PDF hosted at the Radboud Repository of the Radboud University Nijmegen The following full text is a publisher's version. For additional information about this publication click this link. http://hdl.handle.net/2066/129525 Please be advised that this information was generated on 2018-07-07 and may be subject to change. NEWS FROM AN INACCESSIBLE WORLD: THE HISTORY AND PRESENT CHALLENGES OF DEEP-SEA BIOLOGY News from an inaccessible world: The history and present challenges of deep-sea biology Erik Dücker, 2014 ISBN: 9789461087201 Printed by: Gildeprint Drukkerijen, Enschede, the Netherlands Cover: Umbellula encrinus, the first deep-sea species discovered. Drawing by Ernst Haeckel (1834-1919) from Kunstformen der Natur (1904). NEWS FROM AN INACCESSIBLE WORLD: THE HISTORY AND PRESENT CHALLENGES OF DEEP-SEA BIOLOGY Proefschrift ter verkrijging van de graad van doctor aan de Radboud Universiteit Nijmegen op gezag van de rector magnificus prof. mr. S.C.J.J. Kortmann, volgens besluit van het college van decanen in het openbaar te verdedigen op dinsdag 23 september 2014 om 10.30 uur precies door Erik Cornelis Petrus Dücker geboren op 18 mei 1979 te Veldhoven Promotoren Prof. dr. C.H. Lüthy Prof. dr. H.A.E. Zwart Manuscriptcommissie Prof. dr. J.M. van Groenendael (voorzitter) Prof. dr. L.T.G. Theunissen (Universiteit Utrecht) Prof. dr. ir. H.J.W. de Baar (Rijksuniversiteit Groningen) CONTENTS GENERAL INTRODUCTION 11 CHAPTER I THE BIRTH OF DEEP-SEA BIOLOGY 17 CHAPTER II EXPLORING BIODIVERSITY 61 CHAPTER III MAN’S DESCEND 109 CHAPTER IV MODERN BASIC RESEARCH 145 CHAPTER V DEEP-SEA BIOLOGY IN CONTEXT 181 GENERAL CONCLUSIONS 219 SUMMARY 223 SAMENVATTING 227 CURRICULUM VITAE 233 DANKWOORD 235 GENERAL INTRODUCTION HEN we are asked to think about natural life on Earth, the most probable images that emerge are those of the vibrantly coloured life of tropical rainforests and coral reefs, the large iconic mammals that inhabit the African plains, or dolphins swimming through crystal clear oceans near some exotic island. Much less probable is that images emerge of a dark abyss inhabited by fang-toothed fishes, bioluminescent jellyfish and flourishing ecosystems around volcanic vents. The deep sea is unfamiliar to us and does not usually spring to mind when we think of nature and wildlife. Still, if we define as the deep sea all those regions of the ocean that are deeper than 1,000 metres, then the deep sea constitutes around 78.5% of the earth’s biosphere 1, and it is teeming with life at all depths. Estimates state that the deep sea may contain the greatest number of animal species, the greatest biomass and the greatest number of organisms in the living world 2. It is thus the dominant part of our biosphere. And yet we have no relationship with it, it falls outside the scope of our daily life, and we have for a long time imagined any human impact as being inconsequential. Life in the deep sea has only been the subject of study by naturalists, and later by professional scientists, since the nineteenth century. Before that time, there had been some cursory investigations of the physical conditions of the deep sea, most notably in the seventeenth century, but no life was expected to exist since the high hydrostatic pressure and low temperature appeared to prohibit life. It was not until the 1840’s that the first systematic investigations of marine species descended beyond the reach of sunlight. Depths of more than 1,000 metres were not reached until in the 1860’s it had become clear that the deep sea was teeming with life. The definitive start of deep-sea biology as a modern science is marked by the expedition of HMS Challenger (1872-1876), which proved that there was no depth-limit to marine life. We must therefore speak of a very recent field of study when compared to the study of life on land and in coastal areas. However, at various levels the deep sea has over the last decades become more visible, first and foremost at a scientific level. As the deep sea lies so far beyond the boundaries of our daily experience, it is a significant undertaking to explore it at all. As a result, the people 11 GENERAL INTRODUCTION primarily engaged in producing knowledge of the deep sea have been naturalists first, and scientists later. Even today deep-sea exploration is the exclusive domain of professionals, who are uncovering new aspects with almost every dive, every net and every sample taken. In addition, the deep sea has turned out to be rich in traditional resources such as oil, gas and minerals, as well as in biological resources such as genes and proteins. Because the exploitation of such resources has become more viable in recent decades, the deep sea is steadily gaining in economic importance. As a result, human civilization has an increasing impact on these remote ecosystems. Direct human impact does not only occur because of the exploitation of deep-sea biological and mineral resources, but also through the disposal of waste (3-5 and 6 p. 137-224). Indirect impact through biodiversity loss, ocean acidification and climate change also occur, and they appear to be more significant than has previously been assumed (7-9 and 6 p. 176-196). These developments raise questions about the conservation of the deep-sea environment. Deep-sea conservation, however, requires, due to its particular nature 10, more than merely an extension of existing conservation efforts from coastal areas and fisheries into the deep sea. Knowledge of deep-sea life, however, continues to be limited. But if the estimates are correct and the major occurrence of animal life is predominantly found in the deep sea, then the question arises as to why biology, which is after all the study of life, pays so little attention to this vast environment. The primary reason why knowledge of the deep sea progresses at such a slow pace is that any investigation of it requires a huge investment of time, money and resources. These are not generally available to researcher teams. Worse yet, only a few countries have the capability to reach great depths. This capacity requires technology that is expensive to develop and equally expensive to use. Operating costs are typically US $30,000 per day and cruises generally last a few weeks, so that the costs per cruise can quite easily reach US $1 million 4. The remoteness of the deep sea and the technological difficulty of investigating has as a further consequence that the deep-sea biologist is almost the exclusive producer of knowledge of the deep sea. On land there is a multitude of sources of knowledge, ranging from historical descriptions of species distribution over hobby ecologists who enjoy spending their weekends in the woods to people who follow a centuries-old tradition of living in symbiosis with nature, for instance, such as found in the Japanese Satoyama landscapes 11. No such non- scientific sources of knowledge and experience are available for the deep sea, with the possible exception of a few attentive deep-sea fishermen. Due to these circumstances, deep-sea biologists have always had to depend heavily on support from governments and commercial partners. Although this situation renders this discipline quite unique in the life sciences, the consequences of this strong dependence have to date never been examined. Unless I am mistaken, this dissertation is the first monograph dedicated to an examination of the choices and dilemmas of deep-sea biology from its beginnings in the nineteenth century to today. In order to write such a study, it was necessary to combine historical research with philosophical reflection, creating what we might term a ‘philosophical history of deep-sea 12 GENERAL INTRODUCTION biology’. As the history of oceanography, the field to which deep-sea biology is intimately related, is well established, this dissertation draws on the work of such prominent historians of oceanography as Philip Rehbock, Eric L. Mills, Helen Rozwadowski and Naomi Oreskes. It re- evaluates and adjusts their findings in the context of deep-sea biology as being discipline within general biology, rather than oceanography, thereby adding a fresh perspective. Due to my personal background as a biologist, I have been able to incorporate the scientific literature beginning with prominent Victorian naturalists such as Edward Forbes (1815-1854) and ending with the latest scientific publications in the field of deep-sea biology. Along the way I managed to uncover hitherto untreated material, including forgotten nineteenth-century French deep-sea physiological research and recent discoveries of extraordinary deep-sea creatures that are not always acknowledged as important. The historical narrative developed in this dissertation shows how the role of the deep-sea scientist has changed according to the external factors to which it was exposed in each period of its development. Initially, the naturalist exploring the deep sea was a ‘gentleman naturalist’ exploring at his own expense and leisure; the required independent wealth obviously limited access to this science. This role changed as deep-sea biology developed into a professional science supported by government grants. Subsequent transformations of the deep-sea biologists’ role turned them first into fisheries experts and later into cold war allies, and nowadays they must be everything at once: scientists, explorers, advisors and environmental advocates. These new roles raise questions that do not limit themselves to empirical matters, to which the deep-sea biologist nonetheless continues to be sensitive. These new questions will be discussed through the use of case studies of recent developments in deep-sea manganese nodule mining as well as bioprospecting, that is, the search for biological compounds that are of use for commercial purposes.