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Edward Forbes and His Azoic Hypothesis for a Lifeless Deep Ocean

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Edward Forbes and His Azoic Hypothesis for a Lifeless Deep Ocean Review Endeavour Vol.30 No.4 Deserts on the sea floor: Edward Forbes and his azoic hypothesis for a lifeless deep ocean Thomas R. Anderson and Tony Rice George Deacon Division, National Oceanography Centre, Southampton, UK SO14 3ZH While dredging in the Ægean Sea during the mid-19th ship. The captain’s cabin became a laboratory-cum- century, Manxman Edward Forbes noticed that plants museum, where Forbes dissected and drew the animals and animals became progressively more impoverished collected and preserved them in bottles of alcohol. The the greater the depth they were from the surface of the captain and crew were there to help, although it is likely water. By extrapolation Forbes proposed his now infa- that some of the sailors were less than enthusiastic about mous azoic hypothesis, namely that life would be extin- the work involved. However, this work provided evening guished altogether in the murky depths of the deep entertainment for Forbes at least, who frequently turned ocean. The whole idea seemed so entirely logical given his artistic hand to drawing cartoons (Figure 2). the enormous pressure, cold and eternal darkness of this Forbes’ main aim was not, however, to describe the apparently uninhabitable environment. Yet we now animals themselves, but rather to understand their dis- know that the sea floor is teeming with life. Curiously, tribution across different types of habitat. Gathering it took 25 years for the azoic hypothesis to fall from together his results from the Ægean, the influence of depth grace. This was despite the presence of ample contrary was of obvious importance. ‘The sea-bed presents a series evidence, including starfishes, worms and other organ- of zones or regions,’ he wrote ‘each peopled by its peculiar isms that seemingly originated from the deep seabed. inhabitants’ [1]. Seaweeds dominated the uppermost lit- This is a tale of scientists ignoring observations that ran toral zone, below which came the laminarian zone – named counter to their deep-seated, yet entirely erroneous, for its characteristic large brown kelps but occupied also by beliefs. a myriad of fish and invertebrates. Next came the corraline zone, where the abundance of plants declined dramatically The azoic hypothesis in the fading light. Finally, a zone of deep-sea corals that Cruising in the Ægean Sea looking for weird and wonderful existed down to 230 fathoms provided refuge for only a few life-forms on the seabed, Edward Forbes (Figure 1) had an animals, including sponges and molluscs [2]. ‘The number unparalleled opportunity to pursue his great interest – the of species and of individuals diminishes as we descend,’ distribution of animals and plants in the sea. The year was exclaimed Forbes, ‘pointing to a zero in the distribution of 1841 and Forbes, a brilliant naturalist who is considered by animal life as yet unvisited’ [3]. By extrapolation, he many to be the founder of British marine biology, had reasoned that life would be extinguished altogether at a boarded the HMS Beacon under the captaincy of Thomas depth of around 300 fathoms (approximately 550 metres). Graves as part of a surveying mission to the eastern Mediterranean. Dredging in deep water in search of an Evolution of an idea underworld beneath the waves was, however, a major Born on 12 February 1815, it was not long before the undertaking. Although steam engines had been fitted in romantic beauty of his native Isle of Man was to captivate ships from the early years of the 19th century – by 1840 the the young Forbes and inspire him as a naturalist. He spent Royal Navy had no fewer than 50 paddle steamers – most hours arranging, classifying and drawing all manner of naval vessels, like the Beacon, were still driven entirely by objects, including minerals, fossils, shells, dried sea-weeds, the wind and had only man-powered capstans. In these hedge-flowers and dead butterflies. Indeed, by the age of 12 circumstances, it was difficult to maintain a constant he had founded a museum in his own home, and at 16 he course and speed for the duration of a deep dredge haul; left home for London intending to become an artist. The and hand-hauling a heavy, wet rope and a mud-filled Royal Academy refused to admit him. Instead, he followed dredge was time-consuming, backbreaking work. his mother’s wishes and enrolled at the University of Nevertheless, during the 18-month expedition, the crew Edinburgh to study medicine. As things turned out, Forbes of the Beacon made over 100 hauls of the dredge down to a never completed his course because he spent his every depth of 230 fathoms (some 420 metres). In the main, they spare minute pursuing his first love: natural history. Dred- conducted these operations from the Beacon’s tenders, ging the seabed in search of animal life held a particular which were marginally easier to control than the mother captivation for him, with expeditions to the Firth of Forth Corresponding author: Anderson, T.R. ([email protected]). in May 1832, the Irish Sea around the Isle of Man in 1834 Available online 13 November 2006. and the Orkney and Shetland Islands in 1839. www.sciencedirect.com 0160-9327/$ – see front matter ß 2006 Elsevier Ltd. All rights reserved. doi:10.1016/j.endeavour.2006.10.003 132 Review Endeavour Vol.30 No.4 oaks and shrubs at higher altitudes [4]. In similar fashion, Forbes had noted the importance of the influence of eleva- tion above sea level on the distribution of pulmoniferous Mollusca (land snails) in Britain. The number of species declined with altitude until just a single species, Helix alliaria, could tolerate the exposed conditions near moun- tain summits [5]. Aquatic Mollusca were, however, Forbes’ favourite creatures (Figure 3), and it was his dredging studies around the British coasts that inspired the idea of depth-related zones of submarine life [6]. The parallel between terrestrial and marine systems was obvious: extreme environments, be they on the tops of mountains or at the bottom of the sea, would surely lead to extinction of life forms. In fact, the English geologist Henry de la Beche (1796– 1855) had already suggested the idea of a lifeless deep sea as early as 1834. ‘The surface of the deep ocean teems with animal life,’ he wrote, ‘and this is probably not less well proportionably tenanted down to depths where, from the want of necessary conditions, it ceases to exist’ [7]. The work of de la Beche would have been well known to Forbes, from his days as a student in Edinburgh. But it was to be Forbes’ own work in the Ægean, ten years later, that propelled the azoic hypothesis to prominence. His data provided clear evidence for an azoic zone; or so he believed. The scientific community readily embraced Forbes’ azoic hypothesis. It was, after all, so entirely logical. Figure 1. Portrait of Edward Forbes by J.H. Maguire, circa 1850. ßThe Royal Society. How could animals possibly withstand the pressure, cold and eternal darkness of the deep sea floor? The case was stated succinctly by the renowned Swiss-born naturalist By this time, the concept of zonation in land plants had and geologist Louis Agassiz (1807–1873): ‘The depths of the become established. Following his travels to Latin America ocean are quite as impassable for marine species as high between 1799–1804, the great German naturalist and mountains are for terrestrial animals...Not only are no explorer Alexander von Humboldt (1769–1859) remarked materials found there for sustenance, but it is doubtful if that the ‘organic development and abundance of vitality’ of animals could sustain the pressure of so great a column of plant life increased as one went from the poles towards the water’ [8]. Or, as eminent geologist David Page (1814– equator, with different zones of vegetation having their 1879) put it: ‘According to experiment, water at the depth own distinctive character. Furthermore, the same zonation of 1000 feet is compressed 1/340th of its own bulk; and at could be seen on tropical mountains: the palms and bana- this rate of compression we know that at great depths nas found at sea level being replaced by cypresses, pines, animal and vegetable life as known to us cannot possibly exist – the extreme depressions of seas being thus, like the extreme elevations of the land, barren and lifeless soli- tudes’ [9]. Yet we now know that the deep sea floor supports rich animal populations that in terms of species richness might outnumber tropical rainforests and shallow-water coral reef communities many times over [10]. So where did Forbes go wrong? Why did he not find an abundance of animals when dredging the depths of the Ægean Sea? A combination of two factors conspired against him. First, that area is one of the most oligotrophic (nutrient-poor) in the world, resulting in low abundance and few species. Second, the dredges available to Forbes were inefficient, a problem exacerbated by the relatively small size of species that are found there [11]. Danish biologist O.F. Mu¨ ller (1730–1784) might have been the first to use a dredge for scientific purposes. Mu¨ ller’s dredge was a rather coarse netting bag attached to a square iron frame with towing points at each corner Figure 2. Cartoon depicting dredging for marine fauna by Edward Forbes (frontispiece to [2]). Reproduced courtesy of the National Oceanic and (Figure 4a). With such a gaping mouth, it was vulnerable Atmospheric Administration/Department of Commerce. to losing its catch during hauling. By Forbes’ time the www.sciencedirect.com Review Endeavour Vol.30 No.4 133 Figure 3.
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