THE ROLE OF SOLAR ULTRAVIOLET RADIATION IN MARINE ECOSYSTEMS

NATO CONFERENCE SERIES

I Ecology II Systems Science III Human Factors IV Marine Sciences V Air-Sea Interactions VI Materials Science

IV MARINE SCIENCES

Volume 1 Marine Natural Products Chemistry edited by D. J. Faulkner and W. H. Fenical

Volume 2 Marine Organisms: Genetics. Ecology. and Evolution edited by Bruno Battaglia and John A. Beardmore

Volume 3 Spatial Pattern in Plankton Communities edited by John H. Steele

Volume 4 Fjord Oceanography edited by Howard J. Freeland. David M. Farmer. and Colin D. Levings

Volume 5 Bottom-interacting Ocean Acoustics edited by William A. Kuperman and Finn B. Jensen

Volume 6 Marine Slides and Other Mass Movements edited by Svend Saxov and J. K. Niewenhuis

Volume 7 The Role of Solar Ultraviolet Radiation in Marine Ecosystems edited by John Calkins

THE ROLE OF SOLAR ULTRAVIOLET RADIATION IN MARINE ECOSYSTEMS

Edited by John Calkins Albert B. Chandler Medical Center University of Kentucky Lexington, Kentucky

Published in cooperation with NATO Scientific Affairs Division

PLENUM PRESS· NEW YORK AND LONDON

Library of Congress Cataloging in Publication Data

Main entry under title:

The Role of solar ultraviolet radiation in marine ecosystems.

(NATO conference series. IV, Marine sciences; v. 7) "Proceedings of a NATO conference on the role of solar ultraviolet radiation in

marine ecosystems, held July 28-31, 1980, in Copenhagen, Denmark" - Verso t.p.

"Published in cooperation with NATO Scientific Affairs Division." Bibliography: p. Includes index. 1. Ultra-violet rays-Environmental aspects-Congresses. 2. Marine ecology­

Congresses. I. Calkins, John, 1926- . II. Series. OH543.6.R64 574.5'2636 82-3797 ISBN 978-1-4684-8135-8 ISBN 978-1-4684-8133-4 (eBook) AACR2 DOI 10.1007/978-1-4684-8133-4

Proceedings of a NATO conference on The Role of Solar Ultraviolet Radiation in Marine Ecosystems, held July 28 - 31, 1980, in Copenhagen, Denmark

© 1982 Plenum Press, New York

Soflcover reprint of the hardcover 1 st edition 1 982

A Division of Plenum Publishing Corporation 233 Spring Street, New York, N.Y. 10013

All rights reserved

No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming" recording, or otherwise, without written permission from the Publisher

DEDICATION

This monograph and the preceding NATO Advanced Research Institute, Copenhagen, Denmark, July 28-31, 1980, are dedicated to the two Danish scientists who have contributed so greatly to the understanding of the role of solar ultraviolet radiation in aquatic ecosystems: Professors

N. G. Jerlov, Institute of Physical Oceanography, University of Copenhagen, Denmark

and

E. Steemann Nielsen, Freshwater Biological Laboratory, University of Copenhagen, Denmark

Professors Jerlov and Steemann Nielsen are notable for the attention they have'given to the study of the properties and actions of the ultraviolet portion of the solar spectrum on aquatic systems when their con­temporary scientists gave this region of the spectrum only passing consideration.

The Organizing Committee, on behalf of the Advanced Research Institute and the authors of this monograph, wish to express our gratitude and respect for their pioneering work in this important area.

v

PARTICIPANTS

K. S. Baker, U.S.A. J. A. Barcelo, U.S.A. D. Berger, U.S.A. A. Bogenrieder, W. Germany A. M. Bullock, Scotland M. M. Caldwell, U.S.A. J. Calkins, U.S.A. J. Chavaudra, France T. P. Coohill, U.S.A. P. Cutchis, U.S.A. D. M. Damkaer, U.S.A. E. C. DeFabo, U.S.A. B. L. Diffey, UK A. Eisenstark, U.S.A. B. Goldberg, U.S.A. A.E.S. Green, U.S.A. K. R. Gundersen, Sweden D. Hader, W. Germany P. V. Hariharan, U.S.A. N. K. H~jerslev, Denmark J. R. Hunter, U.S.A. N. Jerlov, Denmark R. Klein, W. Germany G. Kullenberg, Denmark

vii

R. D. Ley, U.S.A. F. M. Luther, U.S.A. S. Malmberg, Iceland D. S. Nachtwey, U.S.A. D. R. Norris, U.S.A. M. J. Peak, U.S.A. M. Polne, U.S.A. P. Reynisson, Iceland F. S. Rowland, U.S.A. A. Ruhland, W. Germany C. S. Rupert, U.S.A. R. C. Smith, U.S.A. R. S. Stolarski, U.S.A. F. Stordal, Norway A. H. Teramura, U.S.A. M. Tevini, W. Germany R. M. Tyrrell, Brazil E. Wellman, W. Germany G. ~. Wells, U.S.A. R. C. Worrest, U.S.A. C. Yentsch, U.S.A. R. G. Zepp, U.S.A. S. Zigman, U.S.A.

PREFACE

The inspiration for this monograph derived from the realization that human technical capacity has become so great that we can, even without malice, substantially modify and damage the gigantic and remote outer limit of our planet, the stratosphere. Above the atmosphere of our ordinary experience, the stratosphere is a tenuous layer of gas, blocked from rapid exchange with the troposphere, some twenty kilometers above the surface of the earth, seldom reached by humans, and yet a fragile shell which shields life on earth from a band of solar radiation of demonstrable injurious potential. It is immediately obvious that if stratospheric ozone were reduced and consequently the intensity of solar ultraviolet radiation reaching the earth's surface were increased, then human skin cancer, known to be related to solar ultraviolet exposure, would also be increased. But how does one even begin to estimate the impact of changed solar ultraviolet radiation on such a diverse. interacting, and complex ecosystem as the oceans?

Studies which I conducted in Iceland focused on this question and were noted to the Marine Sciences Panel of the Scientific Affairs Committee of NATO by Professor Unnsteinn Stefansson, leading to a request to investigate the possibility of organizing a NATO sponsored Advanced Research Institute on this topic. An Organizing Committee was formed including myself and Drs. A.E.S. Green, Per Haldall, Raymond Smith, and Robert C. Worrest. Professor Halldall subsequently resigned from the Organizing Committee due to a scheduling conflict and Drs. Gunnar Kullenberg and Charles Yentsch agreed to serve on the Committee. Because of the leadership of Danish scientists in the area, Copenhagen was always considered the natural site for the meeting. Professor Kullenberg and the Institute of Physical Oceanography generously agreed to host the meeting, an intensive four-day meeting without

ix

x PREFACE

subdivision of the participants was planned. The grant application was completed, received approval and the arrangements £or the con£erence were made by corres­pondence or telephone, without an actual meeting o£ the Organizing Committee.

It should be noted that in addition to developing and conducting the meeting in Copenhagen, the Organizing Committee, Drs:

Alex E. S. Green, The University o£ Florida Gunnar Kullenberg, The University o£ Copenhagen Raymond Smith, Scripps Institute of Oceanography Robert Worrest, Oregon State University Charles Yentsch, Bigelow Labora~ory £or Ocean

Science

also agreed to serve as an Editorial Board £or the monograph, reviewing the manuscripts of all participants as well as writing numerous contributions to the mono­graph themselves.

I wish to express my gratitude to three people who contrib~ted very generously to the success o£ the con£erehce and the production o£ the monograph: Ms E. Hallden, Secretary at the Institute £or Physical Oceano­graphy was most help£ul with the arrangements in Copenhagen; Dr. Jeanne Barcelo served as Executive Secretary £or the con£erence and as Associate Editor in the production of the mono"graph. lowe a very special debt to my wi£e Ruth Calkins; she prepared the grant application, typed all correspondence, beginning with the Organizing Committee and later correspondence with the participants o£ the con£erence. She has typed the £inal monograph, o£ten under di££icult cir­cumstances; I have no doubt that she more than any other single person is responsible £or the issuance o£ this monograph and the completion o£ this enterprise.

John Calkins August, 1981

CONTENTS

ATMOSPHERE AND BASIC BIOLOGICAL DOSIMETRY

Preface to Section I

UV-B Reaching the Surface Alex E. S. Green and P. F. Schippnick

Possible Anthropogenic Influences on Stratospheric Ozone . .

F. Sherwood Rowland

Ozone Depletion Calculations . . . . . . Frederick M. Luther, Julius S. Chang, Donald J. Wuebbles and Joyce E. Penner

On the Latitudinal and Seasonal Dependence of

1

5

29

49

Ozone Perturbations . . . . . . . 73 Richard S. Stolarski

Middle Ultraviolet Irradiance at the Ocean Surface: Measurements and Models 79

Karen S. Baker, Raymond C. Smith and A.E.S. Green

The Effect of Perturbation of the Total Ozone Column Due to CFC on the Spectral Distribution of UV Fluxes and the Damaging UV Doses at the Ocean Surface: A Model Study . . . .. . . . . . . 93

Frode Stordal, ~ystein Hov and Ivar S. A. Isaksen

Multi-Wavelength Determination of Total Ozone and Ultraviolet Irradiance 109

Alex E. S. Green

xi

xii

Radiometric Measurements in the UV-B Region of Daylight . . . . . . . . . .

Bernard Goldberg

Photobiological Dosimetry of Environmental Ultraviolet Radiation ...... .

Claud S. Rupert

Action Spectra ........... . John Calkins and Jeanne A. Barcelo

Some Thoughts on UV Action Spectra .. Martyn M. Caldwell

Action Spectra and Their Role in Solar UV-B Studies . . . .

A. Eisenstark

Action Spectra: Emphasis Mammalian Cells Thomas P. Coohill

Dosage Units for Biologically Effective UV-B:

CONTENTS

121

131

143

151

157

161

A Recommendation . . . . . . . . . . . . 167 Martyn M. Caldwell

Measuring Devices and Dosage Units. . . . . 169 John Calkins

The Sunburn UV Network and its Applicability for Biological Predictions . . . . . . . . 181

Daniel Berger

Description and Application of a Personal Dosimeter for Measuring Exposure to Natural Ultraviolet Radiation 193

Brian Diffey, Tony Davis, and Ian Magnus

Nomograms for Biologically Effective UV 205 David M. Damkaer and Douglas B. Dey

A Formula for Comparing Annual Damaging Ultraviolet (DUV) Radiation Doses at Tropical and Mid-Latitude Sites

Pythagoras Cutchis 213

CONTENTS

THE HYDROSPHERE

Preface to Section II ...... .

Spectral Irradiance Penetration in Natural Waters . . . . . . . . . .

Karen S. Baker and Raymond C. Smith

A Method for the Estimation of the Penetration of Biologically Injurious Solar Ultra-

xiii

229

233

violet Radiation into Natural Waters . . 247 John Calkins

Yellow Substance in the Sea N. K. Hpjerslev

Note on the Role of to Effects of Environment

Gunnar Kullenberg

Vertical Mixing in Relation UV Radiation on the Marine

Photochemical Transformations Induced by Solar

263

283

Ultraviolet Radiation in Marine Ecosystems 293 Richard G. Zepp

Penetration of Solar UV-B into Waters off Iceland . . . . . . . . . . .

John Calkins and Thorunn Thordard6ttir

THE BIOSPHERE

Preface to Section III . . .

Lethal Effects on Biological Systems Caused by Solar Ultraviolet Light: Molecular

309

321

Considerations . . . . . . . . . . . . . 325 Meyrick J. Peak and Jennifer G. Peak

DNA Damage of the 5,6-Dihydroxydihydrothymine Type Induced by Solar UV-B in Human Cells 337

P. V. Hariharan and E. R. Blazek

Mechanisms of Actions of Longwave-UV on Marine Organisms ..... . 347

Seymour Zigman

xiv

Survival of the Brine Shrimp, Artemia Salina, After Exposure to 290-NM Ultraviolet Radiation, with and without Maximum Photoreactivation ..... .

Meyrick J. Peak and H. E. Kubitschek

Photoreactivation: Mammalian Cells Thomas P. Coohill

The Amelioration of UV-B Effects on Productivity

CONTENTS

357

by Visible Radiation. . . . . . . . . .. 367 Alan H. Teramura

Error Prone Repair-·Emphasi s Weigle Reacti vati on 383 Thomas P. Coohill

Sensitivity to UV-B Irradiation as Related to Bacterial Life Cycles . . . . 389

A. Eisenstark

Pigment Dispersion by Light in the Melanophores of the Fiddler Crab . . . . . .. 393

Thomas P. Coohill

Yellow Lens Pigment: An Adaptation for Activity in Bright Sunlight . . . . . . . . . . .. 399

Seymour Zigman

Photomovement of Aquatic Organisms in Response to Solar UV . . . . . . . . . . . 407

Jeanne A. Barcelo

Movement Reactions of Blue-Green Algae in Response to Their Photoenvironment .

Donat-Peter Hader 411

Short-Term Responses of Some Planktonic Crustacea Exposed to Enhanced UV-B Radiation. . . . . 417

David M. Damkaer and Douglas B. Dey

Review of Literature Concerning the Impact of UV-B Radiation Upon Marine Organisms 429

Robert C. Worrest

Assessment of Effects of UV Radiation on Marine Fish Larvae . . . . . . . .. 459

John R. Hunter, Sandor E. Kaupp and JohnH. Taylor

CONTENTS

The Effect of UV-B Irradiation on the Integument of the Marine Flatfish Pleuronectes

xv

Platessa 1. . . . . . . . . . . . .. 499 Alistair M. Bullock

Assessment of the Influence of Enhanced UV-B on Marine Primary Productivity ..

Raymond C. Smith and Karen S. Baker

Modeling Light Loss Versus UV-B Increase for Organisms Which Control Their Vertical Position in the Water Column .....

John Calkins

Effects of UV-B Radiation on Thalassiosira

509

.539

Pseudonana: A Preliminary Study . . 543 M. L. Geiger, D. R. Norris, J. H. Blatt and R. D. Petrilla

The Effects of Ultraviolet Irradiation on Photosynthesis by Ruppia Maritima 1. (Widgeon Grass) . . . . .

Gary N. Wells and D. S. Nachtwey

Effects of UV Radiation on Photosynthesis of

555

Natural Populations of Phytoplankton 563 C. W. Modert, D. R. Norris, J. H. Blatt and R. D. Petrilla

The Effect of High Intensity U.V. Radiation on Benthic Marine Algae . . . . . . . . . . 573

M. Polne and A. Gibor

The Effects of UV-B Irradiation on Higher Plants 581 M. Tevini, W. Iwanzik and U. Thoma

Preliminary Results Regarding the Spectral Efficiency of UV on the Depression of Photosynthesis in Higher Plants 617

Arno Bogenrieder and Richard Klein

Possible Errors in Photosynthetic Measurements Arising from the Use of UV-Absorbing Cuvettes: Some Examples in Higher Plants 621

Arno Bogenrieder and Richard Klein

xvi CONTENTS

Biological Interactions Between Wavelengths in the Solar-UV Range: Implications for the Predictive Value of Action Spectra Measurements . . . . . . . . . . . . . .. 629

Rex M. Tyrrell

Does Solar UV Influence the Competitive Relationship in Higher Plants? . .

Arno Bogenrieder and Richard Klein

A Model to Evaluate the Impact of Cyclically

64-1

Recurring Hazards on Selected Populations 651 John Calkins

Solar UV Radiation as a Selective Force in the Evolution of Terrestrial Plant Life 663

Martyn M. Caldwell

Atmospheric Evolution and UV-B Radiation . . . . . 677 Richard S. Stolarski

Some Considerations on the Ecological and Evolutionary Effects of Solar UV . .

John Calkins

The Attenuation of Light by Marine Phytoplankton with Specific Reference to the Absorption

685

of Near-UV Radiation . . . . . . . . . 691 Charles S. Yentsch and Clarice M. Yentsch

Possible Influences of Solar UV Radiation in the Evolution of Marine Zooplankton 701

David M. Damkaer

List of Contributors 707 Index 713