Table Of ContentCRC
Handbook
o f
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. Essentially the entire plant body has nutritional value since only a minor portion of
the cell is indigestible.
2. 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.
3. 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.
4. 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. David O. Hall, Ph.D.
Professor Professor
Department of Biology Department of Plant Sciences
Queens College King’s College London
Flushing, New York London, United Kingdom
Eithan Hochman, Ph.D.
Aharon Abeliovich, Ph.D.
Professor
Senior Lecturer
Department of Economics
The Laboratory of Environmental
Ben-Gurion University of the Negev
Microbiology
Beer-Sheva, Israel
The Jacob Blaustein Institute for Desert
Research
Drora Kaplan, Ph.D.
Ben-Gurion University of the Negev
Lecturer
Sede Boqer. Israel
The Laboratory of Environmental
Microbiology
Yael J. Avissar, Ph.D. The Jacob Blaustein Institute for Desert
Lecturer Research
Department of Biology Ben-Gurion University of the Negev
Ben-Gurion University Sede Boqer, Israel
Beer-Sheva, Israel
S. Herman Lips, Ph.D.
Professor
E. W. Becker, Ph.D.
Department of Biology
Senior Scientist
Beer-Sheva and the Salinity and Plant
Institute of Chemical Plant Physiology
Physiology Lab at Sede-Boqer
University of Tubingen
Ben-Gurion University of the Negev
Tubingen, Federal Republic of Germany
Israel
Zvi Cohen, Ph.D. Amos Richmond, Ph.D.
Lecturer Professor
Microalgal Biotechnology Laboratory Microalgal Biotechnology Laboratory
The Jacob Blaustein Institute for Desert The Jacob Blaustein Institute for Desert
Research Research
Ben-Gurion University Ben-Gurion University of the Negev
Sede Boqer, Israel Sede Boqer, Israel
Carl J. Soeder, Ph.D.
Joseph C. Dodd, Ph.D.
Professor
Consulting Engineer
Institute for Biotechnology
Microalgae Technology
Kernforschungsanlage GmbH
Brawley, California
Julich, Federal Republic of Germany
Zvi Dubinsky, Ph.D. Y. Tsur, Ph.D.
Senior Lecturer Professor
Department of Life Sciences Department of Economics
Bar-Ilan University Ben-Gurion University of the Negev
Ramat-Gan, Israel Beer-Sheva, Israel
L. V. Venkataraman, Ph.D. Avigad Vonshak, Ph.D.
Scientist and Area Coordinator Senior Scientist
Autotrophic Cell Culture Discipline Microalgal Biotechnology Laboratory
Central Food Technology Research The Jacob Blaustein Institute for Desert
Institute Research
Mysore, India Ben-Gurion University of the Negev
Sede Boqer, Israel
TABLE OF CONTENTS
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 Factors.................................................................................69
Productivity of Algae Under Natural Conditions................................................................107
Laboratory Techniques for the Cultivation of Microalgae...................................................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 Microalgae........................................................................................................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
