HYCOVIRUS
BIBLIOGRAPHY
U.S. ENVIRONMENTAL PROTECTION AGENCY
OFFICE OF RESEARCH AND DEVELOPMENT
ENVIRONMENTAL RESEARCH CENTER
CINCINNATI, OHIO
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RESEARCH REPORTING SERIES
Research reports of the Office of Research and Development, U.S. Environmental
Protection Agency, have been grouped into nine series. These nine broad cate-
gories were established to facilitate further development and application of en-
vironmental technology. Elimination of traditional grouping was consciously
planned to foster technology transfer and a maximum interface in related fields.
The nine series are:
1. Environmental Health Effects Research
2. Environmental Protection Technology
3. Ecological Research
4. Environmental Monitoring
5. Socioeconomic Environmental Studies
6. Scientific and Technical Assessment Reports (STAR)
7. Interagency Energy-Environment Research and Development
8. "Special" Reports
9. Miscellaneous Reports
This document is available to the public through the National Technical Informa-
tion Service, Springfield, Virginia 22161.
COVER PHOTO:
Electron micrograph of the virus AS-1 which infects the unicellular
blue-green algae Anacystisnidulansand Synechococcus cedrorum.
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EPA-600/9-77-008
May 1977
PHYCOVIRUS
BIBLIOGRAPHY
"Compiled by
Robert S. Safferman Ph.D.
and
Mary-Ellen Morris
1977
VIROLOGY SECTION
BIOLOGICAL METHODS BRANCH
ENVIRONMENTAL MONITORING AND SUPPORT LABORATORY
OFFICE OF RESEARCH AND DEVELOPMENT
U.S. ENVIRONMENTAL PROTECTION AGENCY
CINCINNATI, OHIO 45268
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DISCLAIMER
This report has been reviewed by the Environmental Monitoring and
and Support Laboratory, U.S. Environmental Protection Agency, and
approved for publication. Mention of trade names or commercial
products does not constitute endorsement or recommendation for use.
11
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PREFACE
A literature search program was developed for phycoviruses as part of earlier efforts to
provide adequate coverage of relevant research. It was accomplished primarily through the use
of an in-depth data base from which source material could be compiled in a continuous and
comprehensive form. The citation index that resulted has been shared with those needing
background information on algal lysing agents. The need for a formal publication soon became
evident from the many requests for such material. The first edition of the bibliography was
published in 1974 and updated in 1975.
The present edition retains the organization and scope of preceding editions with the
citations divided into two sections. The first, pertaining to viruses of procaryotic algae, has
been more thoroughly researched and thus is better understood. Owing to these advances, the
term "phycoviruses", introduced to denote algal viruses in general, has become almost
synonymous with blue-green algal viruses. The second section, dealing with viruses of
eucaryotic algae, is virtually unchanged from that of the preceding edition. Clearly, isolation of
these viruses has presented a formidable obstacle to researchers. Of the reports listed, none
have penetrated the obscurity that has surrounded their identification and assessment.
Reflecting on the procaryotic organization shared by bacteria and blue-green algae, it is
evident that many morphological and chemical similarities are likely to exist between the
viruses infecting them. Some authors have in fact followed bacterial workers and used the term
"cyanophage" for viruses of blue-green algae. As with "cyanophages" and bacterial viruses, a
parallel relationship might be foreseen between viruses of eucaryotic algae yet to be confirmed
and those viruses known to infect higher plants. At present, available evidence is plagued by
vagueness and confusion with no promise of an early answer.
The number of citations in the bibliography has nearly doubled during the three years that
have elapsed since publication of the first edition. This exhaustive survey of the literature
reflects on the different directions that phycovirus research has taken. It provides an insight to
the current state-of-the-art which we hope will serve as a stimulus for future research and
discussions on the subjects.
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ACKNOWLEDGEMENTS
We are grateful to the following colleagues from Environmental
Research Center Library Services who have assisted in the phycovirus
literature search program: Dottie Brofft, Lillian Bosworth, Mary
Calkins, Dora Rusk and Maxine Smith. We also owe a special debt to
Rebecca Blazek for her invaluable help in providing secretarial and
typing assistance.
IV
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INTRODUCTORY REMARKS
These introductory remarks are intended to bring to the readers' attention the significant
trends in the developing field of phycovirus research. Although this field may appear to be of
limited scope in the overall context of virology, its domain is nevertheless extensive. It includes
divergent lines of development that open to exploration the many facets of basic and applied
virus research. Many early problems have yet to be resolved affording investigators a wide
variety of research opportunities.
Although phycoviruses were still unknown in the early 1960's, we considered it unlikely
that the biological characteristics of the algal cell were such that the entire group would be
immune to viral infection. During this time waste stabilization ponds became the focal point of
our search for this viral type. Supporting a dense and fluctuating algal population, these ponds
offered a unique environment from which to isolate the elusive phycovirus. In 1963 a pond in
Indiana yielded the first definite evidence that such a disease existed in algae. The discovered
infectious agent was the LPP-1 virus which caused lysis in several blue-green algal species of
the genera Plectonema, Phormidium, and Lyngbya. Confirmation of data supporting the
existence of this virus group soon followed, as did the interesting aspect of their wide
geographic distribution.
Historically, no notice was taken of phycoviruses during the nearly 50 years following the
first description of a filterable virus of bacteria. This time lapse between the two discoveries was
perhaps inevitable in that, unlike bacteria, algae had neither been closely associated with
animal or human diseases nor had they been demonstrated as a pathogenic agent of
agricultural crops. Also important was the fact that culturing was non-essential in many earlier
studies as algae were readily recognizable in samples collected directly from their natural
habitats.
The events which ultimately led to the detection of phycoviruses originated from a need to
better understand the underlying factors responsible for algal decomposition. With the
discovery of the phycovirus, a new concept in algal pathology evolved since these viruses had
the capacity to exercise a far-reaching effect on the development of algal communities. Our
knowledge is still too limited to relate their activity directly to specific incidences of algal
fluctuations. However, the magnitude and distribution of one group of these viruses have led
some to believe that their lytic activity precludes several susceptible algal species from attaining
objectionable proportions. At a time of increasing concern about algal overabundance in the
aquatic environment, the very nature of the phycovirus clearly points to its possible
development as an alternative to chemical control. Such application has popular appeal and
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support in offering a basic approach to algal population ecology and control. Early attempts to
establish a broad collection of these viruses led to extensive surveys of the environment that
consistently yielded the LPP viral types from which our early concepts were developed.
Considering their distribution and ease of detection, it is difficult to comprehend the
infrequency with which other phycovirus groups have been isolated. These ecological
considerations, however, afford only one view of the experimental aspects realized from their
discovery.
Much of the current interest is focused on the inherent properties of the phycovirus-host
system. It is not surprising that this work has closely followed the lead taken by bacteriophage
studies. Like the phage system, the phycovirus system can be manipulated with ease and
provide clearly-defined conditions often not attainable in higher organisms. Its basic
application as a model to further our understanding of the function and organization of
photosynthesizing cells seems destined to lead to the numerous inroads in molecular biology
that have already been realized through studies with phage-bacteria systems.
VI
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VIRUSES OF PROCARYOTIC ALGAE
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Adolph, K.W. and Haselkorn, R. Photosynthesis and the development of blue-green algal virus N-l. Virology
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Adolph, K.W. and Haselkorn, R. Comparison of the structures of blue-green algal viruses LPP-1M and LPP-2
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Adolph, K.W. and Haselkorn, R. Isolation and characterization of a virus infecting a blue-green alga of the genus
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Allen, M.M. and Hutchison, F. Effect of some environmental factors on cyanophage AS-1 development in
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Booth, S.J. Isolation, identification, and partial characterization of cyanophage LPP-2N. Ph.D. Thesis,
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Brown, R.M., Jr., Smith, K.M., and Walne, P.L. Replication cycle of the blue-green algal virus LPP-1. Nature
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Cannon, R.E. The effect of stress and non-stress conditions upon the interaction of Plectonema boryanum and the
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Goldstein, D.A. Some biological, chemical and physical properties of blue-green algal virus LPP-1. Ph.D. Thesis,
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Goldstein, D.A. and Bendet, I.J. Physical properties of the DNA from the blue-green algal virus LPP-1. Virology
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Goldstein, D.A., Bendet, I.J., Lauffer, M.A., and Smith, K.M. Some biological and physicochemical properties of
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Goryushin, V.A. and Chaplinskaya, S.M. Existence of viruses of blue-green algae. Mikrobiol. Zh. Akad. Nauk.
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Luftig, R.B. Studies on cyanophage LPP-1. Ph.D. Thesis, University of Chicago, 1967.
Luftig, R. and Haselkorn, R. Morphology of a virus of blue-green algae and properties of its deoxyribonucleic
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Moskovets, S.M., Mendzhul, M.I., Nesterova, N.V., Dyachenko, N.S., Vantsak, N.P., and Lysenko, T.G.
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Singh, R.N., Singh, P.K., Kashyap, A.K., Sarma, T.A., Dhar, B., Chaube, I.J., and Choudhury, I.D. Isolation and
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VIRUSES AND VIRUS-LIKE AGENTS OF EUCARYOTIC ALGAE
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TECHNICAL REPORT DATA
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Phycovirus Bibliography
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16. ABSTRACT
The volume comprises a comprehensive survey of the phycovirus
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present time.
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KEY WORDS AND DOCUMENT ANALYSIS
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Viruses
Algae
Bibliography
Phycoviruses
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