 33 Biomass is a jargon term used in the context of energy for a range of products which have been derived from photosynthesis; the products can be recognized as waste from urban areas and from forestry and agricultural processes, specifically grown crops such as trees, starch and sugar crops, hydrocarbon plants and oils, and aquatic plants such as water weeds and algae (Table 3). Thus, everything which is derived from photosynthesis is a potential source of energy since it is a solar energy conversion system. The problem with solar radiation is that it is diffuse and intermittent. Therefore if we are going to use it we must capture a diffuse source of energy and store it; plants accomplished this long ago. The process of photosynthesis embodies the two most important reactions in life. The first is the water-splitting reaction which evolves oxygen as a by-product. All life depends on this reaction. Second is the fixation of C 0 2 to organic compounds. All food and fuel is derived from C 0 2 fixation in the atmosphere. When looking at an energy process we need to have some understanding of what the efficiency of this process will be; one needs to look at the efficiency over the entire cycle of the system, and for the process of photosynthesis we mean incoming solar radiation converted to a stored end-product. Most people agree that the practical maximum efficiency of photosynthesis is 5 to 6%. This may not seem like very much, but one should be mindful that this represents stored energy (Table 4). Photosynthetic efficiency will determine biomass dry weight yields. For example, in the U.K. at 100 W m-2 incoming radiation a good potato crop growing at 1% efficiency (usually not higher than this in temperate regions), will yield about 20 t dry weight per hectare per annum. Obviously if we can grow and adapt plants to increase photosynthetic efficiency the dry weight yields will increase and alter the economics of the crop. One of the more interesting

7 Table 4 PHOTOSYNTHETIC EFFICIENCY AND ENERGY LOSSES34 Available light energy (%) At sea level 50% loss as a result of 400— 700 nm light being photosynthetically usable 20% loss due to reflection, inactive absorption, and transmission by leaves 77% loss representing quantum efficiency requirements for C 0 2 fixation in 680 nm light (assuming 10 quanta/C02),a and remembering that the energy content of 575 nm red light is the radiation peak of visible light 40% loss due to respiration

a

100 50 40 9.2

5.5 (Overall photosynthetic efficiency)

If the minimum quantum requirement is 8 quanta/C02, then this loss factor becomes 72% instead of 77%, giving the final photosynthetic efficiency of 6.7% instead of 5.5%.

areas of research is understanding what the limiting factors are in photosynthetic efficiency in plants both for agriculture and biomass energy. Currently the production of liquid fuels from biomass is of great interest, but yield limitations are becoming paramount in the economics of the overall processes. Table 5 highlights some important parameters. There is another aspect of photosynthesis that we should all appreciate, i.e., the health of our biosphere and our atmosphere is totally dependent on the process of photosynthesis. Every 300 years all the C 0 2 in the atmosphere is cycled through plants; this occurs every 2000 years to all the oxygen, and every 2 million years to all the water. Thus, 3 key ingredients in our atmosphere are dependent on cycling through the process of photosynthesis.

FOOD PRODUCTION, USE, AND PRODUCTIVITY 25 36 60 World agriculture is already producing enough to feed everyone on a Western standard of diet, however, inequalities and inefficiencies in distribution seem to make it impossible for the poorest to ensure their share. The current food stock is sufficient to feed the 1977 population of 4.2 billion at a level of 2570 kcal/day (about 20% above requirements) and supply 69 g/day protein. The disparity between developed ('/4 world population) and de­ veloping (3/4 world population) countries is seen in Table 6, both in the amount of calorie intake and the contribution from animal sources (5 times lower). Miller50 has estimated that of the total primary food production (62% grains) equivalent to 4514 kcal/person/day, only 2149 kcal is directly consumed by man and 2365 kcal is fed to animals, who in turn contribute 414 kcal to man to give an average worldwide total daily intake of 2563 kcal per capita. Over one third of the world’s cereal production (about 1500 t/year of which only 12% enters world trade) and over one half of the soybean production is fed directly to animals. If only V4 of the world soybean harvest was used for direct human consumption instead of animal feed it would provide 5 kg/year of a high protein food (750 kcal energy per day) for everyone in the world.2 The problem with using these facts to help solve the world’s food problem are well discussed by FAO in its recent Agriculture: Toward 2000A1 These facts, which are incon­ sistent with what is morally right, are often put forward to support the case for giving less cereals to animals and more to people. Unfortunately, neither the facts nor the solutions are simple. The great bulk of the feed use of cereals occurs in the developed countries, followed at a great distance by the more prosperous developing countries; the low-income developing countries use less than 2% of their total cereal supplies for animal feed (Table 7). In the

8

Table 5 ENERGY CROP YIELDS35 Conversion yield* (% by weight)

« ce

Energy crop

>
£

ce

Ô

CD CO

75 35 20 20

125 70 50 50

5.5 6.8 14 10

Ethanol

Fuel yield (GJ/ha) O» ce L*

u >

r and the optimal decision is not to harvest. As time progresses, population density increases and growth factors become limiting, suppressing growth until at some time instant tr dG/dB - r

(4)

From there on the optimal harvesting decision is obviously to keep population density at a level that satisfies the condition of Equation 4. Figure 2 describes the above situation. At population level Bs growth is maximal. Hence Bs is generally referred to as Maximum Sustainable Growth (MSG) level. At that level dB/dB = O and the B vs. time curve passes through an inflaction point. With interest rate equal to r, optimal biomass level (i.e., the level of B that equate vl to dG/dB) is B"2. In the same manner Br2 is optimal under r2. Note that for any positive interest rate the optimal biomass level, Br, is smaller than the MSG level Bs. Only at zero interest rate will Bs be economically optimal. Note also that if any growth factor changes over time independent of biomass level, then the optimal biomass level will change over time as well. To see this, assume B = G(B,t) = F(B)g(t) where g(t) reflects factors that suppress growth as a result of aging independent of biomass level B. Hence 0 < g(t) < 1 and g(t) decreases with time. Optimality condition Equation 4 becomes F'(B) = r/g(t)

(5)

With fixed interest rate r, r/g(t) increases over time, hence optimal level must be decreased in order to increase F'(B) and keep the equality above valid. Let us now relax the “ costless” assumption and see how the introduction of cost affects optimal behavior. Two general types of cost exist: fixed and operating costs. The first is assumed given (in the short run) and does not affect operating behavior. Operating costs in algal production involves direct production cost (such as supplementing materials, inducing

47 6

Handbook of Microalgal Mass Culture

FIGURE 2. The relationships between growth, interest rate, price, and cost in determining management decisions. B, B, Bs as in Figure 1; r = interest rate; Br = optimum population density corresponding to interest rate r; Bcr = Br, including operating cost; t = time; ts = time required to reach Bs; tr = time required to reach Br; C = unit operating cost.

water flow), harvesting cost, and drying and processing cost. Different algal products (e.g., human health food, animal food) entail different production, harvesting, drying, and proc­ essing methods and hence a different structure of operating cost and different product price as well. Let us postpone the discussion of these issues to the next section, where an empir­ ical example will be given. At this point we prefer to speak generally and let C(B) indicate unit operating cost. The two-period decision scenario depicted above is slightly changed. The gains from harvesting one (marginal) unit of biomass now is P — C(B). The gains from waiting and harvesting next period is comprised of two parts; the left in biomass unit have produced G'(B) during that period which amount to a value of [P — C(B)] (1 + G'(B)); the change in cost due to total growth at that period is approximately C'(B).G(B) (C'(B)defC/B). The discounted gains from waiting is then given by: 1/(1 + r) [P - C(B)] (1 -h G'(B)) - C'(B)G(B)

4 77

Optimal behavior requires that both gains be equal. This provides the following optimality conditions:

If C'(B) = 0, i.e., if unit operating costs are independent of biomass level, then Equation 6 reduces to Equation 4, the costless case. If C'(B) < 0 then the left hand side of Equation 6 increased and G'(B) must decrease in order to keep the equation valid, which implies (under G"(B) < 0) increasing optimal biomass level. In other words, if unit operating costs decrease with biomass level, then the incentive to leave a biomass unit another period is enhanced since it involves, in addition to growth, the effect on reduced operating costs. The same idea works in the opposite direction if C'(B) > 0. A rigorous treatment of the opti­ mization problem is given in the Appendix. The above problem of managing renewable inventory is similar to the classical capital investment problem in which some production process involves capital input, K. Assume no deterioration and let F(K) denote the net productivity of capital which is the rate of return from a capital stock of size K. The investor wishes to decide whether to invest and increase K by an additional (marginal) unit. An investment decision will provide a return of F'(K) for each dollar invested in buying the additional (marginal) unit. On the other hand the investor could put the money in the bank and receive r, the interest rate, as a return. Obviously, the decision is to invest as long as F'(K) > r. As the investment takes place, K increase and, generally, its efficiency decreases (each additional unit of K contributes less than the previous one to the production process) that is

In economic terms we assume decreasing marginal productivity of K. Eventually for some level of K

and at that point it does not pay to invest any more and the capital stock level is determined by this condition. Returning to our renewable biomass managing problem, the optimality condition (Equation 6) identifies three basic components: 1. 2.

3.

The biological growth process (Equation 1,which in economic terms is the production process. The cost structure of (a) harvesting and drying technologies, and (b) production tech­ nologies. Each of these costs involves fixed cost (infrastructure, ponds, harvesting facilities, etc.) and variable costs (nutrients, labor, energy, etc.). Market considerations as represented by P, the product price, and by prices of inputs to the production process.

The economic feasibility of producing a given biomass is a consequence of a combination of these three components. The first two are highly affected by managerial and human factors and we shall dwell on these issues. The third component is exogenous to the system assuming competitive markets. The next section is devoted to a discussion of these components in relation to algal biomass production accompanied by an empirical example.

478

Handbook of Microalgal Mass Culture AN EM PIRICA L APPLICATION

A model for the optimal management of algal biomass production was developed in the second section and the formal presentation is given in the Appendix. In this section we simulate an application of this model to the case of Spirulina production. We selected the data used in this application from reports concerning the production of Spirulina in the Negev desert. (In the simulation we are grateful to have permission to use a dynamic optimization algorithm developed at the World Bank from Alex Mirhause." As pointed out in the preceding section the structure of equations and parameters of the model can be classified into three main components: (1) growth (production) equations and their parameters; (2) cost functions and their parameters; and (3) inputs and outputs markets. Growth and production of the algal biomass are depicted by Equation 16 in the Appendix and in our applicaton had the specific form ABt = p(l - (Bt/M))Bt - ht

(9)

where p is the growth parameter, Bt is the level of biomass at time t, ht is the harvesting at time t, and ABt = Bt — Bt_, is the additional biomass growth minus harvesting at time t. Thus, the biomass at time t is determined by the level of biomass at time t - 1, plus the growth at time t and minus the amount harvested. The growth function consists of a linear coefficient and an asymptotic level M, and were estimated from a logistic growth function at the labs of the Desert Research Institute.6 The amount harvested is controlled by the grower and is a decision-variable to be determined by the optimization rules. These growth and production functions are heavily influenced by the human factor. The grower’s abilities can be classified into two groups; one includes the agro-technical abilities to produce the maximal possible amount of algae and the second relates to his managerial and economic abilities to manage the operations with minimum costs and maximum profit. The managerial abilities play an important role in the cost and revenue components which constitute the second and third groups of equations. In our application they are represented in the profit function (see also Equation 15 in the Appendix): N

VN = 2

[P - C(B,)] • h . - d / d + r))>

(10)

t= 1

where P is the price per kg of harvested Spirulina, C(Bt) is the cost per kg of harvested Spirulina, r is the interest, and VN is the present value of the sum of profit for a planning horizon of N periods. Since both the price and the cost per unit of Spirulina do not depend on the quantity sold, it is possible to separately calculate the operating profit per kilogram,* and then obtain the total operating profit per period by multiplying by the amount harvested. The sum over the N periods of the discounted value of operating profit per period yields the quantity VN. Now, the unit cost equation is represented in our case by C(Bt), which is equal to harvesting costs per m3 ^

C = Bt

plus production and drying costs per kilogram, indicated by d. Thus, we have decreasing marginal costs with respect to the level of algal biomass. As noted above the costs depend, among other factors, on the managerial ability of the producer. Operating profit measures the profit before deducting the fixed costs.

479

They also depend on the destination and final use of the algae. Thus, if the Spirulina is produced for human consumption as health products, the costs will be much higher than if the final use is for fish or animal food. The obvious reasons will be higher standards of quality as well as required government health regulations, e.g., the U.S. Food and Drug Administration. If the Spirulina is used to feed fish nearby, drying and overall production costs can be reduced drastically by leaving a high percentage of water in the produce. The price received from sales for human consumption is many times higher than for feeding fish. This brings us to another component, the demand for the final product. In our application this factor is represented by the price P in Equation 12, since we assume the producer to have only a small share of the market and is hence unable to influence the price. If the producer has a large share of the market, and therefore exercises an oligopolistic or a monopolistic power, he may be facing a negative sloped demand curve, i.e., D(p) with [D(p)]/p < 0, instead of a fixed price p for all levels of quantities sold. However, though it may make the calculations more complicated, the nature of solution will not change. Up to this stage, the procedure to calculate the operating profits over time was described. As in the static case of the conventional cost-benefit analysis fixed costs of investments must be taken into account. Since these are incurred at the initial state of operation in a fixed amount, they only determine the decision whether to start production or not. If the discounted net value of operation is greater (smaller), than the fixed costs production will take place (cease). The level of fixed costs is determined by the investment in the ponds, i.e., land scrapping, insulation of floor and walls, equipment needed for turbulation of water, as well as other required facilities for harvesting, drying, and producing the final product. Again, the final use of the product will influence the nature of investments, e.g., covering the ponds may be an important requirement for the quality of the final product in the case of human consumption. Two basic types of products are considered in the application: (1) expensive or high standard or quality product (e.g., for human consumption), and (2) cheap product (e.g., for fish feeding). These two basic models were evaluated for various changes in parameters. The empirical set of data was used to reproduce Equations 11 and 12 in the following equations and this basic model was evaluated for various changes in parameters. The cor­ responding basic equations are a human consumption: 0.27(1 - (Bt/1.6))Bt = ABt

(11)

and N

^ t =

(16 - C(Bt))ht(i/(l + 0.15))1 = VN per unit of product

(12)

i

where C(Bt) = 0.03/Bt + 5 An optimal solution is fully characterized by the trajectory of optimal biomass density over time. Since we assume growth, in a given production regime to be dependent on biomass level only (no aging effects), the optimal density level reaches a steady-state level (see discussion in the second section). The optimal harvesting trajectory is chosen in order to keep density level at its optimal trajectory. If harvesting is allowed to take negative values (i.e., it is possible to add biomass after the process starts), then optimal density level can be reached instantaneously. We assume this is the case with algal culture. Optimal solution therefore contains optimal steady-state density level (denoted by bss) and its corresponding

480

Handbook of Microalgal Mass Culture Table 1 OPTIMAL PROFIT AND STEADYSTATE LEVELS OF OPTIMAL DENSITY AND OPTIMAL HARVESTING: THE CASE OF A QUALITY PRODUCT Param eter changes Basic M = 0.2 M = 0.35 d = 10 d = 1 a

Optimal profit1* 51.92 37.99 67.86 28.09 71.02

optimal

hss optimal 0.108 0.080 0.140 0.108 0.108

0.796 0.795 0.798 0.798 0.796

($ )/m \

Table 2 THE “INEXPENSIVE PRODUCT” CASE Param eter changes Basic C = 0.03 C = 0.15 M = 5; C = 0.02; d = 0.05 M = 0.4; C = 0.02; d = 0.05

Optimal profit11

Bss optimal

hss density

12.44 13.45 11.16 8.46

0.842 0.816 0.879 0.810

0.319 0.320 0.317 0.200

6.59

0.815

0.160

a ($)/m\

optimal harvesting (denoted by hss), which is the growth at the density level bss. Table 1 presents optimal solutions and their corresponding profits for the quality product for different levels of the growth parameter p and the cost parameter d. From this table we can conclude: 1. 2.

An increase in the rate of growth, p, cet. par. increases the steady-state level of biomass Bss, and the amount of harvesting at steady state, hss. An increase in [P — C(B)], cet. par., leaves Bss and hss unchanged, but increases optimal profits.

Thus, optimal behavior and profit were found to be very sensitive to changes in the growth function, i.e., the production function and this on the human factors and extension services discussed above. In the previous example we considered a case of a quality algal product, Spirulina pills marketed as health food, with relatively high price (e.g., p = 16) and high unit costs of production (e.g., d = 5). For the sake of comparison we consider a case of a less refined product, that can be used to feed fish and this will receive a lower price, but will require lower costs of production having to meet lower quality requirements. In such a case to have a profitable production a higher rate of growth is required, and we assumed for the sake of demonstration that such a rate of growth is attainable. Thus, the example of a “ cheap product” is depicted by the basic Equations 13 and 14 and the various parametric changes by Table 2. 0.8(1 -

(B t/1 .6 ) ) B t = A B t

(13)

481

(14) where C(Bt) = 0.08/Bt + 0.05 and r is the daily equivalent of 10% annual interest rate. APPENDIX

A Mathematical Formulation of the Optimization Problem The (discounted) value of the algal biomass is PV =

Jo

e ' 5t(P - C(B))h(t)dt

(15)

where 8 is the (continuous) discount rate, P is price of a unit biomass, C(B) is unit production cost (include also harvesting, drying, processing and packing), h(t) is biomass harvested at time t. (We have assumed for a while an infinite planning horizon.) The growth process is given by dB/dt = G(B,t,U(t)) - h(t)

(16)

where U(t) represents all factors other than B that affect growth and which may be controlled by the grower. We assume the environment and technology are given so h(t) indicates production policy. The optimization problem can be stated as: Max PV U(t),h(t)

subject to the growth Equation 16

(17)

Assume first that U(t) is given so that Equation 17 is maximized over h(t) alone. Since the state Equation 16 is linear in h(t), the control variable, it is possible to directly apply the Euler condition (see “ A Historical Outline of Applied Algology” , Chapter 2) and to get the following optimality condition: (18) Equation 18 is with complete analogy to Equation 6. In many situations it is possible, due to biological or economical limitations, to grow only part of the year, say T days. Assuming identical conditions over the years the optimal solution for 1 >ear is also optimal for all other years. The problem can be stated as Max I e'8t[P - C(B)]h(t)dt + e T[P - CT]BT Jo

(19)

subject to Equation 16, where Cx is unit cost of impulse harvesting, i.e., emptying the pond. Note that T, the cycle length, can enter the maximization argument if the determination of optimal cycle length is required. In solving Equation 19 we define the Hamiltonian H = e8t[P - C(B)]h(t) + X(t)[G(B) - h(t)]

( 20)

482

Handbook of Microalgal Mass Culture

where \(t) is an auxialiary variable with an important econmic interpretation.8 Pontryagin’s Maximum Principle9 states that h(t) is determined so as to maximize H for each time period 0 < t < T. For time T, the transversality condition9 states \(T ) = d/dB{e8T[P - Ct]Bt

(21)

By rewriting Equation 20 as follows H = e" ‘[P - C(B)] - (t) h(t) + (t)G(B) it is easy to verify that the maximum principle implies for 0 < t < T h„,in if (t) < 0 Optimal harvesting policy (t) = < h*(t) if (t) = 0

( 22)

hmax if (0 > 0 where hmin (hmax) is the lower (upper) bound of h(h*(t) is the harvesting level that satisfies Equation 18 and 8(t) defe _St[P — C(B)] — Aft). The transversality conditions imply for time T \(T ) = e “5T(P - CT)

(23)

Along the singular solution \(t) = 0, i.e., \(t) = e “ l[P - C(B)], hence Equation 23 will be effective only if C(B(T)) 7 ^ Cx (i.e., only if the unit cost of impulse harvesting differs from the unit harvesting cost), which is not the case in algaculture. As noted earlier, T can be an argument of the maximization in order to determine optimal cycle length. This can be done by a numerical search method. Up to now we have assumed the vector of other control variables U(t) to be fixed. This vector includes variables such as quantities and nutrient content of supplemented fed ma­ terials, water flow rate, indoor or outdoor pond, etc. The subset of variables of U which are truly continuous and time dependent should be treated in the maximization process the same as h(t). This involves the generalization to multiple control theory,9 which in principle creates no problem, but for practical purposes is intractable. Those variables of U which are time independent can be treated in the maximization problem via standard numerical methods.10 LIST OF SYMBOLS t Bt = B(t) B = dB/dt ABt G(B) P M P r Bs K C 5

Time index Biomass population density at time t Rate of change of B over time Discrete change in B between (t-1) to t Growth rate Growth parameter Maximum population density that allows positive growth Price of algal product Interest rate Biomass population density at which growth is maximum Capital Unit cost Continuous discount rate

48 3

h(t) U T H \(t) hmin hmax h*(t) PV VN bss hss d

Biomass harvested at time t Control vector Cycle length The Hamiltonian Auxiliary (shadow price) variable Lower bound of h(t) Upper bound of h(t) Optimal harvesting policy Present value of the stream of future profits Present value of sum of profits for planning horizon of N periods Steady-state level for optimal biomass density Optimal harvesting level Unit drying and processing costs

REFERENCES 1. Binswanger, H. P. and Ruttan, V. W., Induced Innovation, John Hopkins University Press, Baltimore, 1978 (see chap. 1 to 5). 2. Schumpeter, J. A., The Analysis of Economic Change, Reprinted in Readings in Business Cycle Theory', The Blakiston Co., Philadelphia, 1944. 3. Clarck, C. W ., Mathematical Bioeconomics. The Optimal Management of Renewable Resources, Wiley Interscience, New York, 1976. 4. Rausser, G. C. and Hochman, E., Dynamic Agricultural Systems: Economic Prediction and Control, North Holland, New York, 1979. 5. Hochman, E. and Lee, I. M ., Optimal Decisions in the Broiler Producing Firm: A Problem o f Growing Inventory, Giannini Foundation Monogr. No. 29, University of California, Berkeley, 1972. 6. Richmond, A., Vonshak, A., and Arad, S., Environmental limitations in outdoor production of algal biomass, in Algal Biomass, Shelef, G. and Soeder, C. J., Ed., Elsevier/North Holland, Amsterdam, 1980, 65. 7. Vonshak, A., Abeliovich, A., Boussiba, S., Arad, S., and Richmond, A., Production of Spirulina biomass: effects of environmental factors and population density. Biomass, 2, 175, 1982. 8. Dorfman, R., An economic interpretation of optimal control theory, AER V LIX, N. 5, 817, 1969. 9. Pontryagin, L. S., Boltyanskii, V. G., Gamkrelioze, R. V., and Mishenko, E. F., The Mathematical Theory o f Optimal Processes, Interscience, New York, 1962. 10. Talpaz, H. and Tsur, Y., Optimizing aquaculture management of a single-species fish population, Agric. Syst., 9(2), 127, 1982. 11. Mirhause, A., Personal communication, 1981.

485

FUTURE PROSPECTS A. Richmond

The culture of algae for industrial purposes is a novel biotechnology, naturally prompting the question of whether it will ever become an important means of production of food, feeds, and chemicals. The author believes there is solid evidence that the utilization of algae for various economic purposes will gradually develop into an important industrial endeavor, particularly in warm, sunny regions where the water available is unsuitable for the production of conventional agricultural crops. A crucial question in analyzing the future prospects of algaculture concerns the market potential for algal products. No comprehensive answers are available yet, but certain points have become clear. The nutritional value of various algae as animal feeds and as food for humans has been substantiated by hundreds of research studies. In addition, scores of studies have described various therapeutic effects of microalgae. There is a growing list of chemicals that can be extracted from microalgae for commercial purposes. Most notably are glycerol and (3-carotene from Dunaliella, sulfonated carrageenan-like polysaccharides from Porphyridium sp. and phycocyanin from Spirulina which seem well on their way to commercial production. Other products, such as linoleic acid from Spirulina and arachidonic acid from Porphyridium represent potential products from microalgae. Most important in this respect is the fact that while tens of thousands of algal species have been taxonomically defined, only a small fraction have been surveyed for their possible economic potential. Yet, it seems possible to produce from these microorganisms nearly all the products that are currently obtained from conventional crops. Such crops do not usually produce continuously yearround, and most importantly, require a supply of Sweetwater, a resource which is becoming scarce in arid and semi-arid lands all over the world. Being unicellular and having a short life cycle, microalgae in general and cyanobacteria in particular represent a most suitable material for genetic manipulation. Transfer of genes for the synthesis of specific molecules should eventually allow large-scale production of many products that are presently obtained rather inefficiently from terrestial plants or from heterotrophic microorganisms cultured in expensive reactors. It seems clear that the pressing issue in commercial algaculture is not so much the market potential of algae products as it is the cost of their production. The current prices of commonly used animal feeds, i.e., various grains, and of high-protein plants such as soybean for human consumption, are approximately one-tenth the cost of production of algae in the small industrial plants in operation today. What, then, are the chances for microalgae in the competition with conventional crops for land, water, and capital? There are several indications that the cost of production of microalgae, which today confines them to the health food market, will gradually but con­ sistently decline, in parallel with continuous expansion of the market. In general, the history of agriculture and industry provides numerous examples for the natural course of development of new fields of endeavor. Repeatedly, be it with rubber trees or wheat, experience has shown that with time, productivity increases and the cost per unit production decreases by an order of magnitude or more. The scale of production has the most decisive effect on the cost of unit-product. A costbenefit analysis for the production of Scenedesmus in Peru1bears this out. With a cultivation area of 2500 m2, the total production cost of 1 kg of drum-dried Scenedesmus harvested by centrifugation was $7.50. With a fourfold increase in cultivation area with carbon feeding by self-produced C 0 2 and harvesting by flotation, the production cost per kilogram of dry matter was reduced about 50%. A further tenfold increase of the cultivation area to 100,000

486

Handbook of Microalgal Mass Culture

m2 decreased the calculated production cost to $1.17/kg. The capital investment was cal­ culated as $2,851,000, depreciation; maintenance, $309,000; and variable costs, $703,000. The annual output rate was considered to be approximately 1000 tons, i.e., 100 tons/ha/ year or approximately 27 g dry wt m _2/day. It is almost self-evident that due to the substantial capital investment needed per unit pond area, the cost of production is greatly dependent on the yield per area. In the above example, a rather high output rate was assumed and this was a major reason for the relatively low cost calculated for large-scale production. In reality, to date only about 30 to 50% of these outputs have actually been obtained over a long-term production period, making the actual cost per unit product being 5 to 10 times higher. Nevertheless, yields greater than 10 kg/ m _2/year are possible and have been obtained for periods lasting a few weeks, (see “ Outdoor Mass Cultures of Microalgae” ). Theoretically, the photosynthetic efficiency can be ap­ proximately 8% where the temperature does not limit growth, which implies a tremendous output rate, i.e., about 200 tons of dry matter/ha/year. However, even a consistent output one third the theoretical maximum will significantly reduce the current cost of production, which ranges between $5 to $10 (U.S.)/kg of drum- or spray-dried algal powder. The current low yields of 15 to 30 tons/ha/year that are usually obtained in outdoor mass cultures reflect the present state of ignorance in large-scale production of microalgae. It seems appropriate to compare the present know-how in algaculture with that thousands of years ago at the beginning of agriculture, when the cultivated plant species and the cultivation methods used only reflected man’s preliminary selection of species and rudimentary skills in cultivating them. Even a superficial view of the methods and machinery currently used for commercial production of microalgae would reveal that the industry is still in the initial phase of trial and error, adapting machines and materials from other industries. Every phase of this biotechnology may be significantly improved. One example concerns pond-lining, which represents a major investment cost. Lining made of PVC with a long durability is being developed today. If lining could be safely used for two or three decades, the capital investment for pond construction would be significantly curtailed. The cost of nutrients, which comprises some 30% of the running cost of production, could also be reduced. A major contribution to this end would be the provision of an inexpensive source of C 0 2. Industrial waste would be one such source, being suitable, however, only where industry is in close proximity to the production site. Another possibility with a wider application is to extract C 0 2 by burning a 10:1 mix of calcium carbonate and crude oil (using the heat produced for dehydrating the product). Recently, tremendous reservoirs of C 0 2 have been discovered in the southwestern U.S. Tapping such resources for algaculture should be considered. In addition, with algae such as Spirulina, which are grown on an alkaline medium with a pH of 10.5 a substantial amount of carbon required for an average output may be obtained from the C 0 2 in the air. The use of species selected for growth in highly alkaline pH would reduce the cost of carbon nutrition. The most expensive nutritional input except carbon is nitrogen. An attainable target with this nutrient would be the use of nitrogen-fixing microorganisms in conjuction or in symbiosis with the cultured algal species, thus reducing or even eliminating the need for a nitrogen fertilizer. It can be expected that improved methods for harvesting the algal mass will become available in the future. To aid in harvesting unicellular microalgae, simple flocculation techniques, based on physical methods such as electro-flocculation and flotation, will have to be developed. Such means should replace the cumbersome and costly chemical methods commonly used to affect flocculation. New, edible chemical flocculants, as well as novel techniques for screening and filtration could be expected to become significantly improved for the harvesting of microalgae.

487

Expenses on power would be cut if the conventional electric grid could be dispensed with, and photovoltaic power (in the not too distant future) could be used for stirring, pumping, filtration, and even drying of the algal mass. In principle, photovoltaic power can be used most effectively for algaculture, since power is needed mainly during the day and a good parallel exists between peak production of solar power and peak power demand by the algal system. Storage capacity of power, which at the present hinders the economics of photovoltaic power, must be minimal when photovoltaic power is used in algaculture. Natural energies could be also used for dehydration of the harvested algal mass. In arid, sun-rich areas (the natural sites for algaculture) air is relatively dry, being suitable for rapid drying processes. Pretreatment of the algal mass with solar steam will be advantageous in sun-rich areas once solar technologies for steam production, which are being developed today, are put into general use. As long as the commercial aspects of microalgal culture were not recognized, there were no opportunities to relate technically to problems associated with large-scale operation. There is every reason to assume that with the establishment and gradual expansion of a microalgal industry, this biotechnology will continuously improve, becoming increasingly more spe­ cialized and more efficient, with immediate effects on reduction of the cost of production. For example, a breakthrough in commercial algaculture resulted from the increased demand for some microalgal species as health foods, which prompted establishment of large-scale algaculture in several countries. The experience gained in the present production plants will have far-reaching effects on advancing this novel biotechnology. Also, the know-how ob­ tained in large scale commercial production of health foods will be instrumental in the development of other uses for microalgae, e.g., supplements for human food and various special chemicals. A significant cut in the cost of algal production could be expected when the market is further expanded, e.g., through demand for special feed supplements for animals, particularly fish and other aquatic organisms, but also for poultry, hogs, and ruminants. One of the most important advantages of algaculture which will have a decisive effect on its future prospects is the ability of many algal species to thrive in saline water, unsuitable for the production of most agricultural crops. When experience and understanding in this biotechnology becomes such that production is greatly simplified and its cost significantly reduced, algaculture in brackish water will have clear economic advantages in many lands. Somewhat paradoxically perhaps, it is in the hot, dry, usually poor lands of the world which often have untapped sea and brackish water resources, and where only meager conventional agriculture now exists, that algaculture may become a highly productive venture.

REFERENCE 1. Becker, W ., Personal communication.

489 IN D E X

A Abbreviations, list of, 445 Abortion, induction of, 424 Absolute temperature (T), 79 Absorbed light, wavelengths of, 52 Absorption scan, 71 Absorption spectra, 64 Accumulation of algal mass, 303 Acetamide, 165 Acetate, 162— 163, 203 Acetate-limited chemostat cultures, 210 Acetic acid, 31, 212 Acidic heteropolymer, 238 Acid polysaccharides, 27 Acid tolerance, see pH Acrylic acid, 443 Action spectrum, 64 Activation energy (E), 78— 80 Active contractions, 72 Active state colonies, 239 Adaptation, 94 to new light intensity, 70 to salinity, 90 Adaptive enzymes, 211 Adaptive reactions, 69 Adaptive response to salt stress, 95 A/dC, 65 Addresses of culture collections, 143 Adenosine, 165, 190 ADP, 50 Aerobic bacteria, 322 Aerobic growth conditions, 208, 301 Aerobic respiratory metabolism, 298 Africa firewood collection, 2 grain production, 10 Agar, 27, 235 Agar plates, 124 Aging symptoms, 230 Agmenellum quadruplicatum, 164, 183 Agriculture, 1—5, 11, 13, 14, 21 Agro-forestry, 3, 21 Agronomic practices and blue-green algae growth, 459—461 Air, 304 preincubation in, 302 temperature, 314 Airlift device, 31, 252 high rate oxidation pond, 335 Akinetes, 56— 57 ALA formation, 65 Alanine, 162, 164, 165, 169, 171, 227 Alanine dehydrogenase, 169, 171— 172 Albazod, 35—36, 285 Alcohol, 17, 347 Alcoholism, 230 Algaculture, 485—487

Algae-bacteria cultures, 33 Algae-bacteria interactions, 322 Algal-bacterial matter, 285 Algal biomass, see Biomass Algal blooms, see Blooms Algal cell concentration, relationship with incident light and extinction coefficient, 291 Algal cells absorbance of incident light, 286 calculation of number, 325 concentration of, 288 initial number, 325 maximum possible number, 325 Algal concentration, outdoor culture, 288 Algal cultures, 326 Algal dominance, 322 Algal filament, 324 Algal growth media, 126— 134 Algalization technology, 29, 462—464 Algal mass production, 153 Algal movement, 72—74 Algal nuclei, see also Nucleus, 61 Algal nutrition, see Nutrition Algal ponds, see also Ponds, 30, 35— 36 Algal populations, 323 outdoor mass culture ponds, 313— 314 supersaturation with oxygen, 299 Algal strains, selection of, 312 Algal succession, 457—458 Algal suspension, light measurement in, 137— 138 Algal technology, 29, 462—464 Algal ultrastructure, 62 Algal yields, see Yields Algatron, 33 Alginates, 27 Alginic acid, 27, 199 Alien algae, 377— 378 Alien organisms, 317 Alkaline lakes, Spirulina, 217 Alkaline saline lakes, Spirulina, 214 Alkaline water, 215 Alkalinity, 113 cyanophage tolerance to, 320 Spirulina, 217— 218, 220 Alkinate, 179 Allen and Amon medium, 134 Allen’s medium, 126— 127 Allergic effects, 385 Allophanate lyase, 172 Allophycocyanin, 58, 59, 66 , 350 Allophycocyanin B, 58 Alphanothece, 166 Aluminum, 181, 186, 4 0 0 -^ 0 1 Ambient temperature, 82 Amides, 164 Amino acid oxidase (AAO), 437 Amino acids, 164, 169, 172— 173, 180, 304, 342— 344

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203 fortification with, effect of, 369— 370 industrial production of C h lo r e lla , 212 patterns, microalgae, 33 synthesis, 55 Aminooxyacetate (AOA), 172 Ammonia (NH4+), 109, 126, 164, 167, 173, 189 assimilation, 168— 171 C h lo r e lla yields, 210 high rate oxidation pond, 332— 333 S p ir u lin a mass culture, 222 S p ir u lin a production, 224 Ammonium, 164, 165, 167, 168, 172 Ammonium-assimilating enzymes, 169 Ammonium hydroxide treatment, 327 Ammonium N, 168 Ammonium salts, S p ir u lin a , 220 Ammonium sulfate, S p ir u lin a production, 224 Amoeba, 29, 72, 323 C h lo r e lla ,

A m oeba

323, 324 324, 325 sp., 322, 327

d is c o id e s , r a d io s a ,

A m p h id in iu m c a r te r a e , A m p h ip h o ra a la ta ,

69

164

Amphitrophy, 147 sp., 430 (3-Amylase, 349 Amylolysis, 353 Amylopectin, 350 Amylose, 350

A m ph ora

A nabaena c a ta n u la ,

437

c y lin d r ic a

enzymes, 437 lipids, 430 macronutrients, 168, 170— 171, 173, 181 micronutrients, 185, 188 vitamins, 188 flo s a qu ae

light and, 74— 75 light intensity and dissolved oxygen, 305 maintaining nonalgal cultures, 317 pest control, 321, 326 products from, 437, 440 toxicological aspects, 372 o s c illa r io id e s , 437 sp. as biofertilizer, 456 macronutrients, 165— 167, 170— 171 outdoor mass cultures, 317 products from, 438, 439 strain 7120, 168, 170— 171 s p ir o id e s , A l l v a r ia b ilis

hydrogen ion concentration, 86 light and, 73, 89 nutrition, 160, 164 outdoor mass culture, 299 products from, 437 salinity and osmoregulation, 89

temperature and, 80, 82, 89 166, 456 A n a c y s tis , 189, 356

A n a b a e n o p s is , n id u la n s

laboratory techniques for cultivation, 120, 135 nutrition, 156, 157, 168— 169, 189 outdoor mass culture, 300, 302, 305, 323 products from, 437, 438 salinity and osmoregulation, 94 temperature and, 84— 85 Anaerobic growth, 163 Angstrom, 45 Animal feed, 392—405 aquaculture, 402—405 cropland used for, 12 insects, 401—402 nutritional value of algae for, 485 pigs, 398— 399 poultry, 393— 398 ruminants, 399—401 soybean harvest, 7 supplementary effect of algae, 368 Animal feeding tests, 354— 356 Animal nutrition, microalgae, 35 Animal studies, toxicological, 381— 386 A n k is tr o d e s m u s sp., 32, 173, 31 1, 357, 438 Annual output rate, 486 A n o p h e le s , contamination by, 378 Anosine, 165 Anoxic conditions, 109 Antenna molecules, 50 Antenna pigments, 49 Antennas, 58, 64 Antheraxanthin, 350, 432— 433 Antibiotics, 125, 443—444 Antihypertensive, 425 Antisecretory agent, 425 A p h a n iz o m e n o n , 75 f lo s a q u a e , 372 Aphanizophyll, 350 A p h a n o c a p s a 6308, 173 A p h a n o th e c a h a lo p h y tic a , 440 A p h a n o th e c e , 456 h a lo p h y tic a , 438 A p h e lid iu m , 322, 325, 326, 378 Applied algology historical outline, 25—41 microalgae, 25— 26, 28— 35 seaweeds, see also Seaweeds, 25— 28 Aquaculture, 35, 402—405 Aquatic biotope, 101 Aquatic fungi, 321 Aquatic habitats, 240 Arachidonic acid, 239, 424, 445, 485 Areal output rate, 289 Areal yield, S p ir u lin a , 224 Arginine, 57, 165, 171, 173, 227 Arrhenius equation, relationship of chemical reac­ tions to temperature, 79 Arrhenius plot, relationship between growth rate and temperature, 79

491 Arsenic, 181, 186, 228 402 Arthritis, 230

A rte m ia ,

A r th r o s p ir a

215 215 Artificial heat, 260— 261 Artificial seawater medium, P o r p h y r id iu m , 238 Artificial systems for algal production, 154 Artificial upwelling method, 34 As, see Arsenic Asexual reproduction, 230, 235 Ash, 227, 346 Asparagine, 164— 165 Aspartate, 162, 165, 187 Aspartic acid, 57, 164, 173, 227 A sp e r g illu s , 379 Assimilation number, 86 A s ta s ia lo n g a , 430 m a x im a ,

p la te n s is ,

A s te r io n e lla

142 443

fo r m o s a , n o ta ta ,

A s te r o m o n a s g r a c ilis ,

92, 440

Asthma, 424 443 Asymmetric molecular organization, 51 Atmosphere, oxygen replenishment, 47 Atmospheric pressure, 298 Atomizer technique, 123— 124 ATP, 45, 50— 51, 53— 54, 167, 170, 174, 306 determination, 436—437 generating system, 51, 52 inorganic polyphosphate synthesis, 180 light energy conversion into phosphate bonds of, 178 A u lo s ira , 166, 456 Autocatalysis, 55 Autoclaving, 353 Autocolony, 235 Autoflocculation, 258 Automatic synchronization device, 135 Autospore release, mode of, C h lo r e lla , 201— 202 Autospores, 200— 202, 208 Autosporogenesis, 200 Autotrophic algae C h o r e lla , 212 reactors for mass cultivation of, 245— 249 vitamin needs, 189 Autotrophic C h lo r e lla biomass, 209 Autotrophic growth, 181 Autotrophic metabolism, 162 Autotrophs, 147 Autotrophy, 147— 148, 189 Autumn, 294, 296— 297, 316 Auxins, 189 Auxotroph, 147 Auxotrophic algae, 189 Auxotrophy, 147— 148, 188, 189 Available irradiance per cell, 315 Available lysine, S p ir u lin a , 227 Average yield, C h lo r e lla culture, 205 A s tr io n e lla ja p p o n ic a ,

Axenic clones of blue-green algae, 141— 142 Axenic cultures, 125, 137 P o r p h y r id iu m cru en tu m , 238 S c e n e d e s m u s , 235 Axial fibers, 61 Azaserine (AS), 172 A z o lla , 455, 464— 465 A z o to b a c te r , 186

B Bacillariophyceae, 61, 65—66, 200 see Diatoms Bacteria, 29, 317 ammonium assimilation, 170 control of, 320, 322 high-rate algal ponds, 36 mixed population of microalgae and, 285 photosynthetic assimilation of C 0 2, 43 potassium, 182 Bacterial contamination, 379— 381 Bacterial count, 322 Bacterial load in algal cultures, 379, 380 Bacteriophages, 320 Balanced growth, 117 Barium, 181 Barometric pressure, 298 Bases, 421 Batch cultures, 118— 119, 155, 311 B-complex vitamins, 225 Beef liver, 349 Beer-Lamberth law, 106 Benomyl, 326 Benson-Calvin Cycle, 53 3,4-Benzypyrene, 228 Beryllium, 181, 186 Betaine, 165 BG-11 medium, 127 Bicarbonate concentration, 317— 319 Bicarbonate ion, 85 Bicarbonates, 217, 222, 317 B id d u lp h ia a u r ita , 430 Biflagellate alga, 240 Bilayer lipids, 85 Bilayer membranes, 62 Biliproteins, 58, 188, 220 Bioconversion processes, 441 Biofertilizer, 455—471 agronomic practices, 459— 461 algalization technology, 462—464 availability of fixed nitrogen, 461—462 biological factors, 459 factors affecting growth, 457—461 grain yield, 465—467 historical background, 455—457 N 2 fixation physiology, 455—457 symbioses, 464— 465 physical factors, 457—459 prospects, 467 B a c illa r io p h y ta ,

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CRC Handbook of Microalgal Mass Culture

soil factors, 459 Biogenic toxins, 372— 373 Biological contaminations, 377— 381 Biological factors in blue-green algae growth, 459 Biological growth process, 474— 477 Biological principles for outdoor mass cultures, see also Outdoor mass cultures, 285— 312 Biological steady state, high rate oxidation pond, 332— 334 Biological value (BV), 358— 360, 362, 369— 370, 388 Bioluminescent systems, 436 Biomass, 1— 24, 310 advantages, 4 commercial production, 81 composition, 78 concentrations, 293, 311 procedures for determination of, 120— 123 conversion sources, 6 cyanobacterial, 224 dark respiration, influence of temperature on, 80 defined, 6— 8 developed countries, 1 developing countries, 1 DOC-loaded systems, 285 Dunaliella, 233 estimation of, 142 Europe's energy requirements, 1 food production, use, and increased productivity, 7, 9— 15 free energy, 307 fuels, costs of production of, 19 greatest yield per pond area, 292 high temperature, 80 high yield, 289 increased productivity and, 15— 19 macroalgae, 200 macronutrient elements, 156— 188 microalgal, 30, 34— 35, 200 oil and hydrocarbon plants and fungi, 19— 20 output, 81, 310 per unit area, 306 Phaeodactylum, 238 photosynthesis, 1 ,3 , 6— 7 problems, 4 processing, 350— 354 production Chlorella, 209 organic carbon nutrition, 162 RBP carboxylase, 53 summer, 296 thermophilic algae, 206 various depths, 292 productivity, 1 programs Brazil, 17 Europe, 18— 19 United States, 17— 18 rate of increase in, 285 solar irradiance, available quantity of, 286 Spirulina production, 223— 224

stored, 3 world’s energy, 1 yields, 306 Biomass protein (BMP), 339 Biophotolysis system, 439 Biosynthesis, glycinebetaine, 94 Biosynthetic pathway for chlorophyll, 65 Biotic factors, 111 Biotin, 180, 189, 227, 349 Biri, 339 Bismuth, 181 Blackbody, 46 Blackened thermocouples, 46 Black sulfide deposits, 109 Blanket materials, 278 Blood cholesterol reduction, 425 Blood platelet aggregation, inhibition of, 425 Blooms conditions leading to, 104— 108 cyanobacterial, 320 natural conditions, 101— 104, 239 processes leading to, 104— 108 species-specific, 326 spontaneous disintegration, 299— 300 unialgal, 217 Blue deserts, 108 Blue-green algae (BGA), see Cyanobacteria Blue light, 64, 69, 71 Blue pigment phycocyanin, 66 BOD concentration, high rate oxidation pond, 332— 333 Bold’s basal medium, 129— 130 B o m b y x m o r i P . , 401 Boron, 181, 183— 184 B o ty r o c o c c u s b r a u n ii, 20, 101— 102, 239, 431— 432 Boundary layers, 223 B ra c h io n u s , 327, 378, 402 c a ly c if lo r u s , 326 p lic a tilis , 35 r u b e n s, 326 Brackish water, 240 algaculture in, 487 B o ty r o c o c c u s b r a u n ii, 239 P o r p h y r id iu m , 238 S p ir u lin a , 214, 217 Brazil, biomass programs, 17 Bread, supplementary effect of algae, 365— 370 Breeding, B o ty r o c o c c u s b r a u n ii, 239 Brine lakes, 230 Brittany, 26, 27 Broiler chickens, feed for, 394, 397 Bromine, 186 Bronchitis, 424 Brown algae, 61, 63— 64, 6 6 , 148, 158 Buffering capacity, 220 Bulgaria, microalgae production units, 31 B u m ille r io p s is , 165 Buoyancy, 74— 77, 234, 239 Burma, microalgae, consumption of, 28 Bums, 406

493 Butter, 19 Butyrate, 162

c C3, 55, 56, 304 C4, 53, 55— 56 Ca, see Calcium Ca+, S p ir u lin a mass culture, 222 Cadmium, 186, 228 Calcification, 61 Calcified bodies, 59 Calcium, 155, 157, 181— 182, 346 S p iru lin a , 227 teeth formation, effect on, 384 Calcium carbonate, 199, 486 Calculated production, 293 California albazod processing, 36 high-rate algal ponds, 35— 36 Calorie consumption, 10 Calories, 46 C a lo th r ix

444 186 s c o p u lo ru m , 95 sp., 166, 347, 456 Calves, feed for, 400 Calvin Cycle, 53— 56 CAM, 53 Canada, C h o n d r u s production, 27 Cancer, 229— 230 Candela, 46 Candle, 46 Canthaxanthin, 432 Capital costs, S p ir u lin a , 224 Carbamoyl phosphate synthetase, 169, 171 Carbohydrates, 72, 169, 188, 342, 345, 349 accumulation, 211 C h lo r e lla , 201 concentrations, 240 content, 71, 210 photosynthesis of, see also C 0 2 assimilation, 52—55 production under red light, 71 synthesis, 52 Carbon, see also specific types, 157— 163, 183 assimilation, C h lo r e lla , 210 buoyancy, 77 C h lo r e lla yields, 210 concentration, D u n a lie lla , 231 culture medium, 126 D u n a lie lla , 231 growth rate of S c e n e d e s m u s , 237 inorganic, 158— 160 limitation, 289 metabolism, interaction of nitrogen with, 174— 111 nutrition cost, 486 b r e v is s im a ,

p a r ie tin a ,

D u n a lie lla , 231 organic, 160— 163 organic matter through photosynthesis, 47 species, 85 Carbonates, 27, 217, 222 Carbon compounds, 167 Carbon dioxide, see C 0 2 Carbon fixation, 85 Carbonic anhydrase, 90, 160 Carboxylation of RBP, 53— 54 Carboxysomes, 57 a-Carotene, 199, 227, 350 (3-Carotene, 64, 66 , 71, 199, 350, 432—433, 441, 445, 485 accumulation, 94 cancer protection from, 230 D u n a lie lla , 233—234 microalgae, 35 S p ir u lin a , 225, 227 (3-Carotene-containing globules, 94 (3-Carotene to chlorophyll ratio, 234 Carotenes, 199, 421 production, 231 S p iru lin a , 227 Carotenoids, see also specific types, 64, 6 6 , 71, 199, 301, 432—434 reduction in total, 72 S p ir u lin a , 225, 227 thylakoid lamellae, 59 Carrageenans, 27, 441, 445 Carriers of technological changes, 473 C a r te r ia sp., 151 Cascade screens, 256 Catalases, 299 Cattle, feed for, 400— 401 Cd, see Cadmium Cell breakage, S p iru lin a , 224 Cell chlorophyll, 234 Cell color, C h lo r e lla , 203 Cell composition, 78 Cell content, phycocyanin, 220— 221 Cell counting chambers, 121 Cell counts (CC), 83, 120— 121, 142 C h lo r e lla , 205 colony counts, 121 direct microscopic counting, 120— 121 electronic counting, 121 Cell death, 80, 85, 292, 309 Cell density, 290, 295, 315— 316 C h lo r e lla , 205, 212 high, 306 mass culture, 209 Cell diameter C h lo r e lla , 203 high light, 96 low light, 96 low nitrogen, 96 S p ir u lin a , 213 Cell dimensions C h lo r e lla , 201— 202 S p ir u lin a , 213

CRC Handbook of Microalgal Mass Culture

494

Cell division, 63, 85, 231 C h lo r e lla , 207— 208 delay in, 95 inhibition, 85, 94— 95, 208 P h a e o d a c ty lu m , 236 salinity, effect of, 94— 95 Cell enlargement, C h lo r e lla , 207— 208 Cell excretion, 316 Cell fission, 217 Cell growth, procedures for determination of, 120— 123 Cell hydration, 90 Cell length light, 95— 96 temperature, 95— 96 Cell lysis, 316 Cell mass, C h lo r e lla , 205 Cell morphology, 95— 97 Cell movement, temperature, effect of, 85 Cell number, dark cells, 209 Cell permeability, 78 Cell population, 314 Cell quota, 108 Cell response to environmental factors, see Environ­ mental factors Cell shape, S p ir u lin a , 214 Cell size, 286 C h lo r e lla , 205 S p ir u lin a , 214 Cell suspensions, 292 Cell temperature, see also Temperature, 78 Cellular carotenoid levels, 111 Cellular components, prokaryotes, 56 Cellular damage, 302 Cellular osmoregulation, D u n a lie lla , 232 Cellulose, 59, 61, 199, 348 Cell volume, C h lo r e lla , 205, 207 Cell wall, 59, 216, 348— 349 cellulose, 348 C h lo r e lla , 201— 202 composition, 199 D u n a lie lla , 232 empty, 201— 202 four layers, 56 microfibrils, 348 prokaryotes, 56 structure, 61 Cell weight, C h lo r e lla , 205 Cell yield, S p ir u lin a , 2 2 0 Central Africa, alkaline lakes, 217 Central lumen, 61 Central pyrenoid region, 238 Centrifugation, 255— 256, 336— 337, 485 C h lo r e lla production, 212 Centroplasm, 56—58 Cereal production, 7, 365— 370 Cesium, 181 C2H2, 167 C h a e to c e r u s la u d e r i,

444

p s e u d o c u r v is te u s ,

444

sim p le x ,

164

s im p le x c a lc itr a n s , s o c ia lis ,

430

444

sp., 164 Chain of electron carriers, 48— 49 Channel length-to-width ratio, 270 C h a r a g lo b u la r is , 443 C h a r a c iu m p o ly m o r p h u m , 424 Charcoal, 21 Charophyceae, 62 CH 3CN, 167 CH 3COOH, 212 Cheese, 19 Chelated iron, P o r p h y r id iu m , 238 Chemical composition, 86 amino acid, 342— 344 ash, 346 cell wall, 348— 349 C h lo r e lla , 210—211 environmental conditions, effect of, 341 lipids, 345— 346 microalgae, 341— 350 nucleic acids, 346— 347 pigments, 350 polysaccharide, 349— 350 protein, 341— 342 S p ir u lin a , 225, 227— 228 temperature, effect of, 82— 83 vitamins, 347— 349 Chemical energy, 47—48, 306 Chemical flocculation, 258— 259, 337 Chemical potential, 90 Chemical products, 27— 28 Chemicals, 485 algal movement, 73 Chemical score, 343 Chemical specification, 185 Chemiosmotic hypothesis, 51 Chemoheterotrophy, 147 Chemolithotrophic heterotrophy, 147 Chemostat, 119— 120, 137— 139 Chemostat cultures, 210 Chemotrophs, 147 Chicken feed, 393— 398 Chilling temperature, 84— 85 China kelp farming, 27 microalgae, consumption of, 28 microalgal processes, 26 piggery wastes, 30 reforestation, 17 C h ir o n o m u s , 378 Chitin, 61 Chitosan, 259 Chlamydomonadaceae, 240 C h la m y d o m o n a s

62—63, 73, 80— 81 antibiotic activity, 443 calcium, 181 carbonic anhydrase, 160 growth medium for, 130— 131

r e in h a r d tii,

495 hydrogen production, 439 lipids, 422 magnesium, 181 photorespiration, 305 photosynthesis, 305 rates of 14C 0 2(0) and H 14C 0 3- fixation, 161 sterols, 430 triacontanol, effects of, 189 spp., 60, 65, 96, 240 acetate, 176 amino acid composition, 344 ammonium assimilation, 171 blooms in nature, 101 C 0 2 uptake, 189 hydrogenase activity, 438 myxobacteria attacking, 322 nitrogen utilization, 165 photorespiratory N cycle, 304 synchronized cultures, 120 triacontanol, effects of, 189 variabilis, 184 Chloramphenicol, 65, 302 Chlorella australis, 180— 181 Candida, 430 culture, growth of, 205 dry matter, 212 ellipsoidea amino acid pattern, 344 biochemical and physiological properties, 201 growth rates, 162— 164 magnesium, 182 sterols, 430 yields with carbon and nitrogen sources, 210 emersonii, 92— 94 factories, 212 fusca, 202 biochemical and physiological characters, 204 C 0 2 concentration effect on fatty acid composi­ tion, 346 iron deficiency, 183 nutrient levels, 150 fusca var. rubescens, 203 fusca var. vacuolata, 203 homosphaera, 202— 204 industry, 211 kessleri, 202, 204 luteoviridis, 203, 204 minutissima, 202— 204 protothecoides, 65, 202— 204 pyrenoidosa, 87 amino acid pattern, 344 biochemical and physiological properties, 201 chemical composition, 342 cysteine, 180 growth curves, 162 growth rates, 163— 164 lipid content, 423 lutein, 432 molybdenum, 185— 186 purine uptake, 173

sodium, 181 starch, 350 sulfate uptake, 180, 181 temperature, response to, 206— 207 vitamin content, 349 yields with carbon and nitrogen sources, 210 pyrenoidosa Chick 82, 211 pyrenoidosa Pringsheim 82T, 211 pyrenoidosa strain C-28, 163 saccharophila, 202— 204 nitrogen utilization, 16, 165 sorokiniana, 81, 204, 423 sp. K, 211 spp., see also specific topics, 25, 28— 35, 65, 69— 70, 81, 84, 86 , 92, 199— 212 acceptability, 390— 392 alcohols, 347 amino acid fortification, effect of, 369 anaerobic growth, 163 antibiotic activity, 443 automatic synchronization device, 135 biochemical characters, 204 biochemical properties, 201 biomass production, 209 calcium effects, 155 cell division, 207— 208 cell enlargement, 207—208 chemical composition, 210— 211, 341 cholinesterase activity, 230 circular cultivation ponds, 245 closed cultivation system, 246 commercial production, 211— 212 comparability data, 362 competition with Spirulina, 317, 318 contamination by, 317 contamination of Spirulina, 223 copper, 186 culture systems for production of, 212 dark cells, 208— 209 digestibility, 351— 352, 354, 356 dominance, 317 ecology, 200 fungal contamination, 378 glucose, 176 growth conditions outdoors, 203— 205 growth medium for, 128 growth rates, 209 heavy metal contamination, 375 heavy metal content, 375— 376 heterotrophic growth, 210 high-methionine mutants, 344 hydrocarbons, 347 hydrogenase activity, 438 infection by Stylonichia sp., 322 infestation with, 313 light cells, 208— 209 light-limited chemostat cultures, 307 mesophilic, 206 methane production, 439 microstrainers, 257 mixed culture with Spirulina, 319

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morphology, 200— 203 N -8 medium, 129 nitrate reductase structure, 167— 168 nitrite reduction, 174 nitrogen starvation, 211 nitrogen utilization, 165 nutrient levels, 150 nutrients, effect of, 317 optimizing growth conditions outdoors, 203— 205 organic carbon nutrition, 162 organic toxic compounds, 377 osmoregulation, 441 partial pressure of oxygen, effect on yield, 304 pH, 317 P h a e o p h o r b id e s , 350 phosphate starvation, 179 photosensitized skin irritations, 350 photosynthetic rate, 303 physiological properties, 201, 204 pig feed, 398— 399 poultry feed, 393— 398 processing, 360 protein efficiency ratio, 357, 367— 368 protein extract of, 361 reproduction, 200 ruminants, feed for, 400 salt tolerance as species-specific character in, 203 screening, 257 section A u c h lo r e lla , 201 shade adaptation, 138 side effects, 387— 388 sloped cultivation units, 248 starch, 442 sterols, 347 strontium, 181— 182 sulfur-containing amino acids, 180 supplementary effect of algae, 366— 370 synchronized cultures, 120 taxonomy, 200— 203 temperature, effect of, 205— 207, 317 therapeutic properties, 406 thermophilic, 206 turbulence, 207— 209 world programs for mass cultivation, 306 yield, 207, 294—295 zooplanktons invading, 325 v a r ie g a ta , 427 v u lg a r is, 25, 28, 81, 202 amino acid oxidase, 437 amino acid pattern, 344 chemical composition, 342 fatty acid composition, 345 lipid content, 423 methionine, 180 net photosynthetic reaction rate, 290 nutritive value, 361 phosphate starvation, 179 photorespiration and photosynthesis, 305 temperature, 320

yield,

210

203 200, 201, 204 v u lg a r is B e ije r in c k var. v u lg a r is , 202 z o fin g ie n s is , 202— 204 Chlorella Growth Factor (CGF), 34, 212 C h lo r e lla lis , 200 Chloride, 182, 227 Chlorine, 182 3-Chlorocarbonylcyanide phenylhydrazine (CCCP), 160 Chlorococcales, 200 v u lg a r is

var.

v u lg a r is,

v u lg a r is B e ije r in c k ,

C h lo r o c o c c u m o le o fa c ie n s ,

424

sp., 391 v a c u o la tu m ,

438

96 456 /?-Chlorophenyl-l,l-dimethylurea (CMU), 163 Chlorophyceae, 62—63, 65— 66 alkaline water, 217 amino acid carriers, 172 polar lipids, 422 urea amidolyase, 172 vitamin needs, 189 Chlorophycophyta, 234 C h lo r o p h y ta , see Green algae Chlorophyll-a, 64, 66 , 70, 72, 179, 199, 350, 432 content, euphotic zone, 219 nutrient deficiencies, 188 S p ir u lin a , 227— 228 synthesis, 183 Chlorophyll-a 670, 64 Chlorophyll-a 680, 64 Chlorophyll a/b, 64, 90, 91 Chlorophyllase, 69, 350 Chlorophyll-b, 59, 64, 199, 350, 432 Chlorophyll-c, 64, 432 Chlorophyll/carotenoids, 91 Chlorophyll-d, 64, 432 Chlorophyll-protein complexes, 64 Chlorophyll/P700 ratio, 70 Chlorophylls (Chi), 50, 64— 66, 199, 303, 421, 432— 435 absorbance, determination of, 122— 123 biosynthesis, 65 concentration, 302— 303, 313 content, C h lo r e lla , 211—212 copper depletion, 185 dark cells, 208 destruction of, 69 distribution among algae, 65 dry weight ratio, 319 D u n a lie lla , 234 extraction, 122 formation, 6 6 , 212 increase in, 314 light cells, 208 magnesium, 182 molecule, 48 nutrient deficiencies, 188 C h lo r o g lo e a f r its c h ii, C h lo r o g lo e o p s is ,

4 97 per cell, 90— 91 procedures for determination of, 122— 123 production, 234 reaction-center, 48 separation of cells, 122 spectrophotometrical determination of concentra­ tion of, 65 synthesis, 65—66 thylakoid lamellae, 59 triacontanol, effects of, 189 UNESCO procedure, 123 Chloroplast, 48, 60, 62— 63, 94, 202 band-shaped, 202 C h lo r o p h y ta , 64 cup-shaped, 202, 230 cyclic electron flow, 51 division, 63 endoplasmic reticulum, 62 envelope, 63 evolution, 63— 64 eyespots, 63 genome, 63 girdle-shaped, 202 illuminated, 48 laminate, 234 mantle-shaped, 202 morphology, C h lo r e lla , 201— 202 parietal, 202 photophosphorylation, 50 Photosystem I, 49 saucer-shaped, 202 stellate, 238 thylakoid membrane, see Thylakoid membrane thylakoid structure, 62—63 C h lo r o s a r c in o p s is , 96 n e g e v e n s is , 424 Chlorosulfolipids, 422 Chlorphyceae, 200 CH3NC, 167 C6H 120 6, 212 Cholesten-7-ol-3, 227 Cholesterol, 227, 406— 407 Cholinesterase activity, 230 C h o n d r u s , 27, 441 c ris p u s , 26 Chromatic adaptation, 58, 71 Chromatophores, 50, 203 Chromium, 186 C h r o m o p h y ta , 64 Chromoplasm, 56 Chronic toxicity, 381 Chroococcacean cyanobacteria, 56, 166 C h r o o c o c c id io p s is , 166, 456 C h r o o m o n a s , 430 s a lin a , 423, 426 C h r y s o c h ro m u lin a sp., 164 Chrysomonads, 189 Chrysophyceae, 62, 65, 200, 422 Chrysophycophyta, 236 C h ry s o p h y ta , 140, 164 Chrysophytes, 63

Chu 10, 129 C h y tr id sp., 321 Chytrides, 321 Chytridia, 334 Chytridiales, 321 C h y trid iu m sp., 322, 378 Cicatrization of wounds, 229 C ilia ta , 322, 378 Ciliates, 323 Circular DNA, 61 Circular ponds, 245 Citrulline, 171 Cl, see Chloride C la d o p h o r a

430 439 Clams, feed for, 402 Clay blanket, 278 Clean culture, 285 Climate for pond construction, 265— 266 Clinical applications, S p ir u lin a , 229— 230 Clinical studies of algal nutrition, 391— 392 Closed fermenters, C h lo r e lla production, 212 Closed systems, see also Batch cultures, 82, 118— 119, 245—247 C lo s tr id iu m , 186 C 0 2, 4 6 - 4 8 , 85, 126, 163 algaculture resource, 486 algal respiration, 160 assimilation, 52—55, 90—91, 219, 300 buoyancy, 77 C h lo r e lla production, 212 concentration, 300, 317 C h lo r e lla , 210 fatty acid composition, 346 outdoor mass cultures, 289 photorespiration inhibition, 304 competition with 0 2, 304 condensation, 159— 160 conversion into glucose, 55 cultivation area increase, 485 cycling, 3 depletion through photosynthesis, 313 distribution, 254— 255 D u n a lie lla , 231 fixation, 53, 55, 62, 75, 91, 157, 160, 178 industrial production of C h lo r e lla , 212 limitation, buoyancy, 77 photoassimilation, 320 photosynthetic assimilation of, 43 polyphosphate formation, 180 pond feeding provisions, 282 pure gas, 311 rate of exchange with oxygen, 56 reduction, 48 reduction path, 53 requirement for nitrate, nitrite and ammonium up­ take, 176 S c e n e d e s m u s , 236 S p ir u lin a , 220, 224 sulfate uptake, 181 f le x u o s a ,

g lo m e r a ta ,

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CRC Handbook of Microalgal Mass Culture

supply, 31, 32, 209, 254—255 synthesis of organic compounds, 158 uptake, 189 utilization, 254— 255 withholding, effect of, 319 C 0 2-acceptor molecules, 55 C 0 2-free air, 319 14C 0 2, 53, 76, 173 C 0 3, 85 C 0 3 2, 158 Coagulation, 336 Coarse granular blankets, 278 Cobalamine, 349 Cobalt, 181, 185— 187 C o c c o c h lo r is , 85— 86 p e n io c y s tis , 85, 94 C 0 2 fixation, 160 cobalt, 186 inorganic carbon accumulation, 161 photorespiration and photosynthesis, 305 Coccoliths, 59, 61 C o e la str u m

29, 33 spp., 357, 362, 438 hydrogenase activity, 438 protein efficiency ratio, 357 Coenobia, 234 Coenzyme NADP +, 48 Cold shock, 84 Colonial algae, 239 Colonies of cells, 72, 199— 200, 239 Colony counts, 121 Colorimeter, 121 Colors, 43, 45 C o lp o d a a te in ii, 323 Combined culture systems, C h lo r e lla production, 212 Commercial algaculture, 485 Commercial energy, 15 Commercially grown mass cultures, 84 Commercial production C h lo r e lla , 211— 212 S c e n e d e s m u s , 236 S p ir u lin a , 223—224, 226, 228 Commercial S p ir u lin a plant, 226 Common Agricultural Policy, 18 Common carp, feed for, 403 Community forestry, 3 Compensation depth, 105, 112 Compensation irradiances, 71 Compensation point, 309, 315 Competition, 109— 111, 317— 318 Competitive exclusion, 111 Composition of biomass, 78 Concentrated seawater, P o r p h y r id iu m , 238 Concrete block pond walls, 280— 281 Conductivity, 217, 218 Consciousness, treatment of disturbances of, 425 Constant (A), 79 Constant incident light, 285 Constant temperature, 285 s p h a e ric u m ,

Consumption, 26, 28—29 Contaminants, see also specific types, 317, 320 biological, 377— 381 environmental, 373— 377 S p ir u lin a production, 223 Contamination, see also specific factors, 317 control of, 326 D u n a lie lla , 233 other species, 313 Continuous centrifuge, 235 Continuous cultivation, P o r p h y r id iu m cru en tu m , 239 Continuous culture, 83, 119— 120, 155, 222 Continuous illumination, 51 Continuous light, 71— 72, 288 Continuous light (LL) cultures, 71 Continuous mixed culture, 319 Contraception, 424 Contractile vacuoles, eukaryotes, 59 Conversion efficiency P o r p h y r id iu m , 238 S p ir u lin a , 219 Conversion factors, 46 Conversion of solar energy, 307 C o n y a u la x c a tte n e lla , 427 C o p e d o d a , 325 Copolymers, 57 Copper, 95, 181, 184— 185 Com, C 0 2 assimilation, 56 Corrugated asbestos-cement roofing panels, 280 Corrugated asbestos-cement wall, 281 C o sm a r iu m , 165 la e v e , 423 Cosmetics additives, 425 Cosmetics coloring, 435 Cost-benefit analysis, 485— 486 Costs, see specific types Cost structure, 475— 477 Cost of unit-product, 485 Coulter Counter, 121 Critical depth, 107 Critical temperature regions, 85 Cross wall, 58, 214— 215 C r u c ig e n ia a p ic u la ta , 438 Crude fiber, S p ir u lin a , 227 Crude protein, 341 S c e n e d e s m u s , 236 S p ir u lin a , 227 Crushed rock bottom lining, 281 Crustaceans, 325 Crustacean zooplankton, 327 Cryophilic algae, 78 C r y p te c o d y n iu m c o h n ii, 427 C r y p to m o n a s sp., 427 Cryptophyceae, 62, 65—66 C r y p to p h y ta , 139 C r y p to th e c o d in iu m c o h n i, 430 Cryptoxanthin, 227 a-Cryptoxanthin, 350 C r y s o c h r o m u lin a k a p p a , 423 Crysophytes, 179

499 C te n o c la d u s c ir c in n a lis ,

181 403

C te n o p h a r y n g o d o n id e lla ,

Cu(II), 299 Culture density, 208, 326 depth, 286, 292 deterioration, 219, 303, 313— 314, 316, 322 S p ir u lin a , 220, 223 laboratory techniques, see Laboratory techniques for microalgae cultivation media, 126 NaCl concentration, 92 S c e n e d e s m u s , 235 temperature, see also Temperature, 78 performance, evaluation of, 312— 327 systems, C h lo r e lla production, 212 units, mass production, 30 Cup-shaped chloroplast, 230 C y a n id iu m , 165 c a ld a r iu m , 65, 430 C y a n o b a c te r ia (blue-green algae), see also Biofertil­ izer, 25, 64, 74, 109, 217 agar, 133 alkaline range, 320 alkaline water, 217 Allen and Amon medium, 134 Allen’s medium, 126— 127 ammonium assimilation, 170— 171 axenic clones, 141— 142 biofertilizer, 455— 471 biomass, 224 boron, 183 calcium, 181 cell length, effect of light and temperature on, 95— 96 cell organization, 56 chromatic adaptation, 71 C 0 2 concentrations, 317 competition with green algae, 317 diet for zooplankton or planktonic rotifers, 326 filament, 58 future prospects for use of, 485 hydrogenase, 158 lipids, 428—429 morphological changes, 95— 96 myxobacteria attacking, 322 nitrogen, 164 nitrogenase inhibition, 168 nitrogen deficiency, 173 nitrogen-fixing, 166, 455— 457 nutrient deficiencies, 188 O, concentrations, 157 0 2 evolution, 85 phosphate deficiency, 179 photolithotrophy, 148 photooxidative death, 302 phycobilisomes, 58 pigmentation, 199 polar lipids, 422 restriction enzymes in, 437 selenium, 188

sloped cultivation units, 248 sodium, 181 spontaneous disintegration, 299— 300 succession of, 457— 458 surface plating, 142 viruses attacking, 320 Cyanobacterium Synechocystis DUN 52, 93 Cyanocobalamin (B12), 227 Cyanophages, 320 C y a n o p h o r a p a r a d o x a , 438 Cyanophyceae, 65—66, 189, 200 Cyanophycean starch, 63 Cyanophycin, 179 Cyanophycin granule polypeptide (CGP), 173 Cyanophycin granules, 57— 58, 214 C y a n o p h y ta , 25

fluorescence, 139 growth rates, 164 Cyclic electron flow, 51 Cyclic phosphorylation, 51, 182 Cyclic photophosphorylation, 59 Cyclitols, 228 Cyclohexanetetrol, 440 Cycloheximide, 65, 93, 211 Cyclopropane, 167 C y c lo te lla , 70, 96, 165 c r y p tic a , 164 Cylinder, 61 Cylindrical cells, S p ir u lin a , 212 Cylindrical filaments, 214 C y lin d r o s p e r m u m , 166, 456 lic h e n ifo rm , 171 C y lin d r o th e c a fu s if o r m is , 184, 187— 188, 440 C y p r in u s c a r p ia , 403 Cysteine, 180 Cystine, 180, 227 Cytochrome, 168 Cytochrome b, 50 Cytochrome b563, 50 Cytochrome c reductase activity, 168 Cytochrome f, 50 Cytokinins, 189 Cytoplasm, 61, 85 Cytoplasmic continuity, 61 Cytoplasmic membrane, 85 D u n a lie lla , 232 Cytoplasmic ribosomes, 65 Czechoslovakia microalgae production units, 31 microalgae research, 33

D 235 Daily cell counts, C h lo r e lla , 205 Daily radiation course, 295 Daily temperature course, 82, 295 D a p h n ia , 324, 402 Dark cells, C h lo r e lla , 208— 209 Dark growth, C h lo r e lla , 202— 203 D a c ty lo c o c c u s n a g e li,

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CRC Handbook of Microalgal Mass Culture

Dark-heterotrophic conditions, C h lo r e lla , 210 Dark process, rate of, 209 Dark respiration, 80— 81, 307 Dark time, 288 Day length, 88— 89 Day temperature, C h lo r e lla , 205— 206, 209 DCMU, see 3-(3,4-Dichlorophenyl)-l ,1-dimethyl urea Dead algae, 299 Death, 104, 303 Death phase, 118 Decay, 309 Decolorization of algae, 391 Deep culture, 209 Deep culture in closed fermenter, C h lo r e lla produc­ tion, 212 Deficiencies, 66 Deforestation, 21 Degenerative diseases, 230 Dehydration of algal mass, 260— 262, 487 Dense cultures, P h a e o d a c ty lu m , 238 Density gradients, separation of microalgae by, 142 2'-Deoxy adenosine, 190 Deoxyribonucleic acid, see DNA Depletion-repletion studies, 364— 366 Depth integral, 101, 107 Dermal toxicity, 385 D e r m o c a r p a , 166, 456 5-6-Desaturase, 230 Desert, 47 Detention period, P o r p h y r id iu m , 238 Detritus, high-rate algal ponds, 36 Developed countries biomass production, 1 grain production, 10 Developing countries biomass production, 1 calorie consumption, 10 fuel wood shortages, 2 grain production, 10 importance of biomass energy, 15 scarcities, 3 Dextrin, 349 D ia p h a n o s o m a b r a c h y u ru m , 326— 327 Diatomeae, 321 Diatoms (B a c illa r io p h y ta ), 29, 61, 63, 140, 158 alkaline water, 217 blooms, 109 boron, 183— 184 calcium, 181 heterotrophy, 148 nitrogen, 163 phosphate deficiency, 179 selenium, 188 silicon, 187 sloped cultivation units, 248 valves, 236 vitamin needs, 189 world programs for mass cultivation, 306 Diatom seawater-agar, 133— 134

3-(3,4-Dichlorophenyl)-l ,1-dimethyl urea (DCMU), 93, 160, 320 nitrate uptake, 174, 176 nitrite uptake, 174, 177 S p ir u lin a , 219 sulfate uptake, 181 Diet, 7, 21 Digestibility of algae, 350— 356, 359, 362, 385, 389, 401 apparent, 353 humans, 388 processing and, 356, 358 S p ir u lin a , 227 Digestibility coefficient (DC), 358— 360, 362, 369— 370, 401 Dihe, 221— 222, 228— 229, 339 Dihomo-GLA (DGLA), see Dihomo-'y-linolenic acid Dihomo-7 -linoleic acid (DGLA), 230, 424 Dihydro-7-cholesterol, 227 Dilution rate, 119 Dinitrophenol, 188 Dinoflagellates, 63, 101— 102, 109, 158, 189 Dinophyceae, 59, 62—63, 65—66 Dinoxanthin, 432— 433 1,5-Diphosphate carboxylase, 182 D ip lo c y s tis a e r u g in o s a , 186 Direct illumination, 291 Direct microscopic counting, 120— 121 Direct sunlight, 205 Disc-shaped cistemae, 61 Diseased algal cultures, 322, 326 Dissolved C 0 2, 318 Dissolved inorganic carbon (DIC), 77, 85 Dissolved organic carbon (DOC), 285, 311 Dissolved organic compounds, 104, 108 Dissolved oxygen, 219, 223, 303, 314 Dissolved oxygen concentration, 295 decline in, 302— 303 measurement, 314— 315 outdoor mass cultures, 298 pond maintenance, 312 units of weight, 298 Distal ends, 216 Divalent ions, 62 Divalent reduction of hydrogen peroxide, 299 Divalent reduction of oxygen, 299 Division cycle, 234 DNA, 58, 179, 346, 371 C h lo r e lla strains, 204 determination of, 141 Feulgen staining, 141 S p ir u lin a , 227 DNA molecule, 63 DNA synthesis, 69, 95 Dominance, 317 Dominant species, 317 Double fibers, 61 Double-substrate limited growth, models for, 155— 156 Doubling speed of flow, 290 Doubling time of culture, 30, 86— 87, 118, 239

501 Down welling irradiance, 104 Drum drier, 235 Drum drying, 36, 260— 261, 485 Drying of algae, 487 effect on digestibility, 350— 353, 385 Dry weight of algae, 313 C h lo r e lla , 211 S p ir u lin a , 229 Dry weight measuring, 122 D u n a lie lla

35, 94 amino acid pattern, 344 |3-carotene, 432 fatty acid composition, 345 low chlorophyll-containing cells, 234 massive (3-carotene accumulation, 233 pigment content, 234 pigments, 350 salt concentration, effect on glycerol productiv­ ity, 233 b io c u la ta , 342 p a r v a , 90—91, 95, 131— 132, 422 p r im a le c ta , 344, 423 s a lin a , 94, 111, 231 acceptability, 391 chemical composition, 342 hydrocarbons, 431 lipid content, 423 low chlorophyll-containing cells, 234 spp., see also specific topics, 20, 32, 35, 90, 92, 230—234 (3-carotene, 233— 234 chemicals from, 485 extracellular salt concentration, effect of, 234 glycerol production, 233 habitat, 230— 231 halophilic types, 231 halotolerant types, 231 hypertonic conditions, 231— 232 hypotonic conditions, 231— 232 nonprotein nitrogen, 342 osmoregulation, 231— 233, 440— 441 osmotic metabolites, 440 potential for practical exploitation, 445 taxonomy, 230— 231 te r tio le c ta , 70, 83, 92, 422 fatty acids, 344 growth limitation, 153 Dung, 21 b a r d a w il,

E Earlier growth phase, 285 Earthrise Farms, 224— 225 East Africa, 112 Eastern Mediterranean, 108 Echinenone, 227, 350 Ecology, 326, 474 C h lo r e lla , 220 S p ir u lin a , 217— 219

Economic aspects, 473—487 Economic potential, see also specific types of mi­ croalgae, 199— 243, 307, 485 C h lo r e lla , 200—212 D u n a lie lla , 230—234 P h a e o d a c ty lu m , 236— 238 P o r p h y r id iu m , 238— 239 S c e n e d e s m u s , 234— 237 S p ir u lin a , 212— 230 E c to c a r p u s , 165 Eczema, 406 Edible algae, 26 EDTA, 126, 181— 182, 185, 209 Efficiency of plant species, 304 S p ir u lin a mass culture, 223 Efficient parasite control, 326 Efflux of photorespiratory C 0 2, 304 Efflux rates, D u n a lie lla , 233 Egg protein, S p ir u lin a , 222 Egg yolk color, 35 Egypt, microalgal research, 34 Eicosapentaenoic acid (EPA), 424 5,8,11,14-Eicosatetraenoic acid, see Arachidonic acid Einstein, 43, 45, 52, 307, 311 Ein Yahav Algae, Israel, 225— 226 Electrical power and instrumentation for ponds, 283 Electric stimuli, algal movement, 73 Electroflocculation, 259— 260 Electromagnetic radiation, 46, 64 Electromagnetic spectrum, 43—44 Electron acceptor, 157 Electron-acceptor molecule, 48 Electron carriers, 48— 50 Electron flow, 48— 51 Electron hole, 49 Electronic counting, 121 Electron microscopy, 62, 95, 213, 214 Electrons, 48—49, 52, 157— 158, 306 Electron-translucent DNA-containing areas, 62 Electron transport, see Electron flow Electrostatic bridges, 62 Elementary composition of algae, 151 E le u sin e c o r a c a n a , 366 Ellipsoidal cells, C h lo r e lla , 200— 203 Empirical applications, 478—481 Empty cell wall, 201— 202 End cells, 214 Endogenous glycerol concentration, 92 Endogenous respiration, 94, 207 Endoplasmic reticulum, 59, 61—62 Endospores, 236 Endosymbiont, 64 Endosymbiosis, 64 Endozoic phytoplankton, 101 Energy content, C h lo r e lla , 210 Energy crop yields, 7— 8 Energy of light, 43— 45 Energy limits, 43—47 Energy plantations, 21 Energy-rich electrons, 49

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CRC Handbook of Microalgal Mass Culture

Enhanced photosynthesis, 55 Enriched seawater, algal growth medium, 132— 133 Enterobacteria, 379 Enteromorpha compressa, 186 Envelopes eukaryotes, 59— 61 prokaryotes, 56 Environmental adaptation, Dunaliella, 230 Environmental conditions, 312, 341 cyanophages, 320 Environmental contaminants, 373— 377 Environmental factors, see also specific topics, 69— 99, 111, 317, 320, 474 biosynthesis of lipids and fatty acids, 422— 423 cell morphology, 95— 97 control of, 314 hydrogen ion concentration, 85— 86 light, 69— 77 light and temperature interaction, 86— 89 monitoring, 314 osmoregulation, 89— 95 salinity, 89—95 Spirulina morphological variants, 214 temperature, 77— 85 Environmental pollutants, microalgae production, 32 Enzyme reactions, 78 Enzymes, 91, 436— 437 Ephydra, 378 Erythrocyte-cholinesterase, 230 Erythromycin, 125 Escherichia coli, 439 Essential amino acid index (EAA), 343 Essential amino acids, Spirulina, 227 Essential fatty acids, see also specific types, 424— 425 Essential unsaturated fatty acids, Spirulina, 226 Ethanol, 162 Eucheuma, 27, 441 Euchlorella, 200 Eudorina elegans synchronized cultures, 120 Euglena gracilis, 73, 85 bacillaris variety, 162 chemical composition, 342 dry weight increase, 304 fatty acids, 427 functional glycolate pathway, 304, 306 growth medium for, 130 magnesium, 182 organic carbon, 162 sterols, 430 Vischer strain, 162 wax esters, 426, 429 spp., 65, 180 blooms in nature, 101 hydrogenase activity, 438 methane production, 439 nitrogen utilization, 165 polysaccharides, 350 poultry feed, 396— 397 protein efficiency ratio, 357

wax esters, 430 Euglenoids, 63, 189 Euglenophyceae, 61— 63, 65 Euglenophyta, 140, 321 Eukaryotes, 59— 64, 164, 171, 179 Euphorbia lathyrus, 20 tirucalli, 20 Euphotic depth, 104— 107, 112 Euphotic zone, 101, 109, 219 Europe, biomass programs, 1, 18— 19 European Economic Council (EEC), 12, 18— 19 Eustigmatophyceae, 62 Eutrophication, 76, 85, 108— 109 Evaporation, 245— 246 pond surface, 90 Spirulina mass culture, 222 Evaporative cooling, 82 Excitation energy, 48 Excited electron, 48 Excretion, 104, 220, 309— 312 extracellular, 108, 311 Exponential growth rate, 294 Exponential phase, 117, 119, 285— 286, 288 Export earnings, 21 Extinction coefficient, 65, 291 Extracellular excretion, 108, 311 Extracellular salt concentration, 234 Extracellular substances, 326 Extracellular transport, 61 Extraterrestrial life support, 32— 33 Extreme salt stress, 90 Exuviella sp., 427 Eyespots, 62—63

F Farming systems, 15 Farmland, 47 Fats, 47, 199 Fatty acids, 344— 345, 421—427, 443 accumulation, Chlorella, 211 C 0 2 concentrations, effect of, 346 composition, 82 patterns, 33 Spirulina, 227 Favorable nitrogen concentrations, 220 Fe, see Iron Fe(II), 299 Feed, 325, 393, 398, 485 Feedback inhibition, 168 Ferredoxin, 50, 95, 168, 171, 174, 183, 306, 457 Ferredoxin-NADP oxidoreductase, 50 Ferredoxin nicotinamide adenine dinucleotide phos­ phate reductase, 95 Feulgen DNA staining, 141 Fibers, 61—62 Fibrous proteinaceous material, 63 Filament length, 96 Filamentous bacteria, 322

503 Filamentous blue-green algae, fluorescence micros­ copy for estimation of, 141 Filamentous cyanobacterium, see also S p ir u lin a , 212, 320, 323, 326 Filamentous heterocystous forms of C y a n o b a c te r ia , 166 Filamentous nonheterocystous forms of C y a n o b a c ­ te r ia , 166 Filaments, 56, 72, 199 floating, 75 motile, gliding along axis, 213 sinking, 75 Filose amoebae, 323 Filtration, 30, 224, 256, 317, 336, 487 purification of microalgae, 125 Fine bristles, 234 Firewood collection, 2 F is c h e r e lla , 166, 456 Fish feed, 224, 402—405 Fish kills, 109 Fish meal, 395 Fish ponds, 110, 324 Fixed costs, 475 Fixed nitrogen, see also Nitrogen fixation, 461— 462 Flagella, 61, 63, 230 Flagellated algae, 148 Flagellated bacteria, 322 Flagellates, 188, 199, 322 Flagellum, 62 Flash effect, 289 Flashing light effect, 252, 270, 277, 291 Flash time, 288— 289 Flat spiral, 213 Flavin, 168 Flavin adenine dinucleotide, 158, 167 Flavin mononucleotide (FMN), 158, 168 Flavoprotein, 50 Floating, 110, 485 albazod processing, 36 Floating filaments, 75 Floating scum, 239 Flocculation, 257— 260 albazod processing, 36 Florida, high-rate algal ponds, 36 Floridean starch, 63 Flotation, see Floating Flour, supplementary effect of algal protein on, 366 Fluorescence, aquatic fungi parasitizing algae, 321 Fluorescence microscopy, 139— 141 Fluorescent tube lamps, 290 Fluorine, 186 Fluorochromes, determination of nuclear DNA, 141 Flux measurement, 46 Flux of solar radiation incident, 46 FMN, see Flavin mononucleotide Foam, 316 Foaming, 316 Foils, 277, 279, 291 Folic acid, 226, 349 Following growth phase, 285

Food, see also Human food, 27, 485 commercial production of S p ir u lin a for, 226 incorporation of algae into, 388, 391 Food coloring, 224, 435 Food for humans, 387, 485 Food gaps, 10 Food-grade PVC, 225 Food production, 1—2, 5, 7, 9— 16, 21, 22 Foodstuffs, see Food; Food for humans Food supplement for humans, S p ir u lin a , 221— 222 Food vs. fuel, 2 Footcandles, 46 Forestry, 1— 3, 5, 17, 18, 21 Forests, 47 F o rm a

215 215 Fortification of algae with amino acids, 369— 370 Fossil fuels, 21 Four-celled colonies, 235 g e itle r i, m in o r,

F r a g ila r ia

321, 322 444 Fragments, 314 France, microalgal research, 32 Free ammonia concentrations, 327 Free ammonia-N, 327 Free energy, 48, 50, 307 Free fatty acids, 421 Freeze drying, 261—262 Freeze etching, 62 F r e n y e lla d ip lo s ip h o n , 71— 72 Frequency, 43—44 culture transfers, 126 Fructose, 162— 163, 202, 236 Fucoxanthin, 64, 6 6 , 188, 433 F u c u s s p ir a lis , 186 Fuel production, 5 Fuelwood, 2, 12, 16 Fumarate, 162 Fungi, 29, 109, 314, 317, 320—322, 378— 379 Fused thylakoids, 62 Fusiform cells, 236 c r o to n e n s is ,

p in n a ta ,

G Galactose, 162— 163 C h lo r e lla , 202, 210 S c e n e d e s m u s , 236— 237 Gallium, 181 Gas constant (R), 79 Gaseous C 0 2, 318 Gaseous exchange, 223 Gas mixture, 304 Gasohol, 18 Gastritis, 406 Gas vacuoles, 74, 76, 101, 110 S p ir u lin a , 213 Gas vesicles, collapse of, 75 GDH, see Glutamine dehydrogenase

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Gelatin liquefaction, C h lo r e lla strains, 204 Genes, transfer of, 485 Genetic manipulation, 485 Genetic modifications, 239 Genophores, 62— 63 Geology for pond construction, 266 Geothermal water, 82 Germanium, 181, 187 Germany, microalgae research, 29, 32— 34 Gibberellins, 189 G ig a r tin a , 441 GLA, see "y-Linolenic acid G le n o d in iu m fo lia c e u m , 426 Gliding manner, movement in, 199 Global distribution of energy use, population, and food supply, 9 Globular to subspherical cells, C h lo r e lla , 201 G lo e o c a p s a , 456 G lo e o th e c e , 166, 456 G lo e o tr ic h ia , 456 Glucan, 228, 349 2-0-a-D-Glucopyranosylglycerol, 94 Glucosamine, 228, 349 Glucose, 31, 52— 54, 93, 162— 163, 176, 183, 349 C h lo r e lla , 202— 203, 210, 212 conversion of C 0 2 into, 55 S c e n e d e s m u s , 235— 237 S p ir u lin a , 220 synthesis, 55 Glucose-6 -phosphate dehydrogenase, 95 Glucose respiration, C h lo r e lla p y r e n o id o s a , 207 Glutamate, 162, 165, 187 Glutamate synthase (GOGAT), 169— 172 Glutamate synthetase, 457 Glutamic acid, 164, 169— 171, 227 Glutamic dehydrogenase, 173 Glutamine, 164— 165, 170— 171, 187,457 Glutamine dehydrogenase (GDH), 169, 171— 172 Glutamine oxoglutarate aminotransferase, 170 Glutamine synthetase (GS), 95, 167, 169— 173, 457 a-Glutaric acid, 169 Glycerides, 422 Glycerol, 91, 344, 421, 440—441, 445, 485 accumulation, 94 concentration, 90 diffusion, 92 D u n a lie lla , 232— 233 efflux avoidance, 92 efflux tolerance, 92 leakage, 92, 232 microalgae, 35 osmoregulation with, 90— 92 production, extracellular salt concentration, effect of, 234 synthesis, 92 Glycine, 93, 164— 165, 227 Glycinebetaine, 93, 94 Glyclglycine, 165 Glycogen, 57— 58, 228, 349 Glycolate, 56 Glycolic acid, 56

Glycolipids, 422 Glycoproteins, 61 Glyoxylate, 56 GOGAT, see Glutamate synthase Golgi apparatus, eukaryotes, 59, 61 Golgi bodies, 61 Golgi complex, 61 Golgi membranes, 61 G o n y a u la x , 436 c a te n a ta , 25 ta m a r e n s is , 430 Gout, 370 G r a c ila r ia , 26— 27 lic h e n o id e s , 425 Grain production, 10 Grain trade, 11 Grain yield, effects of blue-green algae on, 465— 467 Gram-negative rods, 322 Grana, 62 Granular proteinaceous material, 63 Granules, 57 Grass carp, feed for, 403— 404 Grassland, 47 Gravity, algal movement, 73 Gravity flow, 251 Grazing, 101— 102, 104, 109, 323— 324 Great Britain, kelp kilns, 27 Green algae (C h lo r o p h y ta ), 53, 61, 64, 326 boron, 183 calcium, 181 CO, concentrations, 317 competition with blue-green algae, 317 fluorescence, 140 growth rates, 164 heterotrophy, 148 hydrogenase, 158 selenium, 188 viruses attacking, 321 vitamin needs, 189 world programs for mass cultivation, 306 Green light, 64, 97 Green revolution, 11 Gross structure, 199 Groups of cells, 199 Growth, see also Laboratory techniques for microal­ gae cultivation, 124— 135, 314 curve, effect of light intensity, 286— 287 factors, 180 inhibition, D u n a lie lla , 233 limitation, 306 irradiance, 310 outdoor mass cultures, 292 medium temperature, effect of, 80 wastewater systems, 285 outdoors, 203— 205, 237 parameters, changes in, 118— 119 pattern, 303 physiology, P o r p h y r id iu m , 238— 239 rate, 288, 309, 314, 324

5 05 239 205, 207, 209 comparison, 164 D u n a lie lla , 231 high temperature strains, 81 incubation temperature, effect of, 78 interrelation of temperature and irradiance, 292 limiting nutrient concentration, 107 low temperature, 79, 81 pH, 86— 87 P h a e o d a c ty lu m , 238 population density in relation to, 315 P o r p h y r id iu m , 239 potential, 295 S c e n e d e s m u s , 236— 237 S p ir u lin a , 219— 220 strong light, 209 synchronized cultures, 87 temperature, effect of, 78— 80 weak light, 209 yield, 306 Growth regulators, 189— 190 Growth-temperature relation, 81 Guayule, 20 Gynecological diseases, 406 B o ty r o c o c c u s b r a u n ii, C h lo r e lla ,

H H +, 52, 167 H + ions, 49 H + movement, thylakoid membrane, 51— 52 H2 evolution, 167 Habitat D u n a lie lla , 230— 231 P h a e o d a c ty lu m , 236 P o r p h y r id iu m , 238 S p ir u lin a , 217— 219 H a e m a to c o c c u s , 85 p lu v ia lis , 430 Half light saturation constant, 310 Halobacteria, 92 Halophilic algae, 90, 231 H a lo s p h a e r a v ir id is , A l l

Halotolerant algae, 90, 231, 440 Halotolerant wall-less algae, osmoregulation in, 232 Halotrophy, 148 Hamsters, mutagenicity study, 382 Hangovers, 230 H a p a lo s ip h o n , 456 la m in u su s, A l l

Haptonema, 62 Haptonemata, 59 Haptopyceae, 59, 62— 63, 65 H a r tm a n n e lla c a s te lla n ii, 323 Harvesting, 112, 255— 260 centrifugation, 255— 256, 336— 337, 485 chemical flocculation, 337 coagulation, 336 efficiency, 316 filtration, 256, 317, 336

flocculation, see also Flocculation, 257— 260 flotation, see Floating, 485 improved methods in future, 486 mass cultures, 95 methods, 336—337 microalgae, 29 microstraining, 336— 337 regime, 316 S c e n e d e s m u s , 235 screening, 256— 257 sedimentation, 256 S p ir u lin a , 223— 224 straining, 256— 257 vibrating screens, 337 Hatch-Slack Pathway, 55— 56 HCN, 167 H C 03, 313 H C < V , 85, 158, 304 H2C 0 3, 158 Health food, 30, 425, 485, 487 C h lo r e lla factories, 212 S p ir u lin a , 224, 228— 229 Heart disease, 230, 425— 426 Heat, 95 Heat of combustion, algae, 310 Heating cultures, 327 Heavily polluted estuaries, 101 Heavy metals, 95, 184, 228, 373— 376 Helical coiling, S p ir u lin a , 214 Helical form, 214 Helical shape of trichome, 213 Helicity, 214 Helicoidal shape, 217 Helicoidal trichomes, 213 H e lio th is

401 401 Helix dimensions, 213 Hemacytometer method, 75 Heme, 167 Heme biosynthetic pathway, 65 Hemicellulose, 59 H e m is e lm is b r u n e sc e n s , 427 H e p a lo s ip h o n , 166 Herbivores, 102, 109 Heterocysts, 56, 168, 170, 173 nitrogen-fixing, 166— 167, 455— 457 suppression of, 179 Heterotrophic algae, 181 C h lo r e lla , 212 high rate oxidation pond, 331— 332 Heterotrophic growth, 31, 181 C h lo r e lla , 210 S c e n e d e s m u s , 236 Heterotrophic metabolism, 162 Heterotrophic nutrition, 331 Heterotrophy, 147— 148 Hexadecanoic acid, 237 Hg, see Mercury High blood pressure, 424 High cell density, 209 v ir e s c e n s ,

zea,

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High energy state of electron, 48 Higher light gradients, 289 Highest output rate per unit area, 222 Highest production per area, 292 High filament densities, S p ir u lin a , 214 High irradiance, protecting effect of {3-carotene, 234 High light intensity, see Light intensity High light saturation constant, 312 High lutein content, 66 High nutrient concentration, 214 High output, S p ir u lin a mass culture, 222 High-productivity agriculture, 11 High radiation intensity, 223 High-rate algal pond (NRAP), 29, 35— 36 High rate oxidation ponds (HROP), 331— 338 High rate ponds, 268, 273, 276 High salinity, 94 High salt algae, 90 High salt concentration, 90 High salt stress, 90 High solar irradiance, 310 High-temperature algae, 206 High temperature-dependent respiration rate, 88 High temperature strains, 33, 81— 82 Hill reaction, 48— 49 Hill reagent, 48 Histidine, 165, 227 H20 , 48, 50 H 20-dehydrogenase, 50 Hog manure after fermentation, industrial produc­ tion of C h lo r e lla , 212 Holochrome, 65 Homocysteine, 180 Homogenization, 224 Horizontal ponds, 245— 247 H o r m id iu m sp., 342, 351 Hormogonia, 216— 217 Host, growth parameters of, 326 Host-parasite interrelationships, 322 Host-parasite pairs, 321 Host-parasite relationships, 321 Host specificity, 320 H2P 0 4- + H P 042“ , 177 HRAP, see High-rate algal pond HROP, see High rate oxidation ponds Human consumption, soybean harvest, 7 Human enzyme reactivation, S p ir u lin a , 230 Human food, see Food; Food for humans Human studies, nutritional, 386— 388 Hunger problem, 5 Hydraulic farming, 282 Hydrocarbon accumulation, B o ty r o c o c c u s b r a u n ii, 239 Hydrocarbon plants, 19— 20 Hydrocarbons, 346— 347, 421, 4 3 0 -^ 3 2 H y d r o d ic ty o n r e tic u la tu m , 439 Hydrogen, 157— 158, 438—439 Hydrogen-accepting molecule, 48 Hydrogen acceptor, 48 Hydrogenase, 157— 158, 201, 204, 438 Hydrogen ion concentration, 85— 86

Hydrogen peroxide, 299 Hydrogen sulfide, 109 Hydromechanics, 111 Hydrophobic bonding, 62 Hydroxyl ions, 85 Hydroxyl radical, 299 Hyperlipidemia treatment, 425 Hypertonic conditions, 231— 232 Hypertonic shocks, 90 Hypertrophic aquatic ecosystems, 101 Hypertrophic pond, 106 Hyperuricemia, 371 H y p n e a , 441 Hypocholesterolemia, 406—407 Hypotonic conditions, 231— 232 Hypoxanthine, 164

I Ik, 310 Illuminated surface, 286 Illuminated upper layer. 289 Illuminating power, 45 Illumination of culture, 45— 46, 125— 126, 290 artificial, 29 frequency of, 288 Imitation chicken flavor, 425 Immune system, 229 Implementation policies, 16 Improved Neaubouer, 121 Incandescent light, 46 Incentives, 22 Incident irradiance, seasonal variations, 293— 294 Incident light, 45, 71, 285, 291 Incident radiation, 312 Inclusions crystalline, 62 prokaryotes, 56— 57 Incubation temperature, growth rate, effect on, 78 India microalgae, consumption of, 28 microalgae production, 30 reforestation, 17 Indifference zone, 73 Induction of synchrony, 118 Industrial endeavor, 485 Industrial production, see Commercial production Infant feed, 391 Infections, 320, 322, 326— 327 Infrared light, 46 Infrared radiation, see IR Infrastructure, 21 Inhibition of growth, 220 Inhibitory action, 86 Initial response to salinization, 94 Initial slope, 86 , 88 , 104, 310 Injector, 252 Inner membrane, 61 Innoculation rate, 240 Inoculum, C h lo r e lla production, 212

507 Inorganic carbon, 76, 158— 160 Inorganic nitrogen compounds, 166— 171 Inorganic nitrogen uptake, 176 Inorganic phosphates, 177 Inorganic sulfate, 180 Inositol, 227, 349 Insect feeding, 401— 402 Insoluble molecules, transport of, 61 Integral irradiance, D u n a lie lla , 234 Intensity of stirring, 289 Intensive respiration, 80 Intermittent illumination, 203, 288 Intermittent light, 209 Internal transport system, 61 Intracellular glyerol, 92 Intracellular pigment content, 69 Intracellular reserve pools, 108 Intracellular transport, 61 Intragastric agent, 425 Invading organisms, 314 Invertebrates, control of, outdoor mass cultures, 322— 327 Inverted microscope, cell counting, 120 Investments, mass culture of microalgae, 32 Iodine, 27, 186, 229 Ionic bridges, 238 Ionic composition, 92 Ionic compounds, culture medium, 126 Ionic regulation, response to salinity by, 92 IR, 45—47, 73 Irish moss, 26 Iron, 47, 181, 183, 188 autotrophic C h lo r e lla biomass, 209 deficiencies, 6 6 , 183 S p ir u lin a , 226— 227 Iron protein, 167 Iron starvation, 183 Iron-sulfur protein, 50, 183 Irradiance, see also Solar irradiance, 29, 111, 309 correlation with temperature, 296 growth limitation, 310 insufficient, 223 interrelation with maximum daily temperature and photosynthetic activity, 297 peak of, 296 relationship to day length, 88 S p ir u lin a , 219 variations in, 292 Irradiant flux, 223 I s o c h r y s is sp., 35, 423 Isocitric dehydrogenase, 95 Isogametes, union of, 231 Isolation algae, 141— 142 thermophilic algae, 206 microalgae, 123— 124 Isoleucine, 227 Israel albazod processing, 36 commercial production of S p ir u lin a for food, 226 microalgae study, 29

Italy algological work, 33 microalgae production research, 31

j Japan commercial production of S p ir u lin a for food, 226 iodine extraction, 27 mass culture systems, 30 microalgae production, 29 seaweeds harvested, 26

K K, see Potassium K \ 93, 178 Kanembu, 221 Kelp, 27 Kelzan, 442 Keratinization, S p ir u lin a , 2 2 9 Kinesis, 73 K ir c h n e r ie lla lu n a ris, 438 K le b s o r m id iu m m a rin u m , 440 Klett-Summersen colorimeter, 122 K/Na ratio, 182 Korea, microalgal biomass for sale, 34 Ks value, 154

L Labor, induction of, 424 Laboratory techniques of microalgae cultivation, 117— 145 addresses of culture collections, 143 algal growth media, see also specific types, 125— 134 automatic synchronization, 135 balanced growth, 117 biomass concentration, procedures for determina­ tion of, 120— 123 cell counts, see also Cell counts, 120— 121 cell growth, procedures for determination of, 120— 123 chemostat, 137— 139 chlorophyll determination, 122— 123 collection of microalgae, 123 culture media, 126 dry weight measuring, 122 estimation of biomass and growth, 142 fluorescence microscopy, 139— 140 frequency of transfer, 126 growth, 125— 134 illumination, 125— 126 isolation of algae, 141— 142 isolation of microalgae, see also Isolation, 123— 124 light measurement in algal suspension, 137— 138

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CRC Handbook of Microalgal Mass Culture

light scattering, 121— 122 liquid culture, 117— 120 maintenance of microalgae, 125— 134 mass cultivation, 135 microalgae cultivation, 117— 145 microscopical methods, 139— 141 oxygen electrode, 138— 139 preservation of microalgae, 125— 134 purification of microalgae, see also Purification of microalgae, 125 separation of algae, 141— 142 special devices, 135— 139 special procedures, 139— 142 specific growth rate, 117— 118 sterilization, 125 sunlight intensity, stimulation of variability in, 136— 137 temperature, 126 thermal gradient device, 135— 136 turbidostat, 137— 139 unbalanced growth, 118 Labor-oriented agriculture, 11 Lactation, 381, 383 Lactic acid fermentation, 204 Lactic bacteria, 212 Lag phase, 118 Lake Aranguadi, East Africa, 112 Lake Biwa, Japan, 109 Lake Chad, Central Africa, 213, 221, 339, 360 recipe for Dihe, 228— 229 Lake Erie, United States, 109 Lake Kinneret, Israel, 102— 103, 108 Lake Tchad, 28 Lake Trummen, 1 11 Lake Washington, United States, 109, 111 Lake Zurich, Switzerland, 109 Lambs, feed for, 400—401 Laminaran, 63 L a m in a r ia , 27 ja p o n ic a , 26 Land use patterns, 11— 12 Langmuir cells, 101, 103 Large-scale culture, S p ir u lin a , 222 Large-scale operations, 487 Large-scale production, 486 Laser leveling, 266 L a u d e r ia b o r e a lis , 427 Laurie acid, 227 Laver, 26— 27 Laying hen feed, 395— 396 Layout of pond facilities, 267— 268 LD100, 327 Lead, 181, 186, 228 Lead hydroxide stain, 57 Legumes, 365— 370 Length, 213 Leprosy patients, 29 L e p to c y lin d r u s d a n ic u s , 88 Leucine, 165, 227 Leucosin, 63 Life cycle, S p ir u lin a , 216— 217

Life histories, 111 Light, see also specific topics; specific types, 69— 77, 304, 322 absorption, 66 algal movement, effect on, 72—74 algal production in mass cultures, 310 attenuation coefficient, 106 buoyancy, effect on, 74— 77 cell length, 95— 96 cell morphology, 95—96 cells, Chlorella, 208—209 continuous, 288 daily quantity, 295 dependence of growth on, 285 distribution, mixing, effect of, 289— 292 Dunaliella, 231 growth, Chlorella, 202— 203 high rate oxidation pond, 331— 332, 335— 336 inhibition, 106 inorganic polyphosphate synthesis, 180 intensity, 46, 55, 87, 293, 301, 310—311, 314 availability to each cell, 291 average, 291 below saturation, 78 buoyancy, 75 cell division, 208 changes in, 71 dark cells, 209 different temperatures, 294 Dunaliella, 231 elevation, 79 functions, 288 growth curve, 286— 287 half-maximal growth, 310 high, 58, 6 6 , 69, 90, 94, 211, 214, 231, 285, 288, 300 increasing, 71, 88 , 233, 293 light cells, 209 low, 231, 285 0 2 concentration and temperature, 297 population density in relation to, 285— 292 Porphyridium, 238— 239 saturating, 207, 286— 287 Scenedesmus, 236 sudden changes, 73 intermittence of, 288 light-shade adaptation, 69— 72 limitation, 222, 295 loss of C 0 2, 304 manganese deficiency, 184 measurement in algal suspension, 137— 138 nitrite reducing steps, 174 organelles influenced, 214 penetration of, 101 phosphorus uptake, 178 photoinactivation, 69 photoinhibition, 69— 70 photo-phobotactic reactions, 73 photosynthesis dependence on, 104 photo-topotactic reactions, 73— 74 regimes, 70, 71, 77

509 requirement, 306 saturation, 86 , 105 Chlorella pyrenoidosa, 207 curve, 86 , 88 photosynthesis rate, 81 seasonal variations, 293— 294 selective factor, 317 sensitization, 350 sole limiting factor, 310 temperature interaction with, 86— 89 total amount, 288 transmission efficiency, 310 utilization of, 112, 289, 310 wavelength, see Wavelength of light Light-dark (LD) cycles, 70—72, 252, 288, 292 Light-dark pattern, 289 Light-dark regime, 310 Light-dependent oxygen consumption, 301 Light dissipation efficiency, 310 Light energy (E \), 44, 50 antennas for capture of, 58, 64 conversion into chemical energy, 48 utilization for carbohydrate synthesis, 52 utilization in Hill reaction, 48—49 Light energy conversion efficiency, Porphyridium, 238 Light flux, 45—46, 286 Light harvesting efficiency, 71 Light-harvesting pigment molecules, 48 Light-harvesting pigments, 49 Light-induced electron flow, 51 Light-limited chemostat cultures, 307 Light-limited cultures, 206 Light-limited growth, 285— 289, 295 Light microscope, 56 Light quanta, 49— 50, 52 Light radiation, 43 Light reactions of photosynthesis, 50 Light saturated rate of photosynthesis, 70 Light saturating intensities, 79 Light saturating points, 81 Light saturating values, 81 Light scattering, 87, 121— 122, 302, 313— 314 Light-shade adaptation, 69—72, 105 Light spectra, 43—47 Light to dark ratio, 288 Light trap, 73— 74 Limitations to production, 292 Limiting growth conditions, Botyrococcus braunii, 239 Limiting nutrient, 107, 109 Linablue, 435 Linear phase, 285— 286 Linings for ponds, see Pond construction and design Linoleic acid, 82, 424, 485 prostaglandin PGE,, 230 Spirulina, 227— 228 a-Linoleic acid, 82 Linolenic acid, 237 a-Linolenic acid, 221, 346 y-Linolenic acid, 346, 424, 426, 445

deficiency, 230 Spirulina, 226— 228 Lipids, 29, 188, 342, 345— 346, 421—432 accumulation, 82, 211 biosynthesis, 211 chlorosulfolipids, 422 Chlorella, 210—211 droplets, 202 environmental factors, 422—423 essential fatty acids, 424— 425 fatty acids, 422—427 fraction, 237 free fatty acids, 421 globule, 58 glycerol, 421 glycolipids, 422 hydrocarbons, 430— 432 neutral, 211 nitrogen starvation, effect of, 423 nitrogen sufficiency, effect of, 423 nonpolar, 421 Phaedoctylum, 236— 238 phase change in, 84 phospholipids, 422, 424 polar, 211, 421—422 prostaglandins, 424— 425 sterols, 426, 430—431 storage, 344 synthesis, 55, 82, 211 temperature dependence, 78 triglycerides, 424 unsaponifiable matter, 346— 347 wax esters, 421, 426, 429—430 yield per unit of illuminated area, 211 Lipid-rich algae, 423— 424 Lipoic acid, 180 Lipoid structures, 421 Lipopolysaccharide, 349 Lipoproteins, 61 Liquid culture, see also Continuous culture, 117— 120 Lithium, 181, 186 Lithotrophs, 147 Lithotrophy, 147— 148 Liver enzyme regeneration, 364— 365 Liver tumor cells, 229 Lobose amoebae, 323 Locomotory structures, eukaryotes, 61— 62 Logarithmic growth factors, 325 Logarithmic growth phase, 87 Logarithmic phase, 117 Lomofungungin, 141 Long Island Sound, 108 Longitudinal division of cells, 230 Long-term feeding experiments, 381, 383 Loroxanthin, 350 Low carbohydrate content, Chlorella, 211 Low cell density, 209 Low chlorophyll-containing cells, Dunaliella, 234 Lower unilluminated strata, 289 Low growth rate, 309

CRC Handbook of Microalgal Mass Culture

510 Low Low Low Low

night temperature, S p ir u lin a , 219 nutrient concentration, 214 rate of decay processes, 312 temperature, 83, 85, 206, 214 L o x o d e s m a g n u s, 323 LPP 6402, 165 LPP73110, 165 Luciferase, 436— 437 Luciferin, 436—437 Lumens, 46 Luminescence, 46 Lutein, 350, 432-^133 Lux, 46 Lycopene, 66 Lymphocytes, 229 L yngbya g r a c ilis ,

444 426, 428, 444

m a ju sc u la ,

L y n g b y a -P le c to n e m a P h o r m id iu m

(L.P.P.), 71, 166,

320, 456 Lysine, 227 Lysing fresh water algae, 322 Lysis, 216

M Macroalgae, 200 27 p y r if e r a , 25 Macroeconomic considerations, 473 Macromolecular aggregates, 58 Macronutrient elements, see also specific topics, 156— 188 ammonium assimilation, 172 calcium, 157, 181— 182 carbon, 157— 163 chlorine, 182 defined, 157 hydrogen, 157— 158 magnesium, 157, 182 nitrogen, 157, 163— 177 oxygen, 157 phosphorus, 157, 177— 180 potassium, 157, 182 sodium, 181— 182 sulfur, 157, 180— 181 Magnesium, 157, 181— 182, 188 concentration, 322 deficiencies, 66 S p ir u lin a , 227 Maillard reaction, 343 Maintenance coefficients, 210 Maintenance energy, 307, 315 Maintenance of microalgae, see also Laboratory techniques of microalgae cultivation, 125— 134 Maintenance of outdoor mass culture, 223 Malate, 162 Malaysia, microalgal biomass for sale, 34 Maltose, 163, 236, 349— 350

M a c r o c y s tis ,

Management aspects, 473—487 Managerial factors, 474 Manganese, 47, 181, 184— 185, 227 Manic depression, 230 Mannans, 61, 199 Mannitol, 440 Mannose, 202, 236 Manure, 30 Marginal areas, 15 Marine coccolithophorids, 61 Marine habitats, 240 Marine water, P o r p h y r id iu m , 238 Market considerations, 477 Market potential for algal products, 485 Mass algal cultures control of infection, 326 photooxidative death, 302 Mass cultivation, see also specific topics autotrophic alga, 245— 249 B o ty r o c o c c u s b r a u n ii, 239 C 0 2 supply, utilization and distribution, 254— 255 dehydration of algal mass, see also Dehydration of algal mass, 260— 262 efficiency of photosynthesis, 251 harvesting the algae, see also Harvesting, 255— 260 microalgae, 135 mixing, 250— 253 pond lining, 250 ponds, 245— 247 reactors for autotrophic alga, 245— 249 shading, 250 sloped cultivation units, 247— 249 solar radiation, 250 technological aspects, 245— 263 turbulence, see also Turbulence, 250— 253 Mass culture, 30 B o ty r o c o c c u s b r a u n ii, 239 buoyancy, 75 cell density, 209 chilling temperature, 84 hydrogen ion concentration, 85 microalgae, 28 morphological modifications, 95 natural conditions, 101— 104 O s c illa to r ia , 216 P h a e o d a c ty lu m , 238 S c e n e d e s m u s , 235— 236 S p ir u lin a , 216, 222— 224 M a s tig o c la d u s , 166, 456 la m in o su s, 437, 439 Mastigonemes, 61 Mathematical formulation of optimization problem, 481—482 Matrix, 62—63 Mature plastids, 63 M a u g e o tia , 141 Maximal day temperature, S p ir u lin a , 219 Maximal photosynthetic efficiencies, 307 Maximal photosynthetic rate, 105

511 Maximum conversion of total solar energy to bio­ mass free energy, 307 Maximum daily temperature, 296, 297 Maximum efficiency, 310 Maximum light utilization efficiency, 310 Maximum output, 302 Maximum output rate, 315 Maximum photon flux, 46 Maximum photosynthesis, 315 Maximum photosynthetic rate (Pmax), 88 Maximum possible growth rate, 310 Maximum temperature, 302 Maximum yield potential, 307 Meander ponds, 245 Meat, 19 Mechanical disintegration, 353 Mechanical power phase, 12 Mechanical stimuli, algal movement, 73 Medical uses, seaweed, 27 Medium composition, S c e n e d e s m u s , 234 Meiosis at zygote germination, 230 M e lo s ir a

172 sp., 439 Membrane, 63, 84, 91 hydrogen ion concentration, 85 permeability, 89—90 Membrane-bound nucleus, 60 Membrane-bound plug, 61 Mercaptothione, 327 Mercury, 181, 186, 228 Mesh screens, 327 Mesohaline lakes, 217 Mesophilic algae, 78, 206 Metabolic regulation, disruption of, 80 Metabolic studies, 356— 362 Metachromatin, 57 Metaphosphates, 178 Methane, 109, 439— 440 Methionine, 165, 180, 227 L-Methionine, 180 L-Methionine DL-sulfoximine (MSO, MSX), 171— 172 Methylotrophy, 148 Methyl viologen (MV), 168 Mexico commercial production of S p ir u lin a for food, 226 microalgae, consumption of, 28 Mg, see Magnesium Mg2+, 167, 178, 222 Mice mutagenicity study, 382 toxicological studies, 384— 385 M ic o r a c tin iu m sp., 439 M ic r a c tin iu m , 306, 357, 375, 377, 383 M ic r a s te r ia s d e n tic u la ta , 73 Microbial contamination, 379— 380 Microbiological analysis, S p ir u lin a , 228 M ic r o c o le u s sp., 437, 456 M ic r o c y s tis , 78, 86 , 372 a e r u g in o s a , 75— 77, 86— 87, 372, 429 n u m m u lo id e s,

in c e r ta , 79 Microeconomic considerations, 473 Microfibrils, 348 Micron, 45 Micronutrients, 183— 188 Micropipette, washing by, 123 Microscopical methods, 139— 141 Microscopic evaluation, 313 Microscopic examination of culture, 314 Microstrainers, 256— 257, 336— 337 Milk, 19 Millimicron, 45 Mineral composition, 348 Mineral nutrients C h lo r e lla , 210 deficiency, 312 Mineral nutrition, 55 Mineral precipitation, 312 Minimal temperature, S p ir u lin a , 219 Minimum quantum, 306 Mitochondria, 50— 51, 56, 60 electron transport system, 178 eukaryotes, 59, 61 Mitochondrial respiration, 48 Mitochondrial respiratory chain, 50 Mixed populations of microalgae and bacteria, 285 Mixing of cultures, 250— 253 inadequacy, 312 light distribution, effect on, 289— 292 net photosynthetic reaction rate, 290 outdoor mass cultures, 285— 292 rate of, 292 Mixing depth, 107 Mixing energy, 314— 315 Mixing velocity, 268, 272, 277 Mixotrophic competitors, S p ir u lin a , 224 Mixotrophy, 147— 148 Modem agriculture, 11 M o in a sp., 379 Moisture, S p ir u lin a , 227 Molasses, 236, 380 Molecular oxygen, 49, 299 Molybdenum, 167— 168, 181, 185— 188 Molybdenum-iron protein, 167 M o n a lla n tu s s a lin a , 440 Mongolia, microalgae, consumption of, 28 Monoalgal cultures, 84, 223, 285, 316— 320, 326 M o n o c h r y s is lu th eri, 83, 440 Monod equation, 107— 108, 153— 154 Monod model, see Monod equation M o n o d u s , 165 s u b te r r a n e o u s , 148, 425, 430 Morphological changes, interactions of environmen­ tal factors, 95— 96 Morphological features, algae, 199 Morphological modifications, 95 Morphological transformations, 214 Morphological types, S p ir u lin a , 213 Morphological variability, 96 Morphological variants, S p ir u lin a fu s if o r m is , 214 Morphology of cell algae, 95—97

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200— 203 212— 216 Mother-cell wall, fate of, C h lo r e lla , 201— 202 Motile cells, 230 Motility, 199 Motor responses, 73 M o u g e o tia , 141 Movement, see also Algal movement, 425 Mucilage, layer of, 56, 59 Mucilaginous polysaccharide, P o r p h y r id iu m , 238 Mucopolysaccharides, 61 Multicellular algae, 61 Multigenerational study, 384 Multiple sclerosis, 426 Muramic acid, 228 Mutagenicity, 381— 382 Mutual shading, 77, 286, 288, 292, 295, 315, 317 Mycotoxins, 372— 373 M y c r o c y s tis , 74 Myristic acid, 227 Myxobacteria, 322 Myxol-glycoside, 350 M y x o s a rc in a , 166, 456 Myxoxanthin, 66 Myxoxanthophyll, 6 6 , 350, 433 C h lo r e lla , S p ir u lin a ,

N N3", 167 N -8 medium, 129 N a+, 93, 178 NAD, 158 NAD+, 171 NADH, 171 NADH to water, 48 NADP, 158 NADP+, 48— 51 NADP+-NADPH redox couple, 48 NADPH, 49— 51, 53— 54, 167— 168, 174 N a+-K+ balance, 94 Na/K exchange, 92 N a n n o c h lo ris , 35, 423 N a n o c h lo r o p s is s a lin a , 423 Nanometer, 43, 45 NaOH, 224 Narrow transparent tubes, 285 N a ss u la o r n a ta , 323, 324 Natural conditions, 101— 115 Natural protoplast, 230 N a v ic u la p e llic u lo s a

lipid content, 423 nickel, 186 photorespiration and photosynthesis, 305 photosynthetic pigment, 188 silicon, 187, 188 sterols, 426, 430 r a d io s a , 85 Necridia, 216— 217 N e o c h lo r is o le o a b u n d a n s , 424

Neomycin, 125 Neon light, 46 Neoxanthin, 350 Nephropathy, 370 Net C 0 2 assimilation, 304 Net growth rate, 309 Net photosynthesis, 304 Net photosynthetic reaction rate, 290 Net Protein Ratio (NPR), 360— 361 Net protein utilization (NPU), 358— 362, 367, 369— 370 S p ir u lin a , 227 Net yield, S p ir u lin a , 225 NH3, 304, 312 NH4+, 167, 169— 170, 176 Niacin (B3), 225 Nickel, 181, 185— 187 Nicotinate, 349 Nicotinic acid, 227 Night temperature, 205, 207— 209 N ite lla h o o k e r i, 444 Nitrate (N O ,-), 126, 164— 165, 173, 180, 189 assimilation, 167— 168, 220 C h lo r e lla , 203, 210 deficiency, 94 S p ir u lin a , 222, 224 uptake, 173, 175 Nitrate N, 168 Nitrate oxidoreductase (NR), 167 Nitrate reductase, 173, 183, 185, 460 Nitrate reductase:NAD(P)H:nitrate oxidoreductase (NR), 167 Nitrate reduction, C h lo r e lla strains, 204 Nitrite (N 02"), 164 assimilation, 167— 168 Nitrite oxidoreductase (NiR), 167— 168 Nitrite reductase, 173, 183 Nitrite reductase:NAD(P)H:nitrite oxidoreductase (NiR), 167 Nitrogen, 47, 157, 163— 177, 183, 298, 304, 486 ammonia assimilation, 168— 171 analysis, 312 assimilation, 166— 174, 181, 183 atmosphere, 302 autotrophic C h lo r e lla biomass, 209 balance, 359, 362, 388 buoyancy, 77 C h lo r e lla yields, 210 comparison of algal growth rates, 164 culture medium, 126 deficiency, 6 6 , 169, 173— 174, 179, 188, 201, 422— 423 depletion, 313 distribution of ability of algae to utilize, 165 economy, 304 excess, 311 fixation, 110— 111, 157, 164, 166— 167, 176, 185 C h lo r e lla , 210 cyanobacteria, 455—457 physiology of, 455— 457

513 salinity, effect of, 94— 95 symbioses involving blue-green algae, 464— 465 inorganic compounds, 169 interaction with carbon metabolism, 174— 177 limitation, Phaeodactylum tricornutum, 236 loss, 312 metabolism, 180 nitrate assimilation, 167— 168 nitrite assimilation, 167— 168 organic compounds, 169 Spirulina, 227 starvation, 173— 175, 345 Chlorella, 211 lipid content, effect on, 423 sufficiency, lipid content, effect on, 423 Nitrogenase, 166— 167, 173, 176, 438, 456—457, 460, 464 inhibition of, 168 N2-fixing cyanobacteria, 179 Nitrogen compounds, Chlorella utilization of, 203 Nitrogen-deficient organisms, 163 Nitrogen-depleted cells, 211 Nitrogen-fixing cyanobacteria, 95, 109, 455—457 Nitrogen-fixing microorganisms, 486 Nitrogen-starved algae, 169 Nitrogenous storage material, 220 Nitzschia alba, 187 closterium, 346 Nitzschia ovalis, 172 palea, 423

sp., 84, 165 N20 , 167 Noctiluca, 436 Nonaggregated phycobiliproteins, 62 Nonbacterial contamination, 377—379 Nonessential amino acids, 227 Nonionized (free) ammonia concentration, 327 Nonpolar lipids, 421 Nonprotein nitrogen (NPN), 341—342 Normandy, seaweed industry, 27 North America, grain trade, 11 Nostoc commune, 28, 347, 430 edule, 28 muscorum, 95, 97

cobalt, 186 growth rate, 164 hydrogenase activity, 438 lipids, 429 sp., 28 lipids, 429 nitrogen-fixing, 166— 167, 456 N/P ratio, 180 Nuclear envelope, see also Envelope, 61 Nuclear pores, see also Pores, 61 Nucleic acid, 56, 177, 188, 199, 342, 346—347, 370—372 acceptable intake, 371 uric acid concentration and, 371

Spirulina, 227 Nucleoplasm, 56, 61 Nucleus, 59— 61 Nutrients, see also Nutrition, 86, 286, 314, 317 algal growth, 326 availability, 76, 77 buoyancy, effect on, 74— 77 composition, 111 concentrations, optimal range of, 148— 150 deficiency, 76, 188, 233 elements, variations in supply of, 47 flux, 223 limitation, 297 Nutrition, 147— 198 autotrophy, 147— 148 auxotrophy, 147— 148 growth regulators, 189— 190 heterotrophy, 147— 148 high rate oxidation pond, 335 limitations, 150— 155 list of nutrients needed by algal cells, 149 lithotrophy, 147— 148 macronutrient elements, see also Macronutrient elements, 156— 188 micronutrients, see also Micronutrients, 183— 188 mixotrophy, 147— 148 models for double-substrate limited growth, 155— 156 modes of, 147— 148 optimal range of nutrient concentrations, 148— 150 paratrophy, 148 phagotrophy, 148 photoheterotrophy, 147— 148 plant hormones, 189— 190 saprotrophy, 148 Scenedesmus, 236 Spirulina, 220— 221 vitamins, 188— 189 Nutritional limitation, 79, 150— 155, 312 Nutritional properties of microalgae, see also spe­ cific topics, 339— 419 acceptability studies, 390— 392 animal feed, 392— 405 animal feeding tests, 354— 356 biological contaminations, 377— 381 chemical composition, 341— 350 digestibility, 350— 354 environmental contaminants, 373— 377 metabolic studies, 356— 362 nutritional studies with humans, 386— 388 processing of algal biomass, 350— 354 protein regeneration studies, 362— 365 studies, 353— 356 supplementation studies, 365— 370 therapeutic properties, 406— 408 toxicological aspects, 370— 373 toxicological studies with animals, 381— 386 Nutritional requirements, 78 Scenedesmus, 236

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514

Nutritional status, outdoor mass culture ponds, 312—313 Nutritional studies with humans, 386—388 Nutritional value of algae, 33, 227, 361, 485

o Obesity, 230 Obligate autotrophy, 162 Obligate heterotrophy, 162 Obligate photoautotrophy, 220 Obligate phototrophs, 148 Ocean biological productivity, 47 Dunaliella , 230 photosynthetic production, 47 production of organic matter in, 47 Ochromonas, 65, 130, 165 danica, 148, 422 antibiotic activity, 443 essential fatty acids, 425 fatty acids, 427 lipid content, 423 sterols, 426, 430 malhamensis, 440 variabilis, 148 Oedogonium, 28, 141 Office of Technology Assessment (OTA), 17 OH~, 304 Oil/energy problem, 4 Oil palm, 19 Oils, 47, 163, 199 Oleic acid, 82, 227 Oligotrophic water bodies, 108— 109 Olisthodiscus luteus, 120 Olive oil, 19 Oocystis, 354, 357, 360, 361 polymorpha, 423 Open pond with agitation, Chlorella production, 212

Open pond with circulation, Chlorella production, 212 Open semi-defined culture systems, 311 Open system, see also Continuous culture, 119— 120

Operating costs, 475—477 Operating culture depth, 270 Optical density of suspension, 78 Optimal decision rules, 474— 478 Optimal density, 222, 315 Optimal intermittence, 291 Optimal temperature, 78, 88, 200, 287 Porphyridium, 238 Scenedesmus, 236 Spirulina, 219 Optimization problem, 481—482 Optimizing growth conditions outdoors, 203—205 Optimum cell density, 290, 316 Optimum conditions for algal growth, 326 Optimum growth

phosphorus requirements, 177 Porphyridium, 239

Optimum oxygen concentration, 322 Optimum population density, 292 outdoor mass cultures, 315—316 Spirulina, 222 Oral toxicity, 385 Organelle ribosomes, 65 Organelles, 214 Organic acid, 327 Organic by-products, 220 Organic carbon, 160— 163, 176, 203, 304 Organic compounds, 376—377 Organic excretions, 310 Organic load, 322, 335—336 Organic matter, 47 Organic nitrogen compounds, 169, 171— 174, 176 Organic osmotica, 90—94 Organic phosphates, 177 Organic sulfur, 180 Organic toxins, 377 Organotrophy, 147 Ornithine, 165 Orthocid 50, 326 Orthophosphates, 178 Oscillatoria agardhii, 323—324 limnetica, 181, 438 nigrovidis, 444 platensis, 215 redekei, 70—72 rubescens, 77

sp., 77, 84 cultivation device for, 137 dominance, 317 mass culture, 216 methane production, 439 nickel, 187 nitrogen-fixing, 166, 456 screening, 256 tenuis, 428 Oscillatoriaceae, 214 Oscillaxanthin, 350 Oscillol-glycoside, 350 Osmoregulatory mechanism, Dunaliella, 231 Osmoregulation, 89—95, 440 Dunaliella, 231—233 halotolerant wall-less algae, 232 Osmoregulators, 440—441 Osmoregulatory compound, 94 Osmoregulatory response, 90 Osmotica, 94 Osmotic changes, 124 Osmotic compensation, 92 Osmotic equilibration, 94 Osmotic metabolites, accumulation in halotolerant algae, 440 Osmotic potentials, 92 Osmotic pressure, Dunaliella, 232 Osmotic regulation, 93, 182 Osmotic shock, 62

515 Osmoticum, 90 Ourococcus, 423 Outdoor cultures, 219, 233 Outdoor mass cultures, 285—329 algal concentration, 288 algal production in, 310 biological principles, see also other subtopics hereunder, 285—312 clean, 285 dissolved oxygen concentration, 298 evaluation of culture performance, 312—327 interrelations between solar irradiance, population density, and mixing, 285—292 interrelations between solar irradiance and tem­ perature, 292—297 light distribution, effect of mixing on, 289—292 light-limited growth, 285—289 maintenance, 223 monoalgal cultures, 285, 316—320 open semi-defined systems, 311 optimum population density, 315—316 oxygen inhibition, 303—304 oxygen toxicity, 299—303 pest control, see also Pest control, 320—327 photooxidation, 299—303 photorespiration, 304— 306 pond maintenance, 312—315 potential of, 310 production systems, 285 productivity of, 306—312 Spirulina, 222

supersaturation with oxygen, 298—299 wastewater systems, 285 world programs, 306 Outdoor pond, 81, 213 Outdoor temperature, Chlorella, 206 Outer wall, 215 Output rate, 290, 295 population density in relation to, 315 Scenedesmus, 235 specific growth rate in relation to, 315 Spirulina, 224 Oval cells, 236 Overall decay factor, 310 Overproduction, 4 Overturn, 109 Ovoid shape, 230 Oxidases, 299 Oxidation lagoons, 101 Oxidation pond, Spirulina, 213 Oxidative phosphorylation, 50—51, 56, 178 a-Oxoglutaric acid, 170 Oxygen, 48, 157— 158, 163 activity, 298 atmosphere, 302 competition with C 02, 304 concentration, 223, 295—297, 301—305, 327 content, 298 decline, 223 evolution, 48—49, 85—86, 219, 296 gas phase, 49

high rate oxidation pond, 336 inhibition, outdoor mass cultures, 303— 304 lethal effect of exposure to light in, 299 molecular, 49 outdoor mass culture ponds, 314— 315 partial pressure, effect on Chlorella yield, 304 polyphosphate formation, 180 production, 81 reduction, 301 replenishment in atmosphere, 47 respiratory consumption, 56 solubility, 298 supersaturation, 298— 301, 314 tension, 295, 303, 327 toxicity, outdoor mass cultures, 299— 303 Oxygen electrode, 138— 139, 314 Oxygen-generating photosynthesis, 56 Oxygen-sensitive C 0 2, 304 Oxy trophy, 148 Oysters, 402 Ozone ( 0 3), 46

P Pi, 177 P430, 50— 51 P680, 50 P700, 50—51, 70 Packed cell volume, 285 Paddle wheels, 31, 222, 225— 226, 235, 245, 251, 271 design and construction, 272— 276 drive unit, 272— 275 high rate oxidation pond, 335 main shaft, 272— 275 paddle design, 275— 276 paddle tip speed, 272 power requirements 274— 275 Palmer Malony, 121 Palmitic acid, 82, 227, 237 Palmitoleic acid, 82, 227 Palmitolinoleic acid, 227 Pandorina morum, 325 d-Ca-Pantothenate, 227, 349 Pantothenic acid, 180 Paper industry, 28 Paramylon, 63, 350 Parasite, 325, 326 Parasitic fungi, 334 Parasitism, 104, 111, 321 Paratrophy, 148 Parental walls, 200, 235 Parent cell, 235 Parkinson’s disease, 426 Parthenium argentatum, 20 Partial pressure of oxygen, 298, 304 Particle phenomenon, 43 Particulate carbon (PC), 83 Particulate nitrogen (PN), 83 Particulate organic carbon (POC), 311

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Pastalina, 228

Pasteurization, 224 Patches, 101 Pathogenic fungi, 322 Pathogenic viruses, 321 Paving materials, 278 Pavlova, 165 PCA pool, 85 Peak wavelengths, 49 Pellicle, 61 Pellicular wall, 56 Penicillin, 125 Penicillium, 379 Pennate diatom, 96 Pentose Phosphate Reduction Pathway, 54 Peptidoglycan, 349 Peptone, 163 Perception, treatment of disturbances of, 425 Perforated frustules, 61 Peridinin, 66, 432—433 Peridinium, 436 cinctum, 103— 104, 108, 188, 423 Perinuclear space, 61 Peripheral fibers, 61 Peripheral thylakoids, 62 Permeability Chlorella, 210 Dunaliella, 232 Peroxidases, 299 Peroxide radical, 301 Peroxisomes, 56 Peru microalgae, consumption of, 28 microalgal biomass for sale, 34 Pest control, 320—327 Pesticides, 326, 376—377 Petfish, 35 Petroff Houser, 121 PGA, 53—56 pH, 200, 300—301, 314, 317, 319, 327 algal growth, 326 alkaline, 304 ammonium, 164 Chlorella strains, 204 culture medium, 126 cyanobacteria, 320 diurnal fluctuations, 85 fluctuations, autotrophic Chlorella biomass, 209 gradient, 51—52 growth rate, 86—87 high, 113 high rate oxidation pond, 336 hydrogen ion concentration, 85 inorganic carbon, 158— 160 optimum, Spirulina, 220 outdoor mass culture ponds, 313 phosphorus uptake rate, 178 photosynthetic 0 2 evolution, effect on, 86 polyphosphate formation, 180 Porphyridium, 239

predators, 323

protozoa, 323—324 response to, 85 rise in, 313, 318 Spirulina, 217—218, 222 toxicity of algae, 86—87 Phacophyceae, 62 Phaeocystis poucht, 443 Phaeodactylum, 35, 131, 165, 187, 236—238 tricornutum, 83—84 dominance, 317 fatty acids, 427 growth regulation, 190 lipid content, 424 lipid productivity, 236 nickel, 187 nitrogen deficiency, 173 solar energy conversion efficiencies, 252 sterols, 430 Phaeophorbide-a, 350 Phaeophyceae, 63, 65—66, 189 Phaeophyta, 61, 63—64, 66, 148, 158 Phagotrophy, 148 Pharmaceuticals, 27—28, 229, 406, 443—445 Phase contrast, cross-walls, 214 Phenols, 443—444 Phenylalanine, 227 Philodina sp., 322, 327, 378 Phobic responses, 73 Phosphatases, 177 Phosphates, 57, 85 bonds, 47 metabolism, 180 uptake, 181, 188 Phosphoesterases, 177 3-Phosphoglycerate, 53 Phosphoglycerate kinase (PGK), 436 Phosphoglyceric acid, 160 Phosphoglycolate, 56 Phospholipids, 422, 424 Phosphorus, 47, 109, 157, 177— 180, 183, 222, 227, 311—313 Phosphorylation, 178, 181 Phosphoryled, 228 Photic zone, 289, 292 Photoassimilation, 301 Photoautotrophic biomass, 314 Photoautotrophic culture, 316 Photoautotrophic growth, 180 Photoautotrophs, Porphyridium, 238 Photobiological utilization of solar energy, 307 Photochemical reaction, 50, 62, 306 Photochemical systems, 46 Photoconversion, chlorophyll to pheophorbide, 35 Photodestruction, 211 Photodynamic death, 109, 111 Photoheterotrophy, 147— 148 Photoinactivation, 69 Photoinhibition, 69—70, 105, 223 Photolithotrophic heterotrophy, 147 Photometric measure of light, 45—46 Photon, 43, 45, 47, 52, 306

517 Photon flux, 46 Photooxidation, 75, 219, 223, 299— 303, 312, 314 Photoperiods, 70, 88 Photo-phobotactic reactions, 73 Photophosphorylation, 50— 52, 174— 175, 178, 180 Photoreceptor, 63, 65 Photorespiration, 55— 56, 189, 219, 304— 306 Photosensitive carotenoid-containing globules, 63 Photosensitization, 301 Photosensitized skin irritations, 350 Photosynthesis, 28— 29, 43— 56, 64— 67, 302 biomass production, 1, 3, 6— 7 buoyancy, 75 C4, 55— 56 carbohydrates, see also C 0 2 assimilation, 52— 55 carbon, 47, 167 Chlorella pyrenoidosa, 207 C 0 2 assimilation, see also C 0 2 assimilation, 52— 55 defined, 47 dependence on light, 104 depth integral of, 101, 107 efficiency of, 251 enhanced, 55 first act of, 64 free energy stored, 307 Hatch-Slack Pathway, 55— 56 high rate oxidation pond, light in, 331— 332 high temperature strains, 81 Hill reaction, 48—49 hydrogen ion concentration, 85 increasing temperature, 88 inhibition, 223 iron, 183 light available for, 292 light intermittence, 288 light-limited rates, 209 light reactions of, 50 light saturated rate, 70, 209 loss of C 0 2, 304 low temperature strains, 81 manganese deficiency, 184 maximal primary productivity, 112 nitrogen assimilating enzymes, 173 optimum temperature, 88 oxygen concentration, 157 oxygen-generating, 56 phosphate uptake, 180 photophosphorylation, 50— 52 photorespiration, 56 Photosystems I and II, 49— 50 process of, 47 relative rates in flashing light, 289 salinity, effect of, 94 saturating-light intensity, 88 severe inhibition, 221 stored reserve, 199 temperature, effect of, 305 transformation of dark cells into light cells, 208 triacontanol, effect of, 189 yield of, 47

Z-scheme of photosystems and electron transport in, 49 Photosynthesis-irradiance curves, 89, 104 Photosynthesizing plants, 64 Photosynthetic activity, 65, 94 buoyancy, 77 dark cells, 208 decrease in, 94, 303 dissolved oxygen concentration, 314 increased photorespiration, 302 intensive, 299 interrelated effects of solar irradiance and temper­ ature, 295 interrelation with maximum daily temperature and irradiance, 297 light cells, 208 NaCl concentrations, 94 pH, 313 protective mechanisms, 301 reduction in, 315 salt, effect of, 95 Spirulina, 217 stirring speed, 289 Photosynthetically active radiation (PAR), 307 Photosynthetic apparatus, 65 eukaryotes, see also Eukaryotes, 62—64 prokaryotes, 57—59 Photosynthetic assimilation of C 0 2, 43 Photosynthetic bacteria, 50 Photosynthetic capacity, 211 Photosynthetic Carbon Reduction Cycle, 53 Photosynthetic efficiency (PE), 6— 7, 306— 307, 309, 312, 486 Photosynthetic electron flow, 50, 105, 174 Photosynthetic electron transport chain, 50 Photosynthetic lamellae, 57— 58, 62, 320 Photosynthetic light, 45 Photosynthetic membranes, 57— 58 Photosynthetic microalgae, 307 Photosynthetic organisms, 53, 55 Photosynthetic oxygen evolution, 300 Photosynthetic photolysis of water, 157 Photosynthetic pigments, see also Pigments, 43, 64— 66, 188, 301, 434 Photosynthetic processes, 62 Photosynthetic products, 63 Photosynthetic rate, 81, 88—90 dissolved oxygen concentration, 299 flashing light, 288 light saturation, 81 oxygen pressure, 303 reduced, 88 Spirulina, 217, 219 Photosynthetic reaction rate, 289 Photosynthetic reserve material, 57 Photosynthetic systems, 50, 312 Photosynthetic unit, 70 Photosystem, 48—49 Photosystems I and II, 49— 52, 59, 62, 64, 73, 95, 183 Photo-topotactic reactions, 73— 74

518

CRC Handbook of Microalgal Mass Culture

Phototrophic microorganisms, 56 Phototrophs, 147— 148 Photovoltaic cells, 307 Photovoltaic power, 487 Phycobilins, 64, 66, 199, 4 3 4 -^ 3 6 Phycobiliproteins, 58, 71, 199, 435—436 Phycobilisomes, 58—60, 62, 72, 174, 238 Phycocolloids, 27, 29 Phycocyanin (PC), 58— 59, 66, 70, 72, 173, 220, 435—436, 445, 485 cancer treatment with, 229 S p ir u lin a , 225 Phycocyanin to chlorophyll-a ratio, 71 C-Phycocyanin, 64, 175, 183, 342, 350 R-Phycocyanin, 435 Phycocyanobilin, 435 Phycoerythrin, 58, 66, 435 R-Phycoerythrin, 64 Phycoerythrocyanin, 435 Phycotoxins, 372— 373 Physical factors in blue-green algae growth, 457— 459 Physiological temperatures, 301 Phytoene, 66 Phytoplankton, 34, 101, 219, 299 Phytotoxins, 372 Pig feed, 398, 399 Piggery wastes, 30, 36, 273, 395 Pigmentation, 71, 199, 324 D u n a lie lla , 234 Pigment-containing structures, 62 Pigments, 64, 91, 350, 432—436 carotenoids, 432— 434 chlorophylls, 432—435 composition, 63 photosynthetic, 301, 434 phycobilins, 434— 436 phycobiliproteins, 435— 436 S p ir u lin a , 225 Piping in pond facilities, 267— 268, 282— 283 Pisciculture, 402 Pitch, 213 Pitch ratio of helices, 214 Pits, 61 Planck’s constant, 44 Planktonic algae, 323 Planktonic diatoms, 34 Planktonic filamentous algae, 137 Planktonic rotifers, 326 Plankton soup, 387 Plant body types, 199 Plant hormones, 189— 190 Plant productivity, 14 Plasmalemma, 56, 232 Plasma membrane, 59, 61, 85 Plastid, 59, 60, 63, 65, 199 Plastid genomes, 65 Plastocyanin, 50 Plastoquinone, 50 P la ty m o n a s , 165 P le c to n e m a sp., 320

166, 456 Pleurocapsalean cyanobacteria, 56, 166 Polar lipids, 421—422 Polyculture effects of diets, 405 Polyethylene linings, 226, 250, 320 Polyethylene tubes, 222 Polyglucan granules, 214 Polyhedral body, 58 Poly-p-hydroxybutyrate (PHB), 345 Poly-p-hydroxybutyric acid (PHB), 443 Polymers, 441—443 Polymorphism, 214, 235 Polyphosphates, 57, 178— 180 Polysaccharides, 61, 63, 163, 174, 176, 349— 350, 441—442, 445 granules, 230 iron starvation, 183 P o r p h y r id iu m , 238 production, 238, 240 Polyunsaturated fatty acids (PUFA), 422—424 Polyvinylchloride, see PVC Pond construction and design, see also Ponds and other specific topics, 265— 283 back-to-back configuration, 269 blanket materials, 278 bottom lining, see also other subtopics hereunder, 276— 279 bottom roughness, 270, 277 channel length-to-width ratio, 270 clay blanket, 278 climate, 265— 266 C 0 2 feeding provisions, 282 coarse granular blankets, 278 concrete block, 280— 281 configuration, 268— 269 corrugated asbestos-cement roofing panels, 280 corrugated asbestos-cement wall, 281 costs, 224 crushed rock bottom lining, 281 deepened section, 272, 276— 277 electrical power and instrumentation, 283 geology, 266 high rate oxidation pond, see High rate oxidation pond hydraulics, 268— 272 laser leveling, 266 layout of facilities, 267— 268 limiting area, depth, and velocity relationships, 271 lining, 486 mixing velocity, 268, 272, 277 operating culture depth, 270 paddlewheel, 271— 276 paving materials, 278 piping, 282— 283 production inputs, 267 PVC lining, 486 rolled crushed rock bottom lining, 280 settleable solids control, 276 sheet membrane materials, 278— 279 site selection, 265— 268

P le u r o c a p s a ,

519 size, 268—272 slipform concrete, 280—281 sloping earth berms, 280 soil characteristics, 266 solids deposition, 268 spray-applied membrane materials, 279 topography, 266 traveling bridge, 277 traveling suction device, 276 turbulence enhancement, 279 wall construction, 276—277, 280—282 wall materials and design, 280—281 wall sections, 280 water quality and quantity, 266—267 Ponds, see also Pond construction and design, 30, 35—36 circular, 245 cleaning, 272 closed structures, 30, 245—247 high rate, 268, 273, 276 horizontal, 245—247 light-dark cycles, 252 lining, see also Polyethylene linings; PVC lin­ ings, 250 maintenance, see Outdoor mass cultures management, 312 meander, 245 outdoor mass cultures, see also Outdoor mass cul­ tures, 312—315 raceways, 245, 247, 249 temperature, 82 wastewater oxidation, 331—338 Population, 2, 21 Population density, 85, 291, 294, 301, 310, 318 algal growth, 326 growth rate in relation to, 315 light intensity in relation to, 285—292 logarithm of, 285 maintenance of, 474— 475 monitoring, 312—313 outdoor mass cultures, 285—292 output rate in relation to, 315 S c e n e d e s m u s , 236 seasons, effect of, 294, 316 S p ir u lin a production, 222 Pore, 58, 61 P o r m id iu m m o lle , 305 P orph yra la c in ia ta ,

26

sp., 26 P o r p h y r id ia le s ,

238

P o r p h y r id iu m

442 73, 75 arachidonic acid, 239, 425 essential fatty acids, 425 fatty acids, 427 growth physiology, 238 polysaccharides, 442 sterols, 430 spp., 60, 73, 238—239

a e ru g in e u m , cru en tu m ,

agar, 133 chemicals from, 485 growth medium for, 132 hydrogenase activity, 438 nitrogen utilization, 165 polysaccharide formation, 238, 442 potential for practical exploitation, 445 superoxide dismutase, 437 Potash, 27 Potassium, 157, 181— 182, 188, 220, 227 concentration, 92, 236, 322 Potassium nitrate, 209 Potassium to sodium ratio, 220 Poultry feed, 393— 398 Power costs, 487 PQ, see Plastoquinone Prasinophyceae, 59, 61—63, 65 Prasiola japonica, 28 yunnanica, 28 Predation, 111 Predators, 29, 233, 323 Premenstrual stress, 230 Premenstrual syndrome (PMS), 425 Preservation of microalgae, see also Laboratory techniques of microalgae cultivation, 125— 134 Primary productivity, 101 Primary xanthophyll, 66 Primitive fungi, 321 Prochloron, 64 Production costs, 34, 224, 485—487 Production systems for outdoor mass cultures, 285 Products from microalgae, 421—454 amino acid oxidase, 437 antibiotics, 443—444 ATP determination, 436— 437 carotenes, 421 carotenoids, 432— 434 chlorophylls, 421, 4 3 2 ^ 3 5 chlorosulfolipids, 422 environmental factors, 422—423 enzymes, 436—437 essential fatty acids, 424— 425 fatty acids, 421—427, 443 free fatty acids, 421 glycerides, 422 glycerol, 421 glycolipids, 422 hydrocarbons, 421, 430— 432 hydrogen, 438—439 hydrogenase, 438 lipids, see also other subtopics hereunder, 421— 432 luciferase, 436—437 luciferin, 436—437 methane, 439—440 nitrogenase, 438 nonpolar lipids, 421 osmoregulators, 440—441 pharmaceuticals, 443—445

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CRC Handbook of Microalgal Mass Culture

phenols, 443— 444 phosphoglycerate kinase, 436 phospholipids, 422, 424 phycobilins, 434— 436 phycobiliproteins, 435— 436 pigments, 432— 436 polar lipids, 421—422 poly-p-hydroxybutyric acid, 443 polymers, 441—443 polysaccharides, 441—442 prostaglandins, 424— 425 proteins, 441 restriction endonucleases, 437 starch, 442 sterol esters, 421 sterols, 421, 426, 430— 431 superoxide dismutase, 437 toxins, 444— 445 triglycerides, 421, 424 waxes, 421 wax esters, 421, 426, 429—430 Prokaryotes, 56— 59, 64, 164, 179 Proline, 92— 94, 165, 227, 441 Propeller, 251 Proplastids, 63 P r o r o c e n tr u m m in im u m , 427 Prostaglandin PGE,, 230 Prostaglandins, 230, 239, 42A— 425 Proteases, 173, 175, 221 Proteins, 59, 61, 72, 179, 188, 199, 341— 342, 441 amino acid release, 304 body composition and, 405 breakdown of, 80 content, 83, 90, 342 C h lo r e lla , 210 microalgae, 28 S c e n e d e s m u s , 236 S p ir u lin a , 222, 225 estimation, 342 hydrolysis, 92 malnutrition, 34 powders, 228 production, 9, 71 requirements, 399 S p ir u lin a , 221, 227 synthesis, 57, 182, 302 C h lo r e lla p y r e n o id o s a 82, 211 D u n a lie lla , 232 S p ir u lin a , 221 temperature dependence, 78 utilization, 363 Protein Efficiency Ratio (PER), 227, 354— 357, 367— 369, 380 Protein-protein interactions, 62 Protein regeneration studies, 362— 365 Protochlorophyllide (PChl), 65 Proton motive force, 85 Protoplast, 56 Protoplast division, 235 P r o to s y p h o n b o ty r o id e s , 423, 443 Prototroph, 147

Protozoa, 314, 320, 322— 324, 378 sp., 437 Provitamin A, 233, 349— 350 Provitamin A (3-carotene intake, 230 P r y m n e siu m p a r v u m , 342, 423 PS I and II, see Photosystems I and II P s e u d o a n a b a e n a , 165, 166, 456 P s e u d o s p o r a , 323 Pteridine, 94 Pumping, 487 Pumps, 251, 335 Pure microalgal meal, 29 Purification of microalgae, 125 Purine derivatives, 164 Purines, 164, 173, 342, 370— 371 Purity of outdoor cultures, 318 Putrescine, 165 PVC linings, 226, 250, 486 Pyrenoid, 201— 202 Pyrenoidal membranes, 63 Pyrenoid cap, 63 Pyrenoids, 59, 62—63, 230, 234 Pyrenoxanthin, 350 Pyridoxine (B6), 227, 349 Pyrimidine, 171 P y r r o p h y ta , see Dinoflagellates Pyruvate, 158, 162 Pyruvic acid, 171

P r o v id e n c ia

Q Q, 50 Q.o, 78 Quantitative estimations of light, 46 Quantum, 44 Quantum efficiency, 306 Quantum requirements, 219 Quantum theory, 43—47 Quantum yield, 112 Quaternary ammonium, 93

R Raceway ponds, 31, 222, 245, 247, 249, 285 Radiant flux, 45— 46 Radiation, 46, 295 Radiation flux, 86, 302 Radioactive carbon dioxide (14C 0 2), 53 Radiolabeled fatty acids, 426 Radiometric measure of light, 46 R a d io s p h a e r a n e g e v e n s is , 423 Random distribution, 291 R a p h id o n e m a , 165 Rapid growth, 97 Rats, mutagenicity study, 382 RBP, see Ribulose 1,5-bisphosphate Reaction center, 48— 51 Reaction rate (K), 79 Reactivity of oxygen, 299

521 Reactors, see also Mass cultivation, 245— 249 Recommended Daily Allowance (RDA) (U.S.) vitamin A, 226 vitamin B 12, 225 Red algae, see also specific types, 59— 61, 63— 64, 66, 139, 158 nutrient deficiencies, 188 photolithotrophy, 148 pigmentation, 199 Red drop, 49 Redfield ratio, 107 Redgwick Rafter, 121 Red light, 64, 71, 97 Reduced electron carrier, 301 Reduced ferredoxin (Fdred), 50, 176 Reduced plastoquinone, 50 Reduced Q, 50 Reductive Pentose Phosphate Pathway, see also Ribulose 1,5-bisphosphate, 53 Reforestation, 3, 17 Regular spirals, 215 Relative abundance, S p ir u lin a , 218 Relative humidity, 82 Renewable liquid fuel, 239 Reproduction, 381 C h lo r e lla , 200 S c e n e d e s m u s , 234— 235 Respiration, 28, 47, 56, 104, 108, 292, 309— 310, 312 high rate oxidation pond, 334 high temperature strains, 81 in the light, 56 low temperature strains, 81 phosphate uptake, 180 salinity, effect of, 94 Respiration rates, 71 Respiratory activity, 94 Respiratory consumption of oxygen, 56 Respiratory system, 175 Resting spores, 321 Resting state colonies, 239 Restriction endonucleases, 437 Rhamnose, 228, 349 R h a p h id iu m spp., 438 Rhaphidophyceae, 65 Rhizoids, 321 Rhodophyceae, 62, 65— 66, 200 Rhodophycophyta, 238 R h o d o p h y ta , see Red algae R h o d o tu r o la , 379 Riboflavin (B2), 225, 227, 349 Ribonucleic acid, see RNA Ribosomes, 58, 59, 61 70S Ribosomes, 62 Ribulose biphosphate, 160 Ribulose 1,5-bisphosphate (RBP), 53— 57, 69, 160, 173 Rice production, 11 Rigid polysaccharide cell wall, 230, 232 RNA, 61, 179, 346, 371 S p ir u lin a , 227

Rolled crushed rock bottom lining, 280 379 Rotating screens, 256 Rotation rate, 291 Rotifers, 322, 326, 327, 378 Rubidium, 181, 186 Rubifloc, 259 Rudimentary grana stacks, 64 Ruminants, feed for, 400—401 Ruthenium red, 201

R o ta r o r ia ,

s 362 Sahara, 28 Saline water, 215, 487 Salinity, 89—95, 298, 313 cyanophages, 320 P o r p h y r id iu m , 238 S p ir u lin a , 214, 217, 220 S a lm o n e lla , 380— 381 Salt-adapted cells, 92 Salt concentration, 94, 233, 239 Salt content, bodies of water, 217 Salt-induced SH-groups, 95 Salt marshes, 230 Salt resistance, 95, 203 Salt stress, 90, 95 Salt tolerance, C h lo r e lla , 203— 204 Salt water ponds, 215 Saprotrophy, 148 Sargasso Sea, 108 Saturated brine, 90 Saturated fatty acids, 82 Saturating light intensity, 286— 287, 310 Saturation effect, 236 Scale of production, 485 Scenedesmaceae, 200 S a c c h a r o m y c e s c e r e v is a e ,

S cen edesm u s a c u tu s

average growth factor, 325— 326 copper, 185 diseased cultures, 322 growth outdoors, 237 therapeutic properties, 406 viruses contaminating outdoor cultures of, 321 a c u tu s var. a lte r n a n s , 235 b iju g a te s , 236 d im o r p h u s , 188, 423 lo n g u s, 235 o b liq u u s, 29— 30, 33— 34 amino acid pattern, 344 antibiotic activity, 443 chemical composition, 342 digestibility, 351— 352 fatty acid composition, 345 manganese, 184 mineral composition, 348 molybdenum, 185 nutritive value, 361

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protein efficiency ratio, 355 sexual reproduction, 235 sterols, 426 toxins, 373 vanadium, 186— 187 vitamin content, 349 obtusiosculus, 440 quadricauda, 185, 342, 347, 349, 351 sp., 28, 30, 32, 34, 65, 83, 96, 199, 234— 236 acceptability, 390— 391 amino acid fortification, 370 Aphelidium sp. infecting, 325 autoflocculation, 258 automatic synchronization device, 135 bacterial load, 380 biological value, 359— 360 comparability data, 362 contamination by fungi, 378 contamination by insects, 378 correlation of temperature and irradiation with yield of, 296 cost-benefit analysis, 485—486 depletion-repletion studies, 364— 365 dermal toxicity, 385 digestibility, 351— 355 digestibility coefficient, 359— 360 drum drying, 260— 261, 485 fatty acids, 345 growth rate, 236— 237 harvesting, 485 heavy metal contamination, 374— 375 heavy metal content, 375 hepatic enzyme levels, 365 hydrogenase activity, 438 hypocholesterolemic effect, 407 industrial production, 236 infection of, 322 Loxodes magnus consumption of, 323 manganese, 184 mass culture, 235— 236 medium composition, 234 methane production, 439 microbial contamination, 380 molybdenum, 186 net protein utilization, 359— 360 nitrogen, 165, 359 nonprotein nitrogen, 342 nutritional requirements, 236 nutritive value, 399, 403 oral toxicity, 385 organic carbon nutrition, 162 organic compounds, 376, 377 parasites of, 322 pathogenic viruses, 321 pig feed, 398— 399 poultry feed, 393— 398 processing, 350— 351, 360 protein efficiency ratio, 357, 367— 368 protein regeneration, 363— 365 reproduction, 234— 235 ruminants, feed for, 400—401

side effects, 387—389 sloped cultivation units, 248—249 S p ir illu m - like bacterium attacking, 322 sulfate uptake, 181 supplementary effect of algae, 366—370 synchronized cultures, 120 taxonomy, 234— 235 therapeutic properties, 406—407 thylakoid membrane, 62 toxicological study, 384— 385 toxins, 373 uric acid, 371—372 world programs for mass cultivation, 306 yield as function of temperature and solar irra­ diation, 295—296 Schizophrenia, 230 S c h iz o th r ix c a lc io la , 444 Schumpeterian system, 473 S c o tie lla sp., 423 Scotland, seaweed industry, 27 Screening, 256—257, 317, 327, 337 S c y to n e m a , 166, 456 Sea lettuce, 26 Seasons or seasonal variations, 109, 293—294, 316 Seawater, S p ir u lin a , 217 Seawater ponds, 34 Seaweeds, 25—28, 200 Secondary carotenoids, 201, 204 Sedimentation, 256 S e le n a s tru m sp., 165, 347, 438 Selenium, 186, 188, 228 Semicontinuous culture, 120 Sensitive species, 302 Separation of algae, 141— 142 Separation disks, 216 Sep Pak, 434 Septa, 213—214 Serine, 93, 164— 165, 227 Settling chambers, 120 Sewage, 35—36, 109, 238 Sexual reproduction, 235 Shaded cells, 291 Shade type plants, 90 Shading, 250 Shallow culture, 209 Shallow culture depth, 292 Sheath characteristics, 214 Sheep, feed for, 400—401 Sheet membrane materials, 278—279 Shock reactions, 73 Shock treatment, 327 Short chains, 216 Short-term feeding experiments, 381—382 Short-term light deprivation, 237 Sialic acid, 228 Silica, 61, 163, 236 Silicon, 111, 184, 187— 188 Silicon dioxide, 199 Silver, 181 Singapore, 36 Single cell protein (SCP), 339—340

523 Single cells, 72 Sinking, 104, 110 Sinking filaments, 75 Siphoaxanthin, 350 Siphonein, 350 S ir o g o n iu m , 141 Site selection for pond construction, see also Pond construction and design, 265— 268 Sitosterol, 227 S k e le to n e m a

70, 82— 83, 427 sp., 35, 70, 164 Skin diseases, 406 Skin metabolism, 229 Slipform concrete pond walls, 280— 281 Sloped cultivation units, 247— 249 Sloping earth berms, 280 Soda, 27 Soda lakes, 112 Sodium, 181— 182, 220, 227 Sodium acetate, 210 Sodium bicarbonate, 231, 313 Sodium carbonate, 217 Sodium chloride, 90— 94, 204, 231, 233 Soil characteristics for pond system, 266 Soil conditioner, 240 Soil factors in blue-green algae growth, 459 Soil-inhabiting bacterium, 235 Soils, 240 Soil water medium, 134 Solar constant, 46 Solar drying, 30 Solar energy, 47, 223, 238, 252, 291, 307 Solar heating in winter, 226 Solar intensity, 295 Solar irradiance, see also Irradiance, 82, 222, 312, 314— 315 outdoor mass cultures, 285— 297 S c e n e d e s m u s , 235— 236 Solar irradiation, 306 Solar radiance, interrelation with temperature, 292— 297 Solar radiations, 46, 250 Solar steam, 487 Solar thermal energy, 307 Solids deposition in ponds, 268 Soluble molecules, transport of, 61 Sonication, 62 Sorbitol, 91, 440 Sorghum, 56 South Africa, 34 South Korea, 17 Soybean harvest, 7 Space research, microalgae, 32— 33 Species competition, 84 Species dominance, 83— 84, 111 Species specificity, 217 Specific growth rate, 78— 80, 117— 118, 157, 294— 295, 315 S c e n e d e s m u s , 237 S p ir u lin a p la te n s is , 219 c o sta tu m ,

Spectral quality of incident light, 71 Spectrophotometer, 121 Speed of light, 43 Speirs Levy, 121 Spherical cells, C h lo r e lla , 200— 201, 203 Spherical walls with girdle-shaped chloroplasts, 202 Spinach, 349 Spines, 61, 96, 234 Spin restriction, 299 Spiral pitch, 216 Spirals, 215— 216 S p irillu m -W k t bacterium, 322 S p ir o g y r a , 28, 141, 322 S p ir u lin a

214 215 je n n e r i, 215 m a x im a , 35, 215 alkaline lakes, 217 amino acid pattern, 344 carbohydrates, 349 cell dimensions, 213 chemical composition, 342 commercial production, 224 fatty acid composition, 345 methane production, 439 mineral composition, 348 nucleic acid nitrogen, 342 vitamin content, 349 p la te n s is , 28, 81, 95— 96, 111, 215 alcohols, 347 alkaline lakes, 217 amino acid pattern, 344 biological value, 360 cell dimensions, 2.13 chemical composition, 342 closed cultivation systems, 246 digestibility, 351, 360 essential fatty acids, 425 fatty acids, 345, 427 helicity, 214 hydrocarbons, 347 intensity of stirring, effect of, 289— 290 interrelation between maximum daily tempera­ ture, irradiance and photosynthetic activity, 297 Lakes of Africa, 218 y-linolenic acid, 226, 228, 426 mineral composition, 348 modifying winter temperature, effect of, 219 net protein utilization, 360 nonvacuolated from Lake Chad, 213 outdoor monoalgal culture, 317 output rate, 295 photosynthetic activity, 295 poly-(3-hydroxybutyrate, 344 poly-P-hydroxybutyric acid, 443 proteases, 173 protein efficiency ratio, 355 relationship between 0 2 concentration, temper­ ature and light intensity, 297

fu s if o r m is , g e ittle r i,

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CRC Handbook of Microalgal Mass Culture

respiration, 219 screening, 257 specific growth rate, 219 sterols, 347 straight non vacuolated trichomes, 213 straight vacuolated trichomes, 213 unialgal bloom, 217 vacuolated, 213 vitamin content, 349 sp., 25, 28— 30, 32, 34— 35, 75, 84, 199, 212— 230, 339 airlift to create turbulence, 252 amino acids, 343, 344, 369— 370 bacterial contamination, 379 bacterial count, 322 bacterial load, 380— 381 bicarbonate in culture, 313 biological value, 359— 360 cancer, 229, 230 cell density, 315 cell wall, 349 centrifugation, 255— 256 chemical composition, 225, 227— 228 chemicals from, 485 chitosan, 259 chlorophylls, 435 clinical applications, 229— 230 closed cultivation system, 247 C 0 2 distribution, 255 commercial production, 223— 224, 226, 228 comparability data, 362 competition with Chlorella, 317— 318 contamination, 378 dense population of, 217 depletion-repletion studies, 366 dermal toxicity, 385 deterioration of culture, 314 digestibility, 351— 352, 354— 355 digestibility coefficient, 359— 360 ecology, 217— 219 empirical application, 478— 481 environmental limitations, 295 fatty acid composition, 345 filamentous bacteria, 322 fish feed, 224 food coloring, 224 food supplement for humans, 221— 222 gas vacuolated, 213 growth medium for, 128, 220 habitat, 217—219 harvesting regime, 316 health food, 224, 228— 229 heavy metal contamination, 374 hepatic enzyme levels, 365 high filament densities, 214 high-methionine mutants, 344 history of, 221 human enzyme reactivation, 230 hypocholesterolemic effect, 407 Lakes of Africa, 218 life cycle, 216— 217

y-linolenic acid stimulation, 230 mass cultures, cultivation of, 222— 224 methane production, 439 microbiological analysis, 228 mixed culture with Chlorella, 319 monoculture of, 318 morphological types, 213 morphology, 212— 216 mutagenicity, 382 net protein ratio, 360 net protein utilization, 359— 360, 367 nitrogen balance, 359 nutrition, 220— 221 oral toxicity, 385 organic compounds, 376, 377 organ weights, 382 outdoor culture, 319 output rate, 316 oxygen concentration, 302, 314 pH, 159 photooxidation, 303 photooxidative death, 314 photosynthetic system, 296 phycocyanin, 435 physical properties, 228 pig feed, 399 pigments, 350 polysaccharide, 349 potential for practical exploitation, 445 poultry feed, 396— 397 processing, 360 prostaglandin stimulation, 230 protein efficiency ratio, 357, 367— 368 proteins, 228, 363— 365, 441 purity of outdoor cultures, 318 ruminants, feed for, 400— 401 screening, 256 sonic dehydrator, 261 specific growth rate, 294 superoxide dismutase, 437 supplementary effect of algae, 367— 370 taxonomy, 212— 216 temperature, 219, 223 therapeutic properties, 229— 230, 406— 407 thyroid stimulation, 229 toxicity, 383 toxins, 373 urea, effect of, 220 vitamin content, 348, 349 world programs for mass cultivation, 306 wound treatment, 229 Spongiococcum, 35, 357 Sporangium, 202, 321 Spray-applied membrane materials, 279 Spray drying, 224, 260 Spring, 294, 296— 297, 316 Stacked thylakoids, 62 Stalked pyrenoid, 63 Standard plate count, Spirulina, 228 Staphylococci, 379— 381 Starch, 47, 60, 91, 188, 199, 350, 442

525 accumulation, 211 grains, 202 sheath of, 63 synthesis, 55 Starch-like compounds, 199 Starter poultry feed, 395 Stationary phase, 118 Stationary-state phase, 118 Steady state, S p ir u lin a production, 223 Steady-state light-saturated conditions, 107 Steady-state phase, 119 Steady-state response to salinization, 94 Stearic acid, 227 S te p h a n o p y s is c o s ta ta , 164 Sterilization, 125 Sterol esters, 421 Sterols, 346— 347, 421, 426, 430— 431 S p ir u lin a , 227 S tic h o c o c c u s m ir a b ilis , 443 S tig e o c lo n o iu im sp., 342, 351 Stigmasterol, 227 S tig o n e m a , 166, 456 Stillage, 17 Stirring, 487 Stirring speed, 289, 299, 314 Stop responses, 73 Storage lipid, 211 Storage of food reserves, 47 Storage polysaccharide, 63 Storage product, 237 Stored biomass, 3 Straining, 256— 257, 336— 337 Stratification, 75, 109 Stratosphere, 46 Streaks, 101 Streptococci, 379 Streptomycin, 125, 180 S tr o b ilid iu m sp., 322, 378 Stroma, 62 Strong light, 209 Strontium, 181— 182, 186 S ty lo n ic h ia sp., 322, 327, 378 Subacute toxicity, 381 Subsistence quota, 108 Substrate, 55 Substratum, 62 Subsurface irradiance, 106 Subzero temperatures, 95 Succinate, 162 Sucrose, 91—94, 162— 163, 350 C h lo r e lla , 202 S c e n e d e s m u s , 236 synthesis, 55, 93 Sugar, 19, 421 Sugar cane, 17, 56 Sulfated polysaccharides, 59, 61, 199 Sulfated sugars, 238 Sulfate uptake, 180— 181 Sulfide, 181 Sulfolipids, 180 Sulfonated carrageenan-like polysaccharides, 485

Sulfur, 157, 180— 181, 188, 443 Summer, 294—297, 302, 315—316 Sun, electromagnetic radiation from, 46 Sun drying, 261 Sunlight, 29, 4 6 - 4 7 , 209, 288 intensity, 136— 137, 288 Sunlight visible region efficiency, 310 Sun type plants, 90 Superoxide anion radical, 299 Superoxide dismutase (SOD), 299, 301, 437 Supersaturated oxygen, 314 Supplementation studies, 365— 370 Surface drying, 260 Surface plating of blue-green algae, 142 Surface scums, 75—76, 110 Surplus food, 18 Swimming algae, 102 Symbiosis, 111 nitrogen fixation, 456 , 464— 465 Symbiotic association, 101, 148 Symbols, list of, 482—483 Synchronized batch cultures, 118— 119 Synchronized cultures, 120 Synchronized suspensions, 208 Synchronous cultures, 155 Synchrony, induction of, 118 S ynechococcus

438 95 sp., 78, 97, 165— 166, 342, 456 S y n e c h o c y s tis sp., 165, 291, 396— 397, 438— 440 Systemic mixing, 291 e lo n g a tu s, liv id u s ,

T Taiwan, 29, 226 Taxis, 73, 124 Taxonomy, 214 C h lo r e lla , 200—203 D u n a lie lla , 230— 231 P h a e o d a c ty lu m , 236 P o r p h y r id iu m , 238 S c e n e d e s m u s , 234— 235 S p ir u lin a , 212—216 Tecuitlatl, 221— 222, 339 Temperature, 29, 55, 77— 87, 111, 286, 301, 311, 314, 317 algal growth, 326 algal movement, 73 algal production in mass cultures, 310 ambient, 82 cell division, 208 cell length, 95—96 cell morphology, 95— 96 cell movement, 85 chemical composition, 82— 83 chilling, 84— 85 C h lo r e lla , effect on, 205— 207 chytrid parasitism, 322 correlation with irradiation, 296

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CRC Handbook of Microalgal Mass Culture

cultivation of microalgae, 126 cyanophages, 320 daily course, 295 dark respiration, 80— 81 decline in, 96 D u n a lie lla , 231 extreme, 94 feeding, effect on, 325 functions of, 288, 295 growth limiting, 222 growth medium, effect on, 80 growth rate, 78— 80 high, 81— 82, 96, 231 high rate oxidation pond, 335— 336 inhibitory effect, 79 interrelation with solar irradiance, 292— 297 length as function of, 97 light interaction with, 86— 89 light saturation parameter, 88 maintenance of monoalgal culture, 318 modifying winter, effect on S p ir u lin a p la te n s is , 219 0 2 solubility as function of, 298 organelles influenced, 214 outdoor mass cultures, 292— 297 output of biomass, 310 photosynthesis and photorespiration, 305 pond, 82 P o r p h y r id iu m , 238 protozoa affected by, 323 relationship with 0 2 concentration and light inten­ sity, 297 rise in, 219 S c e n e d e s m u s , 235— 236 seasonal variations, 293— 294 species dominance, 83— 84 S p ir u lin a , 219, 222— 223 sulfate uptake, 181 thermotaxis, 85 variations in, 292, 314 Temperature-dependent daylength effect, 88 Temperature growth response curve, 79 Tetraterpenes, 66 Teratogenicity, 381 Terminal cell, 215— 216, 234 Terpenoid origin, 239 T e tr a s e lm is , 187 Textile printing, 28 Thailand, 28, 226 T h a la s s io s ir a , 173 p s e u d o n a n a , 83 Thallium, 181 Thames River, England, 109 Therapeutic properties, 406— 408 S p ir u lin a , 229— 230 Thermal gradient device, 135— 136 Thermal springs, S p ir u lin a , 217 Thermistors, 46 Thermodynamic efficiency, 310 Thermophilic algae, 78, 82, 206, 208, 219, 246 Thermophily, 204

Thermopiles, 46 Thermotaxis, 85 Thiamin (B,), 180, 189, 201, 349 C h lo r e lla , 203— 204 S p ir u lin a , 225, 227 Thiol compounds, 187 Threonine, 227 Thrombosis, 425 Thylakoids, 48, 51— 53, 56— 59, 62—63, 65, 174 Thyroid stimulation, 229 T ila p ia , 36, 4 0 4 -^ 0 5 Tin, 181 Titanium, 181, 186 Titerpen alcohols, 227 Tocopherol, 227 p-Toluenesulfonamide, 326 T o ly p o th rix , 166, 456 c o n g lu tin a ta , 429 te n u is, 71 Toxicity, 86— 87, 184— 185, 299— 303, 370— 373, 381— 386 Toxins, 372— 373, 377, 4 4 4 ^ 4 5 Trace elements, 86, 126 Trace metals, 189 Translocation, 101 Transmission electron microscopy, 56 Transparent plastic tubes, 212 Trehalose, 350 1-Triacontanol (TRIA), 189 Triacylglycerol synthesis, 211 T r ib o n e m a , 165 a e q u a le , 430 Trichocysts, 59 Trichoderma, 351 T r ic h o d e sm iu m , 109 T r ic h o d e sm u s, 456 Trichome color, 214 Trichomes, 56, 213, 215— 217 Triglycerides, 345— 346, 421, 424 Triose phosphate molecules, 54— 55 Triradiate cells, 236 Tryptothan, 227 Tungsten, 186 Turbidity, see Light scattering Turbidostat, 120, 137— 139, 318— 319 Turbulence, 207— 209, 277, 290— 291 airlift, see also Airlift, 252 creation techniques, 251— 252 enhancement, 279 flashing light effect, 252, 270, 277 flowing culture, 289 foils, 277, 279 gravity flow, 251 high, 312 injector, 252 light distribution, 315 mass cultivation, 250— 253 paddlewheel, see also Paddlewheel, 251 propeller, 251 pumps, 251 S p ir u lin a , 223

5 27 Turbulent flow, 222, 291 Turgor potential, 93 Turgor pressure, 75 Tyrosine, 227

u Ubiquinone, 50 Ulcers, 406, 424 Ulcers cruris, 406 U lo th rix

427 sp., 342, 351 Ultrastructure of microalgae, 56— 64 U lv a la c tu c a , 26 Uncouplers, 175, 181 a e q u a lis ,

Vestigial cell wall structure, 64 Vibrating screens, 256, 317, 337 Violaxanthin, 350, 4 3 2 -^ 3 3 Viruses, see also Cyanophages, 29, 320— 321 Visible fraction of total sunlight, 310 Visible light, 46-^17, 73, 85 Visible wavelengths, 47 Vision, 46 Vitamin A, 225— 226, 230 Vitamin B12, 186, 189, 225, 348 Vitamin C, 349 Vitamin E, 349 Vitamins, 126, 163, 188— 189, 222, 227, 347— 349 Volutin, 57 V o lv o c a le s , 230 V olvo: c c a r te r i, 422 V o r tic e lla sp., 322, 378

U n d a ria , 21 p in n a tifid a , 26 Undemourishment/poverty problem, 5 Underwater light field, 104, 111 UNESCO procedure, 123 Unicellular algae, 59, 61, 199— 200, 235 Unicellular cyanobacteria, 166 Unicellular eukaryotes, 64 Uninucleate cells, 234 United Kingdom, 12 United States, 12, 17— 18, 27, 226 U .S.S.R ., 11, 25, 31 Unsaponifiable matter in lipids, 227, 346— 347 Urate, 165 Urea, 126, 164— 165, 171— 173, 186 C h lo r e lla , 209, 210 P o r p h y r id iu m , 238 S p ir u lin a , 220, 222, 224 Urea amidolyase, 172— 173 Urea carboxylase, 172 Urease, 172 Uric acid, 164, 389 excretion capacity, 372, 397 levels in humans, 370— 371 Uric acid stones, 370 Uridine, 165 U ro n em a

342, 347, 351 sp., 351, 357, 362 te r r e s tr e , 347 UV, 45— 47, 73, 111, 125, 322 UV-induced mutations, 46 g ig a s ,

v V a c u la ria v ir e s c e n s , 432 Vacuoles, 59, 60, 202 Valine, 227 Vanadium, 181, 185— 187 Vasodepressor, 445 Vegetable oils, 19 Vegetative cells, 56, 200, 231 Velocity of propagation (V), 43

w Wall composition, 199 Warm-temperature algae, 81 Wastewater oxidation ponds, see also High rate oxi­ dation ponds (HROP), 331— 338 Wastewater systems, 285 Wastewater treatment 29, 35— 36 Water blooms, see Blooms Water cleavage, 49 Water potential, D u n a lie lla , 232 Water quality and quantity for pond system, 266— 267 Water retention, 240 Water stress, 90, 95 Water temperature, 314 Wavelength (\), 43-^16, 49, 52 Wavelength of light, 43— 45 Wavelengths of solar irradiation, 307 Wave phenomenon, 43 Waxes, 421 Wax esters, 421, 426, 4 2 9 ^ 3 0 Weight gain, 380, 393, 398 W e s tie lla , 166, 456 W e s tie llo p s is , 166, 456 Wheat output, 11 Windmill, 30 Wine, 19 Winter, 294— 297, 315— 316, 320, 327 S p ir u lin a production, 222 Wood, 2, 21 World Bank, 14 World land use, 11— 12 Wound treatment, S p ir u lin a , 2 2 9

x Xanthine, 164 Xanthophyceae, 62, 65, 189, 422 Xanthophylls, 66, 199, 227

528

CRC Handbook of Microalgal Mass Culture Scenedesmus, 235 sloped cultivation units, 248— 249 Spirulina, 224

Xenococcus, 166, 456 Xenon lamps, 290 Xylan, 61, 199

Y Yeast extract, 163, 203, 234 Yields, 295, 306— 307, 309— 310 Chlorella, 207, 210—211, 304 correlation of temperature and irradiation with, 296 mass culture of microalgae, 32 maximum potential, 307 overall, 320

z Zeaxanthin, 72, 350, 4 3 2^133, 445 Spirulina, 227 Zinc, 181, 184— 185, 227, 230 Zirconium, 181 Zooplankton, 32, 36, 101— 102, 109, 317, 324— 327 Zoospores, 321— 322 Zygote, 230, 231