advancing the chemical sciences

ISBN 0-85404-301-2

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Preface for the ESP series in photochemical and photobiological sciences

"Its not the substance, it's the dose which makes something poisonous!" When Paracelsius, a German physician of the 14th century, made this statement he probably did not think about light as one of the most obvious environmental. But his statement applies as well to light. While we need light, for example for vitamin D production, too much light might cause skin cancer. The dose makes the difference. These diverse findings of light effects have attracted the attention of scientists for centuries. The photosciences represent a dynamic multidiscip-linary field which includes such diverse subjects as behavioral responses of single cells, cures for certain types of cancer and protective potential of tanning lotions. It includes photobiology and photochemistry, photomedicine as well as the technology for light production, filtering and measurement. Light is a common theme in all these areas. In recent decades a more molecular centered approach has changed both the depth and the quality of the theoretical as well as the experimental foundation of photosciences.

An example of the relationship between global environment and the biosphere is the recent discovery of ozone depletion and the resulting increase in high energy ultraviolet radiation. The hazardous effects of high energy ultraviolet radiation on all living systems is now well established. This discovery of the result of ozone depletion put photosciences in the center of public interest with the result that in an unparalleled effort scientists and politicians worked closely together to come to international agreements to stop the pollution of the atmosphere.

The changed recreational behavior and the correlation with several diseases in which sunlight or artificial light sources play a major role in the causation of clinical conditions (e.g. porphyrias, polymorphic photodermatoses, Xeroderma pigmentosum and skin cancers) have been well documented. As a result, in some countries (i.e. Australia) public services inform people about the potential risk of extended periods of sun exposure. The problems are often aggravated by the phototoxic or photoallergy reactions produced by a variety of environmental pollutants, food additives or therapeutic and cosmetic drugs. On the other hand, if properly used, light-stimulated processes can induce important beneficial effects in biological systems, such as the elucidation of several aspects of cell structure and function. Novel developments are centered around photodiagnostic and phototherapeutic modalities for the treatment of cancer, artherosclerosis, several autoimmune diseases, neonatal jaundice and others. In addition, classic research areas like vision and photosynthesis are still very active. Some of these developments are unique to photobiology, since the peculiar physico-chemical properties of electronically excited biomolecules often lead to the promotion of reactions which are characterized by high levels of selectivity in space and time. Besides the biologically centered areas, technical developments have paved the way for the harnessing of solar energy to produce warm water and electricity or the development of environmentally friendly techniques for addressing problems of large social impact (e.g. the decontamination of polluted waters). While also in use in Western countries, these techniques are of great interest for developing countries.

The European Society for Photobiology (ESP) is an organization for developing and coordinating the very different fields of photosciences in terms of public knowledge and scientific interests. Due to the ever increasing demand for a comprehensive overview over the photosciences the ESP decided to initiate an encyclopedic series, the "Comprehensive Series in Photochemical and Photo-biological Sciences". This series is intended to give an in-depth coverage of all the very different fields related to light effects. It will allow investigators, physicians, students, industry and laypersons to obtain an updated record of the state-of-the-art in specific fields, including a ready access to the recent literature. Most importantly, such reviews give a critical evaluation of the directions that the field is taking, outline hotly debated or innovative topics and even suggest a redirection if appropriate. It is our intention to produce the volumes at a sufficiently high rate to generate a timely coverage of both well established and emerging topics. As a rule, the individual volumes are commissioned; however, comments, suggestions or proposals for new subjects are welcome.

Donat-P. Hàder and Giulio Jori Spring 2002

Volume preface

The surge in the systematic study of UV effects on aquatic habitats is contemporary with the discovery of the ozone hole in the 1980s. Since then, and for the last two decades, the number of publications on UV related issues has grown virtually exponentially. Paralleling the explosive development of this new "field", a number of reviews have attempted to summarize the available knowledge in the primary literature. These works have evolved from environmental agencies' reports to symposia volumes to multi-authored edited books, many of which are excellent, some of which are reasonably comprehensive, and a few of which are quite recent. Thus, as soon as we were offered the opportunity of producing a new book on UV effects on aquatic ecosystems, we wondered how a new book on this subject could provide new insights or a different perspective, and perhaps the stimulus or inspiration for future research. In this book, we have attempted to bridge the gap between the environmental studies of UVR effects and the broader, traditional fields of ecology, oceanography and limnology. Our purpose has been to provide evidence to persuade a general ecologist that UV driven processes are relevant to aquatic ecosystems. But, at the same time, we wanted to adopt the point of view that UV is only one of several important ecological processes operating synchronously in the natural environment. If we have succeeded, the message from this book should be that the search for environmental UV effects must be framed within a wider ecological context.

What's in the book

The book is divided into five sections, which are intended to cover the most salient aspects of UV research. The introductory chapter provides an overview of the role of UVR in aquatic systems with a strong emphasis on the interaction between UVR and DOM. This interaction will be a recurrent subject in subsequent chapters, and reflects the contemporary perception of UV researchers about the key role played by DOM in controlling UV optics, and directly or indirectly regulating chemical and biological processes.

The Physics section provides information and discussions on global UVR climatology at the Earth's surface level and the factors controlling the transmit-tance of UVR through the atmosphere (Chapter 2), and the penetration into the water column (Chapter 3). The last chapter within this section (Chapter 4) introduces basic notions of mixing and advection and outlines the implications of vertical water motion for photochemical and photobiological processes.

The four chapters within the Chemistry section discuss the effects of UVR on biogeochemical cycles of various elements (Chapter 5), the photochemistry of DOM (Chapter 6), the photo-activated toxicity of several natural and anthropogenic substances (Chapter 7) and the environmental implications of photoinduced formation of reactive oxygen species (Chapter 8). Once again, the central role of DOM emerges as a unifying theme.

The fourth section focuses on individual and sub-individual effects and responses. The first chapter within this section (Chapter 9) reviews the effects of UVR on DNA, which has long been identified as one of the primary targets of UVR in biological systems. It is followed by a discussion of the main physiological photoprotective mechanisms in aquatic organisms (Chapter 10). Chapter 11 reviews the available literature on UVR effects on autotrophs, while Chapters 12 and 13 present two different and complementary perspectives on the effects of UVR on heterotrophs. This section ends with an extensive review on the role of sensory systems and behavioral responses to UVR (Chapter 14).

Three chapters within the last section address the effects of UVR from the community and ecosystem perspective that has been anticipated in the introductory section. Chapter 15 provides a thorough review of the effects of UVR on species interactions, including predation, competition, parasitism and diseases. Chapter 16 discusses the methods for reconstructing the radiation history of aquatic ecosystems and presents evidence for different UVR paleoclimates. Chapter 17 speculates on potential future UVR scenarios in a world that is experiencing several climatic changes from regional acidification to global warming and the also global depletion of stratospheric ozone.


Many people have contributed to make this book possible. First, we would like to thank all the authors for their time and commitment, and for their great disposition to help us out in every way. It has been a pleasure and a honour to work with them. We are deeply indebted to Ruben Sommaruga, who "volunteered" to work as Associate Editor for the chapters authored by either one of us. His help has been essential to assure the impartial review process of our chapters. Virginia Villafane helped us at every stage of the editing process and her inputs, comments and criticisms have been fundamental.

The contribution of the external reviewers deserves a separate paragraph. We cannot overemphasize how much this book benefited from the comments, opinions and generous suggestions made by the many reviewers: Maria Marta Bianchi, J. Piatt Bradbury, Howard Browman, Cynthia Carey, Ron Douglas,

Bruce Greenberg, David Hamilton, Bruce Hargreaves, Osmund Holm-Hansen, Ron Kiene, Peter Kiffney, Susanne Kratzer, George Losey, John Marra, Diane McKnight, Tim van Oijen, James T. Oris, Isabel Reche, David Schindler, Ralph Smith, Francesco Zaratti, and seven anonymous reviewers.

Finally, we would like to acknowledge the support of European Society of Photobiology, Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Universidad Nacional del Comahue and Fundación Playa Unión.

E. Walter Helbling Horacio Zagarese


Anastazia T. Banaszak

Unidad Académica Puerto Morelos


Apartado Postal 1152


Quintana Roo 77500 México.

Mario Blumthaler

Institute for Medical Physics Muellerstr. 44 A-6020 Innsbruck Austria

Peter Boelen

Department of Marine Biology Center for Ecological and Evolutionary Studies University of Groningen, P.O. Box 14, 1790 AA Haren The Netherlands

Anita G.J. Buma

Department of Marine Biology Center for Ecological and Evolutionary Studies University of Groningen P.O. Box 14 1790 A A Haren The Netherlands

Stephen A. Diamond

U.S. Environmental Protection Agency

Mid-Continent Ecology Division 6201 Congdon Boulevard Duluth, MN 55804 USA

David Fabacher

USGS Columbia Environmental Research Center 4200 New Haven Road Columbia, MO 65201 USA

Félix L. Figueroa

Departamento de Ecología Facultad de Ciencias Universidad de Málaga Campus Universitario de Teatinos s/n

E-29071 Málaga Spain

Bruce R. Ha r greaves

Lehigh University Department of Earth & Environmental Sciences 31 Williams Drive Bethlehem, PA 18015 USA

E. Walter Helbling

David J. Kieber

Estación de Fotobiología Playa

State University of New York


College of Environmental Science


and Forestry

Consejo Nacional de

Chemistry Department

Investigaciones Científicas y

1 Forestry Drive

Técnicas (CONICET)

Syracuse, NY 13210

Casilla de Correos No. 153


(9100) Trelew


Peter R. Leavitt


Limnology Laboratory

Dept. of Biology

Dag O. Hessen

University of Regina

Department of Biology


University of Oslo


P.O.Box 1027

Canada, S4S 0A2


0316 Oslo

Dina M. Leech


Department of Earth and

Environmental Sciences

Dominic A. Hodgson

Lehigh University

British Antarctic Survey

Bethlehem, PA 18015

Natural Environment Research



Present address:

High Cross

Biology Department

Madingley Road

Box 90338

Cambridge, CB3 OET

Duke University

United Kingdom

Durham, NC 27708


Wade H. Jeffrey

Center for Environmental

Edward E. Little

Diagnostics and Bioremediation

USGS Columbia Environmental

University of West Florida

Research Center

11000 University Parkway

4200 New Haven Road

Pensacola, FL 32514

Columbia, Missouri 65201



Sönke Johnsen

Donald P. Morris

Biology Department

Department of Earth &

Woods Hole Oceanographic

Environmental Sciences


Lehigh University

Woods Hole, MA 02543

31 Williams Drive


Bethlehem, PA 18015

Smithsonian Environmental Research Center P. O. Box 28 Edgewater MD 21037 USA

Christopher L. Osburn

Chemistry Division, Code 6115 Naval Research Laboratory 4555 Overlook Ave SW Washington, DC 20375 USA

Barrie M. Peake

Chemistry Department University of Otago Union Place Dunedin New Zealand

Reinhard Pienitz

Paleolimnology-Paleoecology Laboratory

Centre d'Études Nordiques Université Laval, Québec Québec

Canada, G1K 7P4

Norman M. Scully

Chemistry Department Center for Marine Science University of North Carolina at Wilmington

One Marvin K. Moss Lane Wilmington, NC 28409 USA

Ruben Sommaruga

Institute of Zoology and Limnology University of Innsbruck Technikerstr. 25 A-6020 Innsbruck Austria

Kristina Sundbäck

Department of Marine Botany

Göteborg University

Box 461

SE-405 30


Virginia E. Villafañe

Estación de Fotobiología Playa

Unión and

Consejo Nacional de

Investigaciones Científicas y

Técnicas (CONICET)

Casilla de Correos No. 153

(9100) Trelew



Ann R. Webb

Department for Physics University Manchester Institute for Science and Technology Manchester M60 1QD United Kingdom

Robert G. Wetzel

Department of Environmental Sciences and Engineering School of Public Health The University of North Carolina Chapel Hill, NC 27599-7431 USA

Craig E. Williamson

Department of Earth and Environmental Sciences 31 Williams Drive Lehigh University Bethlehem, PA 18015-3188 USA

Horacio £. Zagarese

Laboratorio de Fotobiología Centro Regional Universitario Bariloche

Universidad Nacional del Comahue

Unidad Postal Universidad

8400 Bariloche


Richard G. Zepp

U.S. Environmental Protection Agency

960 College Station Road Athens, GA 30605-2700 USA



Chapter 1

Solar radiation as an ecosystem modulator 3

Robert G. Wetzel


Chapter 2

UVR climatology 21

Mario Blumthaler and Ann R. Webb

Chapter 3

Water column optics and penetration of UVR 59

Bruce R. Hargreaves

Chapter 4

Modulation of UVR exposure and effects by vertical mixing and advection 107

Patrick J. Neale, E. Walter Helbling and Horacio E. Zagarese


Chapter 5

Solar UVR and aquatic carbon, nitrogen, sulfur and metals cycles 137

Richard G. Zepp

Chapter 6

Photochemistry of chromophoric dissolved organic matter in natural waters 185

Christopher L. Osburn and Donald P. Morris

Chapter 7

Photoactivated toxicity in aquatic environments 219

Stephen A. Diamond

Chapter 8

Reactive oxygen species in aquatic ecosystems 251

David J. Kieber, Barrie M. Peake and Norman M. Scully

Individual and Sub-indiviudal Effects and Responses

Chapter 9

UVR-induced DNA damage in aquatic organisms 291

Anita G.J. Buma, Peter Boelen and Wade H. Jeffrey

Chapter 10

Photoprotective physiological and biochemical responses of aquatic organisms 329

Anastazia T. Banaszak

Chapter 11

Photosynthesis in the aquatic environment as aifected by UVR 357

Virginia E. Villafane, Kristina Sundbâck, Félix L. Figueroa and E. Walter Helbling

Chapter 12

UVR and pelagic metazoans 399

Dag 0. Hessen

Chapter 13

UVR-induced injuries in freshwater vertebrates 431

Edward E. Little and David Fabacher

Chapter 14

Behavioral responses - UVR avoidance and vision 455

Dina M. Leech and Sônke Johnsen

Community and Ecosystem Perspectives

Chapter 15

UVR and its effects on species interactions 485

Ruben Sommaruga

Chapter 16

Past UVR environments and impacts on lakes 509

Peter R. Leavitt, Dominic A. Hodgson and Reinhard Pienitz

Chapter 17

UVR effects on aquatic ecosystems: a changing climate perspective 547

Craig E. Williamson and Horacio E. Zagarese

Subject Index

Abbreviations and symbols

O, quantum yield

Oa, apparent quantum yield a*cdom,/b chromophoric dissolved organic matter-specific optical absorption a*chu> chlorophyll-specific spectral absorption factor a*DOo dissolved organic carbon-specific absorption

6-4 PP, (6-4) pyrimidone photoproduct

8-oxoG, 8-hydroxyguanosine aCDOM, absorption coefficient of CDOM

ADP, adenosine di-phosphate

AOP, apparent optical properties

ATP, adenosine tri-phosphate

BAF, bioaccumulation factor

BCF, bioconcentration factor

BED, biologically effective dose

BEE, biologically effective exposure BEI, biologically effective irradiance BLP, biologically-available photoproduct BWF, biological weighting function

CA, carbonic anhydrase CAT, catalase

CCN, cloud condensation nuclei

CDOM, chromophoric dissolved organic matter chl-a, chlorophyll a

C02, carbon dioxide

CO, carbon monoxide

COS, carbonyl sulfide

CPD, cyclobutane pyrimidine dimer

CTM, chemical transport models

DIC, dissolved inorganic carbon

DMS, dimethyl sulfide

DMSP, dimethylsulfonium propionate

Dn, diadinoxanthin

DNA, desoxy ribonucleic acid

DOC, dissolved organic carbon

DOM, dissolved organic matter

DON, dissolved organic nitrogen

DOS, dissolved organic sulfur

Dt, diatoxanthin

DVM, diel vertical migration

Ed, downwelling irradiance

EEMS, excitation-emission matrix spectra

ENSO, El Nino-Southern Oscillation

Eph, euphotic zone (1% of surface PAR)

EPR, electron paramagnetic resonance spectroscopy

Eu, upwelling irradiance

FAD, flavin adenine dinucleotide

FP, fluorescent pigments

FWHM, full-width at half maximum

G3PDH, glyceraldehyde -3-phosphate dehydrogenase

GCM, general circulation models

GST, glutathione transferase

HMW, high molecular weight

HNF, heterotrophic nanoflagellates

HO*, hydroxyl radical

HOMO, highest occupied molecular orbital

HPLC, high-performance liquid chromatography

IOP, inherent optical properties

IR, infrared

Kd, diffuse attenuation coefficient for downwelling irradiance

Kov/, organic-water partitioning coefficient

Ku, diffuse attenuation coefficient for upwelling irradiance

LMW, low molecular weight

LUMO, lowest unoccupied molecular orbital

MAA, mycosporine-like amino acid

MCH, melanin-concentration hormone

MDR, mean damage ratio

MPB, microphytobenthos

NCDOM, non-chromophoric dissolved organic matter

NO, nitric oxide

NR, nitrate reductase

PAH, polycyclic aromatic hydrocarbons

PAM, pulse amplitude modulated (fluorescence)

PAR, photosynthetically active radiation (400-700 nm)

PER, photoenzymatic repair

POC, particulate organic carbon

POM, particulate organic matter

PS II, photosystem II

PUFAs, polyunsaturated fatty acids

PWF, phototoxicity weighting function

QFT, quantitative filter technique

QSAR, quantitative structure-activity relationship

RAF, radiation amplification factor

ROS, reactive oxygen species

RNA, ribonucleic acid

RPA, relative photodynamic activity

RUBISCO, ribulose-l,5-biphosphate carboxylase/oxygenase

SBB, single strand breaks

SOD, superoxide dismutase

SPE, solid phase extraction

SWF, spectral weighting function

SZA, solar zenith angle

TT, thymine dimers

UML, upper mixed layer

UV-A, ultraviolet radiation (315-400 nm)

UV-B, ultraviolet radiation (280-315 nm)

UV-C, ultraviolet radiation (200-280 nm)

UVeff, weighted UVR for photosynthesis inhibition

UVR, ultraviolet radiation

WSC, Weddell-Scotia confluence za, depth (m) of 1% of surface irradiance.

ZEph, depth of the euphotic zone

Z„%, percent attenuation depth

ZUML, depth of the upper mixed layer


• Solar radiation as an ecosystem modulator

Chapter 1

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