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The Microbiomes of Humans, Animals, Plants,
and the Environment 3

Lucas J. Stal
Mariana Silvia Cretoiu Editors

The Marine
Microbiome
Second Edition
The Microbiomes of Humans, Animals,
Plants, and the Environment
Volume 3
This series covers microbiome topics from all natural habitats. Microbiome research
is a vibrant field of science that offers a new perspective on Microbiology with a
more comprehensive view on different microorganisms (microbiota) living and
working together as a community (microbiome). Even though microbial
communities in the environment have long been examined, this scientific movement
also follows the increasing interest in microbiomes from humans, animals and
plants. First and foremost, microbiome research tries to unravel how individual
species within the community influence and communicate with each other. Addi-
tionally, scientists explore the delicate relationship between a microbiome and its
habitat, as small changes in either, can have a profound impact on the other. With
individual research volumes, this series reflects the vast diversity of Microbiomes
and highlights the impact of this field in Microbiology.

More information about this series at https://link.springer.com/bookseries/16462

Lucas J. Stal • Mariana Silvia Cretoiu
Editors

The Marine Microbiome

Second Edition
Editors
Lucas J. Stal Mariana Silvia Cretoiu
Department of Freshwater and Marine University of Utrecht
Ecology – IBED Utrecht, The Netherlands
University of Amsterdam
Amsterdam, The Netherlands

ISSN 2662-611X ISSN 2662-6128 (electronic)

The Microbiomes of Humans, Animals, Plants, and the Environment
ISBN 978-3-030-90382-4 ISBN 978-3-030-90383-1 (eBook)
https://doi.org/10.1007/978-3-030-90383-1

# The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland
AG 2016, 2022
This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher, whether
the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of
illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and
transmission or information storage and retrieval, electronic adaptation, computer software, or by
similar or dissimilar methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication
does not imply, even in the absence of a specific statement, that such names are exempt from the relevant
protective laws and regulations and therefore free for general use.
The publisher, the authors, and the editors are safe to assume that the advice and information in this
book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or
the editors give a warranty, expressed or implied, with respect to the material contained herein or for any
errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional
claims in published maps and institutional affiliations.

This Springer imprint is published by the registered company Springer Nature Switzerland AG.
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
Dedicated to
Prof. dr. Wolfgang Elisabeth Krumbein
Geomicrobiologist
1937–2021
In loving memory
Foreword

Marine microbiology has been a latecomer to the field of microbial ecology. But the
last three decades have been enough to revert the situation. The ocean is now
arguably the best known major microbial ecosystem on Earth. It was to be expected
that its physical structure at first sight is much more amenable to sampling and
interpretation than the richly micro-structured habitats like sediments or soil, not to
mention animal microbiomes. The ocean is a single gigantic lake that is
homogenized by the equally colossal global circulation. It has relatively constant
and largely moderate conditions (apart from extreme oligotrophy, for exceptions see
Chap. 14). On the other hand, it is also the largest, oldest and likely most critical
ecosystem for the environmental health of the planet. It has also been the least
impacted by the arrival of multicellular plants and animals ca. 500 million years ago.
It is possible to envisage vast marine areas in which the conditions have changed
little after the great oxygenation event ca. 2.2 billion years ago. Only in the last
hundred years has anthropic impact started to be noticed in the Pacific and Atlantic
central gyres that remained nearly pristine until the arrival of intensive whaling by
the mid-nineteenth century.
Microbiology is now at a crossroads or, if you wish, a new beginning. It turns out
that the fathers of microbiology were lucky to be able to discover the causative
agents of the major infectious diseases of their time. Human pathogens tend to be
copiotrophs well suited for growth in laboratory pure cultures, but most microbes are
not, and this includes most bacteria, archaea and protists that live in the ocean. We
know that because now we can sequence their genomes directly from the environ-
ment. However, the new microbiology that arose from nucleic acid sequencing is not
without limitations and drawbacks. First and foremost, the capability of annotating
genes and genomes (that is inferring function from sequence) is very unsatisfactory
and largely based on a few model organisms very distant from their wild
counterparts. Too little effort is invested in detecting new functions and too many
mystifying sequences added to the humongous databases. What is worse, we still
miss an evolutionary model that would consider the major impact of the pangenome
of prokaryotic species. The key role played by the pangenome dynamics in the
evolution of microbes and their ecology is yet to be fully incorporated in models of
evolutionary biology or ecosystem functioning.
There is also an overall lack of finesse in microbiome studies that often gravitate
over scale (big data) rather than detailed analysis of individual depth profiles, time
vii
viii Foreword

series or specific microbes. To compound it all, the classification (nomenclature) of

microbes has suffered from indiscriminate abuse by major rearrangements based on
relationships, at the sequence level, of ribosomal components without contemplating
that a microbe is much more than the pedigree of its ribosomes (and that is assuming
that ribosomal RNAs or proteins really provide a reliable relationship). It is often
unclear what the names mean in terms of the genome make-up and the biology of the
microbes.
At this point in time, Lucas Stal and Silvia Cretoiu have undertaken the task of
publishing a book (this) that updates the field and aggregates a large amount of
relevant information; for that they have managed to gather a remarkable set of
distinguished authors who have collectively done an amazing job of using the
‘tsunami’ of data generated by high-throughput sequencing technologies to describe
most of relevant topics in the field, covering from population genomics to biogeog-
raphy. They have introduced novel perspectives like the micro-seascape ‘vast
expanses of extremely dilute background seawater punctuated by rich hotspots of
dissolved and particulate nutrient resources’ (from Chap. 2). Everyone knows I am
not an enthusiast of review books, although I believe they are certainly useful for
newcomers such as PhD or master’s students. The speed at which science is
progressing these days coupled with the time required to collect and review chapters
detracts from their usefulness. However, some books have the power to change a
field. I hope this book will help to move marine microbiology into the next frontier in
which a better understanding of microbes (largely, but not only, derived from
genomics) will allow a more profound understanding of the oldest and largest
microbiome on Earth.

Universitat Miguel Hernández, Francisco Rodriguez Valera

Sant Joan d’Alacant, Spain
Preface

In 2015, Springer invited us to edit a book on marine microbiology. At the time, we

were leading the project ‘MaCuMBA’ (‘Marine Microorganisms: Cultivation
Methods for Improving their Biotechnological Applications’) with a large consor-
tium of European marine microbiologists and with funding of the European Com-
mission. We decided to produce the book with contributions of MaCuMBA
consortium members and entitled it The Marine Microbiome (with the subtitle An
Untapped Source of Biodiversity and Biotechnological Potential). We thought that
with this title we emphasize the importance of considering the total of
microorganisms in the ocean and its adjacent seas, bays and estuaries as an entity,
and that this entity plays a critical role in the functioning of the marine ecosystem,
and that it would stimulate its research. The book appeared in 2016 (Stal and Cretoiu
2016) and was successful, and indeed, the research on the marine microbiome
enjoyed more attention since (see below). The success of The Marine Microbiome
was the reason for Springer to inquire with us already in 2019, whether we would be
willing to edit a second edition. Meanwhile, Springer launched a book series The
Microbiomes of Humans, Animals, Plants, and the Environment and the second
edition of The Marine Microbiome fits perfectly in that series. While in our opinion
the first edition is still recent and quite up-to-date, we decided that the second edition
would need substantial new material, and therefore, only a few of the chapters of the
first edition remained and were rewritten and updated. Therefore, the second edition
of The Marine Microbiome might as well be considered The Marine Microbiome
Part 2 being an important addition to the first.
Most of the authors of this volume agreed to their contributions in late 2019 and
early 2020, not knowing the impact that the COVID-19 pandemic would have on
their work routine. Many authors with young children were faced with closed
schools and while mainly working from home had to divide their attention to their
work as well as to the assistance of home education of their children. Also, their
work and work load changed and from one day to another routine teaching, research
and administration had to be completely reorganized in a way never done before.
Meanwhile we, the editors, were pressing for delivery of the book chapters. We are
therefore really grateful to all contributors for the great chapters that they wrote for
this book, and we apologize to those authors who met the original deadline and had
to wait much longer to see their work in print than originally anticipated.

ix
x Preface

We would also like to take this preface as an opportunity to exchange our

thoughts about the title of this book (and of the previous edition) and about the
use of the concept ‘microbiome’. We used it in the first edition more or less to
replace the term ‘marine microbiology’, and it seemed to become fashionable to use
the term ‘microbiome’ in a variety of situations in which microbiologists want to
describe the whole of all microorganisms in a certain ‘biome’. Now, although we
kept the term ‘microbiome’ as it seems that it has become commonplace in microbi-
ology, we feel that it is semantically incorrectly used.
Let us consider the term ‘microorganism’. ‘Organism’ is defined as ‘a living
being or entity adapted for living by means of organs separate in function but
dependent on one another; any living being or its material structure; any complete
whole which by the integration, interaction and mutual dependence of its parts is
comparable to a living being’. And ‘life’ is the state of an organism characterized by
certain processes or abilities that include metabolism, growth, reproduction and
response. ‘Micro-‘ as a prefix means small or minute, or a millionth of (e.g. a
metre). Therefore, ‘microorganism’ is a small, minute organism. These definitions
were obtained from the Webster’s Dictionary of the English language.
Now, ‘biome’ is defined as ‘a large community of organisms having a peculiar
form of vegetation and characteristic animals’ and usually covers large areas. This
term was coined by Frederic E. Clements in his lecture ‘The development and
structure of biotic communities’ at the Ecological Society of America in 1916 and
published in 2017 in the Journal of Ecology (p. 120–121). He wrote:

The biotic community is regarded as an organic unit comprising all the species of plants and
animals at home in a particular habitat. While plants are regarded as exerting the dominant
influence in the community, it is recognized that this rôle may sometimes be taken by the
animals. The biotic community, or biome, is fundamentally controlled by the habitat, and
exhibits a corresponding development and structure. In its development the biotic formation
reacts upon the habitat, and thus produces a succession of biomes, comparable in practically
all essentials to the succession of plant communities. Every such succession, or biosere,
terminates regularly in a climax. The bioseres of each climax are either primary or second-
ary, and these may be further distinguished as hydroseres, xeroseres, etc.

A ‘microbiome’ would therefore mean a small or minute form of a biome which is

in fact a contradiction in terms. This is not what has become the meaning of it. It has
unfortunately introduced more and unnecessary ‘jargon’ in scientific language—
jargon that is rarely precisely defined and increases confusion among ourselves but
even more to society for which our scientific research should be relevant. Now that
we seem to be stuck with ‘microbiome’, it may be good to briefly review its
definition.
Berg et al. (2020) adopted the definition of Whipps and his colleagues (1988) as
‘a characteristic microbial community’ in a ‘reasonably well-defined habitat which
has distinct physicochemical properties’ as their ‘theatre of activity’ (this book
chapter of Whipps et al. (1988) is cited in Berg et al. (2020), and these authors
made some explanatory comments to it).
Preface xi

BioConcepts (www.biological-concepts.com) gives two definitions of

‘microbiome’:

1. A population of microorganisms inhabiting a specific environment: a microbial

community or ecosystem. This definition has been attributed to Mohr (1952) who
mentioned the term microbiome without, however, giving an explicit definition.
2. The collective genomes of all the microorganisms inhabiting a specific
environment.
The latter was proposed by the 1958 Nobel laureate Joshua Lederberg (2001)
(cited in Hooper and Gordon 2001) (citation: ‘It includes Lederberg’s own recent
coinage of microbiome, to signify the ecological community of commensal,
symbiotic, and pathogenic microorganisms that literally share our body space
and have all but ignored as determinants of health and disease.’) Hooper and
Gordon (2001) citing Lederberg: ‘microbiome’ to describe the collective genome
of our indigenous microbes (microflora).

Shade and Handelsman (2012) introduced the term ‘core microbiome’ in the hope
to find microorganisms common to any biome and hypothesized that it would fulfil
the basic and fundamental roles required for any ecosystem to function. This is an
interesting idea that awaits the discovery and description of examples that fit this
hypothesis. Shade and Handelsman (2012) defined microbiome as ‘an assemblage of
microorganisms existing in or associated with a habitat; includes active and
interacting member as well transient or inactive members’, definition that was
based on Lederberg and McCray (2001). They gave as examples the human
microbiome, earth microbiome, Lake Erie microbiome and soil microbiome (the
marine or ocean microbiome had not yet been discovered; see Fig. 2). Remarkably,
Shade and Handelsman (2012) define biome as ‘the world’s major ecosystems,
defined by temperature gradients in latitude and altitude, precipitation and season-
ality’ and cite Walter and Box (1976). Examples given by them are subtropical,
Mediterranean and polar. It is difficult to see a microbiome with this definition.
The Earth microbiome in fact comprises all microorganisms on Earth and all three
domains of life (bacteria, archaea and eukarya), and we also like to include viruses
and other forms of genetic information that interact with cells by keeping some sort
of functional equilibrium between the different organisms, exchanging genetic
information and generating diversity.
A search in the Web of Science (WoS) using the term ‘microbiome’ in the title of
scientific publications tells us that it appeared for the first time in 2006 (3 hits)
culminating to more than 3500 hits in 2020 (Fig. 1). The combination ‘marine’ and
‘microbiome’ gave the first 2 hits in 2012 and 12 in 2020 (Fig. 2). Just to give a little
more insight in the popularity of microbiome research, a WoS search on 19 May
2021 with ‘microbiome’ in the title returns 16,138 hits. 28% and 10% of those are
with respectively ‘gut’ or ‘human’ in the title and 3.3 and 0.3% with the term ‘soil’ or
‘rhizosphere’ and ‘marine’, respectively. These numbers show that microbiome
research becomes increasingly popular but that most of it focuses on humans and
occasionally other animal model systems and that the second-most investigated
xii Preface

Fig. 1 Hits in the Web of

Science with keyword
‘microbiome’ in the title of
publications by year of
publication

microbiomes are connected to (crop) plants and food production. The microbiome of
the ocean, arguably the largest ecosystem and crucial for the Earth’s climate and
food production, has still received little attention.
The Human Microbiome Project (2008–2012) was dominating microbiome
research and until 2010 only human (gut) microbiome and few other animal gut
microbiomes returned as hits in the title of a WoS search. In 2010, the Earth
Microbiome was launched (Gilbert et al. 2010). Microbiome research stays focused
on the human microbiome up to today. Other microbiomes studied are mostly in
other animal models and to some extent plants or a rare environment or habitat
turned up in the title together with the term microbiome. In 2013, the scientific
journal Microbiome appeared next to a few other even more specialized journals as
well as Environmental Microbiome (2019).
This book uses the term ‘microbiome’ as a description of all microorganisms in a
certain biome. We consider the ocean and its adjacent seas, bays and estuaries as the
‘marine biome’ and refer to the total of microorganisms in it as the ‘marine
microbiome’.
We hope and expect that this book along with the first edition (Part 1) will inspire
microbiologists to speed up the study of marine microorganisms and increase the
knowledge of their diversity and function because it is urgently needed to understand
the changes our planet is facing today and in the coming decennia. Fortunately, since
2016 important new initiatives have been launched such as the Atlantic Ocean
Research Alliance (AORA) that published the ‘Marine Microbiome Roadmap
2020’, the United Nations ‘Decade of Ocean Science for Sustainable Development’
and the EU Ocean Literacy projects ‘Sea Change’ and ‘ResponSEAble’ (see
Chap. 18). We are confident that this book will crank up marine microbiological
research.
One of us (LJS) retired in 2017 but remained as emeritus professor ‘Marine
Microbiology’ affiliated to the University of Amsterdam. This book may be his last
product of an almost 45-year career in marine microbiology, and Chap. 1 of this
book will be his last scientific paper. LJS would like to take this opportunity (again)
to thank all his friends and colleagues in science and particularly in marine
Preface xiii

Fig. 2 Hits in the Web of

Science with keywords
‘microbiome’ AND ‘marine’
in the title of publications by
year of publication

microbiology for the wonderful time he had, accomplishing scientific work and
contributing to the world’s knowledge on the unseen majority.
Finally, we would like to thank the project coordinators of Springer for their help
and encouragement and for giving us the opportunity to publish this second edition
of The Marine Microbiome. We particularly thank Mr. Bharat Sabnani, Mrs. Andrea
Schlitzberger and Mr. Markus Spaeth. We also like to thank Professor Francisco
Eduardo Rodriguez Valera for his kind foreword to this book.

Amsterdam, the Netherlands Lucas J. Stal

Utrecht, the Netherlands Mariana Silvia Cretoiu

References

Berg G, Rybakova D, Fischer D, Cernava T, Champomier Vergès M-C, Charles T,

Chen X, Cocolin L, Eversole K, Herrero Coral G, Kazou M, Kinkel L, Lange L,
Lima N, Loy A, Macklin JA, Maguin E, Mauchline T, McClure R, Mitter B,
Ryan M, Sarand I, Smidt H, Schelke B, Roume H, Kiran GS, Selvin J, Soares
Correa de Souza R, van Overbeek L, Singh BK, Wagner M, Walsh A,
Sessitsch A, Schloter M (2020) Microbiome definition re-visited: old concepts
and new challenges. Microbiome 8:103
Gilbert JA, Meyer F, Jansson J, Gordon J, Pace N, Tiedje J, Ley R, Fierer N, Field D,
Kyrpides N, Glöckner F-O, Klenk H-P, Wommack KE, Glass E, Docherty K,
Gallery R, Stevens R, Knight R (2010) The Earth microbiome project: meeting
report of the “1st EMP meeting on sample selection and acquisition” at Argonne
National Laboratory October 6th 2010. Stand Genome Sci 3:249–253
Hooper LV, Gordon JI (2001) Commensal host-bacterial relationships in the gut.
Science 292:1115–1118
Lederberg J, McCray AT (2001) ‘Ome sweet ‘omics - A genealogical treasury of
words. The Scientist 8:April 2
xiv Preface

Mohr JL (1952) Protozoa as indicators of pollution. Sci Month 74:7–9

Shade A, Handelsman J (2012) Beyond the Venn diagram: the hunt for a core
microbiome. Environ Microbiol 14:4–12
Stal LJ, Cretoiu MS (2016) The marine microbiome. An untapped source of
biodiversity and biotechnological potential. Springer International Publishing,
Cham, 498 p
Contents

1 A Sea of Microbes: What’s So Special about Marine

Microbiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Lucas J. Stal

Part I Diversity and Evolution of Marine Microorganisms

2 Survival in a Sea of Gradients: Bacterial and Archaeal
Foraging in a Heterogeneous Ocean . . . . . . . . . . . . . . . . . . . . . . . . 47
Estelle E. Clerc, Jean-Baptiste Raina, François J. Peaudecerf,
Justin R. Seymour, and Roman Stocker
3 Marine Cyanobacteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
Frédéric Partensky, Wolfgang R. Hess, and Laurence Garczarek
4 Marine Protists: A Hitchhiker’s Guide to their Role
in the Marine Microbiome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
Charles Bachy, Elisabeth Hehenberger, Yu-Chen Ling,
David M. Needham, Jan Strauss, Susanne Wilken,
and Alexandra Z. Worden
5 Marine Fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243
Gaëtan Burgaud, Virginia Edgcomb, Brandon T. Hassett,
Abhishek Kumar, Wei Li, Paraskevi Mara, Xuefeng Peng,
Aurélie Philippe, Pradeep Phule, Soizic Prado, Maxence Quéméner,
and Catherine Roullier
6 Marine Viruses: Agents of Chaos, Promoters of Order . . . . . . . . . . 297
Marcos Mateus
7 Evolutionary Genomics of Marine Bacteria and Archaea . . . . . . . . 327
Carolina A. Martinez-Gutierrez and Frank O. Aylward

Part II Marine Habitats

8 Towards a Global Perspective of the Marine Microbiome . . . . . . . . 357
Silvia G. Acinas, Marta Sebastián, and Isabel Ferrera

xv
xvi Contents

9 The Pelagic Light-Dependent Microbiome . . . . . . . . . . . . . . . . . . . . 395

Julie LaRoche and Brent M. Robicheau
10 Microbial Inhabitants of the Dark Ocean . . . . . . . . . . . . . . . . . . . . 425
Federico Baltar and Gerhard J. Herndl
11 The Subsurface and Oceanic Crust Prokaryotes . . . . . . . . . . . . . . . 461
Mohamed Jebbar
12 The Microbiome of Coastal Sediments . . . . . . . . . . . . . . . . . . . . . . 479
Graham J. C. Underwood, Alex J. Dumbrell, Terry J. McGenity,
Boyd A. McKew, and Corinne Whitby
13 Symbiosis in the Ocean Microbiome . . . . . . . . . . . . . . . . . . . . . . . . 535
Jonathan P. Zehr and David A. Caron
14 Marine Extreme Habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 579
Maria Pachiadaki and Virginia Edgcomb

Part III Marine Microbiome from Genomes to Phenomes:

Biogeochemical Cycles, Networks, Fluxes, and Interaction
15 Marine Biogeochemical Cycles . . . . . . . . . . . . . . . . . . . . . . . . . . . . 623
Samantha B. Joye, Marshall W. Bowles, and Kai Ziervogel
16 A Holistic Approach for Understanding the Role of
Microorganisms in Marine Ecosystems . . . . . . . . . . . . . . . . . . . . . . 673
Gerard Muyzer and Mariana Silvia Cretoiu
17 The Hidden Treasure: Marine Microbiome as Repository
of Bioactive Compounds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 693
Bathini Thissera, Ahmed M. Sayed, Hossam M. Hassan,
Usama R. Abdelmohsen, Rainer Ebel, Marcel Jaspars,
and Mostafa E. Rateb
18 Ocean Restoration and the Strategic Plan of the Marine
Microbiome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 731
Marieke Reuver, Jane Maher, and Annette M. Wilson
A Sea of Microbes: What’s So Special about
Marine Microbiology 1
Lucas J. Stal

Abstract

This chapter investigates what justifies marine microbiology as a discipline in its

own right. Do marine microorganisms really exist? And if so, what distinguishes
them from freshwater- or terrestrial microorganisms, or from microorganisms
living in any other specialized habitat? The marine environment—particularly the
ocean—is the largest continuous habitat on Earth. This makes the ocean different
in terms of scale and sharing space and nutrients, as well as in terms of the
distribution, dispersal, and encounters of microorganisms that inhabit it. The
ocean comprises a large variety of confined sub-habitats. It has an impact on
the function of the planet Earth as a whole. A critical property of seawater is that it
contains a large amount of salt and that this requires microorganisms that live in it
are able to adjust their osmotic pressure. The marine microbiome is composed of
the three domains of life: bacteria, archaea, and eukarya, as well as viruses, all of
which in dazzling numbers and diversity. All of the known microbial lineages are
represented and many are exclusively found in the ocean and there is little doubt
that life originated in the ocean.

Keywords
Marine microbiology · Milestones of marine microbiology · Nitrogen fixation ·
Ocean · Redfield ratio · Salinity

L. J. Stal (*)
Department of Freshwater and Marine Ecology – IBED, University of Amsterdam, Amsterdam,
The Netherlands

# The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 1

L. J. Stal, M. S. Cretoiu (eds.), The Marine Microbiome, The Microbiomes of
Humans, Animals, Plants, and the Environment 3,
https://doi.org/10.1007/978-3-030-90383-1_1
2 L. J. Stal

1.1 Introduction

Nowhere else where life is possible, probably in no other place in the Universe except
another ocean, are so many conditions so stable and so enduring.

Baas Becking (1934) put this quote from Henderson’s book “The Fitness of the
Environment” (Henderson 1913) above Chap. 9 “De Zee” (“The Sea”) of his book
“Geobiologie of inleiding tot de milieukunde” (Fig. 1.1). But immediately, Baas
Becking put this quote in perspective by stating that it might refer only to the
physicochemical characteristics, because the ocean is considered to be stable
because it is composed mainly of water. He continues by stating that when viewing
it from a biological perspective no environment has such a diversity as the ocean
with its adjacent seas, bays, estuaries, and coasts, with their currents and variations in
illumination, temperature, salinity, hydrostatic pressure, and a variety of chemical
components and organisms. Baas Becking’s book, although written in Dutch
(an English translation appeared in Canfield 2015, published by Wiley, and edited
by Don Canfield), became famous by the quote: “. . .alles is overal: maar het milieu

Fig. 1.1 Scan of the title page of Baas Becking’s book “Geobiologie of inleiding tot de
mileukunde” with a photo of the author on board of the RV “Max Weber” of the Zoological Station
Den Helder (predecessor of the Netherlands Institute for Sea Research, NIOZ) while doing research
in the waters around the island of Texel in the Netherlands in 1935. The photo is an original copy
which was made by Dr. H. Oomen, who also was the original owner of this copy of the book which
he obtained in 1935 during the same cruise on board of the RV “Max Weber.” The book is now in
possession of L.J. Stal
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