JOHN T. O. KIRK
Kirk Marine Optics
CAMBRIDGE UNIVERSITY PRESS
Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore,Sa˜o Paulo, Delhi, Dubai, Tokyo, Mexico City
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| Light and Photosynthesis in Aquatic Ecosystems 3rd Edition |
Third edition
Beginning systematically with the fundamentals, the fully updated third edition of
this popular graduate textbook provides an understanding of all the essential
elements of marine optics. It explains the key role of light as a major factor in
determining the operation and biological composition of aquatic ecosystems, and
its scope ranges from the physics of light transmission within water, through the
biochemistry and physiology of aquatic photosynthesis, to the ecological
relationships that depend on the underwater light climate. This book also provides
a valuable introduction to the remote sensing of the ocean from space, which is
now recognized to be of great environmental significance due to its direct
relevance to global warming.
An important resource for graduate courses on marine optics, aquatic
photosynthesis, or ocean remote sensing; and for aquatic scientists, both
oceanographers and limnologists.
john t.o. kirk began his research into ocean optics in the early 1970s in the
Division of Plant Industry of the Commonwealth Scientific & Industrial Research
Organization (CSIRO), Canberra, Australia, where he was a chief research
scientist, and continued it from 1997 in Kirk Marine Optics. He was awarded the
Australian Society for Limnology Medal (1981), and besides the two successful
previous editions of this book, has also co-authored The Plastids: Their Chemistry,
Structure, Growth and Inheritance (Elsevier, 1978), which became the standard text in its field.
Beyond his own scientific research interests, he has always been interested in the
broader implications of science for human existence, and has published a book on
this and other issues, Science and Certainty (CSIRO Publishing, 2007).
Preface to the third edition
Four things are required for plant growth: energy in the form of solar
radiation; inorganic carbon in the form of carbon dioxide or bicarbonate
ions; mineral nutrients; and water. Those plants which, in the course of
evolution, have remained in, or have returned to, the aquatic environment
have one major advantage over their terrestrial counterparts: namely,
that water – lack of which so often limits productivity in the terrestrial
biosphere – is for them present in abundance; but for this a price must be
paid. The medium – air – in which terrestrial plants carry out photosynthesis
offers, within the sort of depth that plant canopies occupy, no
significant obstacle to the penetration of light. The medium – water – in
which aquatic plants occur, in contrast, both absorbs and scatters light.
For the phytoplankton and the macrophytes in lakes and rivers, coastal
and oceanic waters, both the intensity and spectral quality of the light
vary markedly with depth. In all but the shallowest waters, light availability
is a limiting factor for primary production by the aquatic ecosystem.
The aquatic plants must compete for solar radiation not only with
each other (as terrestrial plants must also do), but also with all the other
light-absorbing components of the aquatic medium. This has led, in the
course of evolution, to the acquisition by each of the major groups of
algae of characteristic arrays of light-harvesting pigments that are of great
biochemical interest, and also of major significance for an understanding
both of the adaptation of the algae to their ecological niche and of the
phylogeny and taxonomy of the different algal groups. Nevertheless, in
spite of the evolution of specialized light-harvesting systems, the aquatic
medium removes so much of the incident light that aquatic ecosystems
are, broadly speaking, less productive than terrestrial ones.
Thus, the nature of the light climate is a major difference between
the terrestrial and the aquatic regions of the biosphere. Within the
radiation; inorganic carbon in the form of carbon dioxide or bicarbonate
ions; mineral nutrients; and water. Those plants which, in the course of
evolution, have remained in, or have returned to, the aquatic environment
have one major advantage over their terrestrial counterparts: namely,
that water – lack of which so often limits productivity in the terrestrial
biosphere – is for them present in abundance; but for this a price must be
paid. The medium – air – in which terrestrial plants carry out photosynthesis
offers, within the sort of depth that plant canopies occupy, no
significant obstacle to the penetration of light. The medium – water – in
which aquatic plants occur, in contrast, both absorbs and scatters light.
For the phytoplankton and the macrophytes in lakes and rivers, coastal
and oceanic waters, both the intensity and spectral quality of the light
vary markedly with depth. In all but the shallowest waters, light availability
is a limiting factor for primary production by the aquatic ecosystem.
The aquatic plants must compete for solar radiation not only with
each other (as terrestrial plants must also do), but also with all the other
light-absorbing components of the aquatic medium. This has led, in the
course of evolution, to the acquisition by each of the major groups of
algae of characteristic arrays of light-harvesting pigments that are of great
biochemical interest, and also of major significance for an understanding
both of the adaptation of the algae to their ecological niche and of the
phylogeny and taxonomy of the different algal groups. Nevertheless, in
spite of the evolution of specialized light-harvesting systems, the aquatic
medium removes so much of the incident light that aquatic ecosystems
are, broadly speaking, less productive than terrestrial ones.
Thus, the nature of the light climate is a major difference between
the terrestrial and the aquatic regions of the biosphere. Within the
underwater environment, light availability is of major importance in
determining how much plant growth there is, which kinds of plant predominate
and, indeed, which kinds of plants have evolved. It is not the
whole story – biotic factors, availability of inorganic carbon and mineral
nutrients, and temperature, all make their contribution – but it is a large
part of that story. This book is a study of light in the underwater environment
from the point of view of photosynthesis. It sets out to bring
together the physics of light transmission through the medium and capture
by the plants, the biochemistry of photosynthetic light-harvesting
systems, the physiology of the photosynthetic response of aquatic plants
to different kinds of light field, and the ecological relationships that
depend on the light climate. The book does not attempt to provide as
complete an account of the physical aspects of underwater light as the
major works by Jerlov (1976), Preisendorfer (1976) and Mobley (1994); it
is aimed at the limnologist and marine biologist rather than the physicist,
although physical oceanographers should find it of interest. Its intention
is to communicate a broad understanding of the significance of light as a
major factor determining the operation and biological composition of
aquatic ecosystems. It is hoped that it will be of value to practising
aquatic scientists, to university teachers who give courses in limnology
or marine science, and to postgraduate and honours students in these and
related disciplines.
Certain features of the organization of the book merit comment.
Although in some cases authors and dates are referred to explicitly, to
minimize interruptions to the text, references to published work are in
most cases indicated by the corresponding numbers in the complete
alphabetical reference list at the end of the book. Accompanying each
entry in the reference list is (are) the page number(s) where that paper or
book is referred to in the text. Although coverage of the field is, I believe,
representative, it is not intended to be encyclopaedic. The papers referred
to have been selected, not only on the grounds of their scientific importance,
but in large part on the basis of their usefulness as illustrative
examples for particular points that need to be made. Inevitably, therefore,
many equally important and relevant papers have had to be omitted from
consideration, especially in the very broad field of aquatic ecology. I have
therefore, where necessary, referred the reader to more specialized works
in which more comprehensive treatments of particular topics can be
found. Because its contribution to total aquatic primary production is
usually small I have not attempted to deal with bacterial photosynthesis,
complex and fascinating though it is.
The behaviour of sunlight in water, and the role that light plays in
controlling the productivity, and influencing the biological composition,
of aquatic ecosystems have been important areas of scientific study for
more than a century, and it was to meet the perceived need for a text
bringing together the physical and biological aspects of the subject, that
the first, and then second, editions of Light and Photosynthesis in Aquatic
Ecosystems were written. The book was well received, and is in use not
only by research workers but also in university courses. In the 27 years
since the first edition, interest in the topic has become even greater than it
was before. This may be partly attributed to concern about global
warming, and the realization that to understand the important role the
ocean plays in the global carbon cycle, we need to improve both our
understanding and our quantitative assessment of marine primary production.
An additional, but related, reason is the great interest that has been
aroused in the feasibility of remote sensing of oceanic primary productivity
from space. The potentialities were just becoming apparent with the
early Coastal Zone Color Scanner (CZCS) pictures when the first edition
was written. The continuing stream of further remote sensing information
in the ensuing years, as space agencies around the world have put new and
improved ocean scanners into orbit, enormously enlarging our understanding
of oceanic phytoplankton distribution, have made this a particularly
active and exciting field within oceanography. But the light flux that
is received from the ocean by the satellite-borne radiometers, and which
carries with it information about the composition of the water, originates in
fact as a part of the upwelling light flux within the ocean, which has
escaped through the surface into the atmosphere. To interpret the data
we therefore need to understand the underwater light field, and how its
characteristics are controlled by what is present within the aquatic medium.
In consequence of this sustained, even intensified, interest in underwater
light, there is a continued need for a suitable text, not only for
researchers, but also for use in university teaching. It is for this reason, the
first and second editions being out of print, that I have prepared a
completely revised version. Since marine bio-optics has been such an
active field, a vast amount of literature had to be digested, but as in the
earlier editions, I have tended to select specific papers mainly on the basis
of their usefulness as illustrative examples, and many other equally valuable
papers have had to be omitted from consideration.
In the 16 years since the second edition of this book appeared, interest
in this subject has, if anything, increased. While there has been an
acceleration, rather than a slackening in the rate of publication of new
research it must be said that this has been much more evident in certain
areas than in others. Remote sensing of ocean colour, and its use to arrive
at inferences about the composition and optical properties of, and primary
production going on within, ocean waters has been the standout
example of a very active field. A variety of new instruments for measuring
the optical properties of the water, and the underwater light field, have
been developed, and a number of these are described. So far as photosynthesis
itself is concerned, the most notable change has been the development
of instrumentation, together with the necessary accompanying
theoretical understanding, for in situ measurement of photosynthetic rate,
using chlorophyll a fluorescence. A great deal more is also now known
about carbon concentrating mechanisms in aquatic plants, and these
topics are discussed. The presumptive role of iron as a limiting factor
for primary production in large areas of the ocean has received a great
deal of attention in recent years, and current understanding is summarized.
Nevertheless, quite apart from these specific areas, there has been
across-the-board progress in all parts of the subject, no chapter remains
unchanged, and the reference list has increased in length by about 50%.
I would like to thank Dr Susan Blackburn, Professor D. Branton, Dr
M. Bristow, Mr S. Craig, Dr W. A. Hovis,Mr Ian Jameson, Dr S. Jeffrey,
Dr D. Kiefer, Professor V. Klemas, Professor L. Legendre, Dr Y. Lipkin,
Professor W. Nultsch, Mr D. Price, Professor R. C. Smith, Dr M. Vesk;
Biospherical Instruments Inc., who have provided original copies of
figures for reproduction in this work; and Mr F. X. Dunin and Dr
P. A. Tyler for unpublished data. I would like to thank Mr K. Lyon of
Orbital Sciences Corporation for providing illustrations of the SeaWiFS
scanner and spacecraft, and the SeaWiFS Project NASA/Goddard Space
Flight Center, for remote sensing images of the ocean.
John Kirk
Canberra
April 2010
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ISBN
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Copyright
| John T. O. Kirk 2011 |
Contents
Preface to the third edition page ix
PART I THE UNDERWATER LIGHT FIELD 1
1 Concepts of hydrologic optics 3
1.1 Introduction 3
1.2 The nature of light 3
1.3 The properties defining the radiation field 6
1.4 The inherent optical properties 14
1.5 Apparent and quasi-inherent optical
properties 21
1.6 Optical depth 24
1.7 Radiative transfer theory 24
2 Incident solar radiation 28
2.1 Solar radiation outside the atmosphere 28
2.2 Transmission of solar radiation through
the Earth’s atmosphere 30
2.3 Diurnal variation of solar irradiance 38
2.4 Variation of solar irradiance and insolation
with latitude and time of year 42
2.5 Transmission across the air–water interface 44
3 Absorption of light within the aquatic medium 50
3.1 The absorption process 50
3.2 The measurement of light absorption 53
3.3 The major light-absorbing components
of the aquatic system 61
3.4 Optical classification of natural waters 92
3.5 Contribution of the different components
of the aquatic medium to absorption of PAR 95
4 Scattering of light within the aquatic medium 98
4.1 The scattering process 98
4.2 Measurement of scattering 104
4.3 The scattering properties of natural waters 116
4.4 The scattering properties of phytoplankton 128
5 Characterizing the underwater light field 133
5.1 Irradiance 133
5.2 Scalar irradiance 143
5.3 Spectral distribution of irradiance 144
5.4 Radiance distribution 147
5.5 Modelling the underwater light field 149
6 The nature of the underwater light field 153
6.1 Downward irradiance – monochromatic 153
6.2 Spectral distribution of downward irradiance 159
6.3 Downward irradiance – PAR 159
6.4 Upward irradiance and radiance 168
6.5 Scalar irradiance 178
6.6 Angular distribution of the underwater light field 181
6.7 Dependence of properties of the field
on optical properties of the medium 188
6.8 Partial vertical attenuation coefficients 197
7 Remote sensing of the aquatic environment 199
7.1 The upward flux and its measurement 200
7.2 The emergent flux 215
7.3 Correction for atmospheric scattering
and solar elevation 218
7.4 Relation between remotely sensed reflectance
and the scattering/absorption ratio 225
7.5 Relation between remotely sensed reflectances
and water composition 228
PART II PHOTOSYNTHESIS IN THE AQUATIC ENVIRONMENT 263
8 The photosynthetic apparatus of aquatic plants 265
8.1 Chloroplasts 265
8.2 Membranes and particles 268
8.3 Photosynthetic pigment composition 275
8.4 Reaction centres and energy transfer 298
8.5 The overall photosynthetic process 300
9 Light capture by aquatic plants 308
9.1 Absorption spectra of photosynthetic systems 308
9.2 The package effect 311
9.3 Effects of variation in cell/colony size and shape 314
9.4 Rate of light absorption by aquatic plants 319
9.5 Effect of aquatic plants on the underwater light field 325
10 Photosynthesis as a function of the incident light 330
10.1 Measurement of photosynthetic rate in aquatic
ecosystems 330
10.2 Photosynthesis and light intensity 339
10.3 Efficiency of utilization of incident light energy 360
10.4 Photosynthesis and wavelength of incident light 380
11 Photosynthesis in the aquatic environment 388
11.1 Circulation and depth 388
11.2 Optical characteristics of the water 397
11.3 Other limiting factors 400
11.4 Temporal variation in photosynthesis 430
11.5 Photosynthetic yield per unit area 440
12 Ecological strategies 453
12.1 Aquatic plant distribution in relation to light quality 453
12.2 Ontogenetic adaptation – intensity 469
12.3 Ontogenetic adaptation – spectral quality 479
12.4 Ontogenetic adaptation – depth 488
12.5 Significance of ontogenetic adaptation of the
photosynthetic system 503
12.6 Rapid adaptation of the photosynthetic system 514
12.7 The microphytobenthos 528
12.8 Highly productive aquatic ecosystems 532
References and author index 539
Index to symbols 626
Index to organisms 628
Index to water bodies 632
Subject index 638
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